You are on page 1of 686

Aquatic Biodiversity in Latin America ABLA

Biodiversidad Acuática en América Latina Vol. 3

FOR MANY YEARS,

Neotropical Simuliidae (Diptera: Insecta)


STUDENTS FROM NUMEROUS COUNTRIES HAVE COMPLAINED
ABOUT THE LACK OF “TOOLS” TO IDENTIFY AQUATIC INVERTEBRATES FROM
LATIN AMERICA. KEYS FOUND IN ACCEPTED TEXTBOOKS ARE MOSTLY LIMITED,
SUPERFICIAL OR TOO POPULAR-SCIENCE. ON THE OTHER HAND, MORE ADVAN-
CED KEYS IN ACADEMIC BOOKS ON AQUATIC INVERTEBRATES FAIL TO COVER
NEOTROPICAL REPRESENTATIVES IN DUE DETAIL. THE FEW THAT DO, HOWEVER,
ALSO SHOW TOO MANY DEFECTS IN SCOPE AND PRESENTATION TO BE CONSID-

Sixto Coscarón & Cecilia L. Coscarón Arias


ERED UP-TO-DATE.
AN INTERNATIONAL TEAM OF EDITORS COMBINED THEIR EFFORTS WITH
PENSOFT PUBLISHERS TO LAUNCH A NEW MAJOR SERIES ON THE AQUATIC BIODI-
VERSITY IN LATIN AMERICA. ABOUT 15 MONOGRAPHS, WRITTEN BY SCIENTISTS
FROM VARIOUS COUNTRIES, WILL OFFER A NEW, UNRIVALLED VIEW OF THE MAR-
VELLOUS AQUATIC WORLD OF SOUTH AMERICA!
THE SERIES IS ADDRESSED TO ZOOLOGISTS, ECOLOGISTS, HYDROBIOLOGISTS,
BIOGEOGRAPHERS, CONSERVATIONISTS AND STUDENTS INTERESTED IN AQUATIC
BIODIVERSITY. THE SERIES WILL BE AN IMMINENT TOOL FOR ANY BIOLOGICAL
LIBRARY.

Volume 3
This book is unique in coverage, summarizing all available information con-
cerning the American simuliid fauna south of the United States. It also includes
morphological diagnoses of females, males, pupae and larvae, as well as keys (in
English and Spanish), illustrations of characters, mapped distributions, and bio-
nomics. This region appears to support 359 species grouped in two tribes,
12 genera and 18 subgenera. The description of each taxon is provided with a
list of the available literature, as well as all other relevant information. The book
is addressed to taxonomists, limnologists, ecologists, veterinarians and biologists
in general.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 1

Aquatic Biodiversity in Latin America

Biodiversidad Acuática en América Latina

Volume 3
Neotropical Simuliidae (Diptera: Insecta)

Volumen 3
Simuliidae Neotropicales (Diptera: Insecta)
2 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

BIODIVERSIDAD ACUÁTICA EN AMÉRICA LATINA

Volumen 3

SIMULIIDAE
NEOTROPICALES
(DIPTERA: INSECTA)
por
Sixto Coscarón & Cecilia L. Coscarón Arias

Lector Científico a los Editores:


Peter H. Adler

Editores de la Serie:
Joachim Adis, Jorge R. Arias,
Guillermo Rueda-Delgado & Karl Matthias Wantzen

Sofia–Moscow
2007
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 3

AQUATIC BIODIVERSITY IN LATIN AMERICA

Volume 3

NEOTROPICAL
SIMULIIDAE
(DIPTERA: INSECTA)
by
Sixto Coscarón & Cecilia L. Coscarón Arias

Scientific Reader to the Editors:


Peter H. Adler

Series Editors:
Joachim Adis, Jorge R. Arias,
Guillermo Rueda-Delgado & Karl Matthias Wantzen

Sofia–Moscow
2007
4 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

AQUATIC BIODIVERSITY IN LATIN AMERICA


BIODIVERSIDAD ACUÁTICA EN AMÉRICA LATINA

Series Editors/Editores de la Serie:


Joachim Adis, Jorge R. Arias, Guillermo Rueda-Delgado & Karl Matthias Wantzen

VOLUME 3. VOLUMEN 3.
NEOTROPICAL SIMULIIDAE (DIPTERA: INSECTA) SIMULIIDAE NEOTROPICALES (DIPTERA: INSECTA)
by por
Sixto Coscarón & Cecilia L. Coscarón Arias Sixto Coscarón & Cecilia L. Coscarón Arias
Scientific Reader to the Editors: Lector Científico a los Editores:
Peter H. Adler Peter H. Adler

Front cover:
Male of the newly described Simulium nunesdemelloi HAMADA, PEPINELLI & HERNÁNDEZ, 2006
(Simuliidae) and Amazon River near Manaus, Brazil
(photos: M. Cutrim, W.J. Junk; design: Zheko Aleksiev & Elke Bustorf)

Aquatic Biodiversity in Latin America Vol. 3


ISSN 1312-7276
First published 2007
ISBN: 978-954-642-293-4

The book should be cited as follows:


Sixto Coscarón & Cecilia L. Coscarón Arias. 2007. Neotropical Simuliidae (Diptera: Insecta).
In: Adis, J., Arias, J.R., Rueda-Delgado, G. & K.M. Wantzen (Eds.):
Aquatic Biodiversity in Latin America (ABLA). Vol. 3. Pensoft, Sofia-Moscow, 685 pp.

© PENSOFT Publishers
All rights reserved. No part of this publication may be reproduced, stored in a retrieval system or
transmitted in any form by any means, electronic, mechanical, photocopying, recording or
otherwise, without the prior written permission of the copyright owner.

Pensoft Publishers
Geo Milev Str. 13a, Sofia 1111, Bulgaria
pensoft@mbox.infotel.bg
www.pensoft.net

Printed in Bulgaria, March 2007


ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 5

AQUATIC BIODIVERSITY IN LATIN AMERICA

BIODIVERSIDAD ACUÁTICA EN AMÉRICA LATINA

Series Editors: Joachim Adis, Jorge R. Arias,


Guillermo Rueda-Delgado & Karl Matthias Wantzen

Biodiversity is a key word in science and global management schemes; however very few
people are able to identify the species and their ecology that make up “biodiversity”. For
many years, researchers and students from numerous countries complain about the lack of
“tools” to identify aquatic invertebrates from Latin America. Keys found in accepted
entomological textbooks are mostly highly limited, superficial and rarely cover Neotropical
biota in sufficient detail. On the other hand, specialized information on taxonomy or ecology
is scattered throughout the literature in many single publications.
An international team of editors have combined their efforts with Pensoft Publishers
to launch a new major series on the Aquatic Biodiversity of Latin America (ABLA). Their
goal was to find experts who combine the current state of knowledge in taxonomy and
ecology, in order to produce a concise and affordable handbook for each group. About 15
separate monographs, written by reference scientists from various countries will offer a
new, unrivalled view on the aquatic fauna of South America. Information on the ecology
and status of the taxa (written in English) is combined with illustrated identification keys to
families and genera, in both English and Spanish. The series is aimed at zoologists, ecologists,
hydrobiologists, biogeographers, conservationists and students interested in aquatic
biodiversity. The series will be an essential tool for any biological library.

Volume 1: Amazon Fish Parasites (Second edition) by Vernon E. Thatcher.


Pensoft Publishers, Sofia-Moscow, ISBN-10: 954-642-258-4, ISBN-13: 978-954-642-
258-3, 165x240 mm, 508 pp., including 194 plates of figures in line drawings and
photos (15 plates in color); publication date: March 2006.

Volume 2: Ephemeroptera of South America by Eduardo Domínguez, Carlos Molineri,


Manuel L. Pescador, Michael D. Hubbard & Carolina Nieto.
Pensoft Publishers, Sofia-Moscow, ISBN-10: 954-642-259-2, ISBN-13: 978-954-642-
259-0, 165x240 mm, 646 pp., including 234 plates of figures in line drawings and
photos (16 plates in color); publication date: May 2006.
6 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Volume 3: Neotropical Simuliidae (Diptera: Insecta) by Sixto Coscarón & Cecilia L.


Coscarón Arias.
Pensoft Publishers, Sofia-Moscow, ISBN-13: 978-954-642-293-4, 165x240 mm, 685
pp., including 135 plates of figures in line drawings and 19 maps of distribution;
publication date: March 2007.

Simuliid larvae and pupae inhabit freshwater streams, from sea level to about 5000 m altitude
in the Neotropics. The larvae are filter feeders and, along with pupae, live attached to different
underwater strata for support. These insects play an important role in the bioenergetic cycle
of continental waters. Imagoes are strong flyers, and females are well known for their biting
habits as well public health importance, being vectors of several pathogens.
This book compiles the current known information on the American simuliid fauna
south of the United States. It also includes a morphological diagnosis of females, males,
pupae and larvae, along with keys, illustrations of characters, distributions with maps, and
bionomics for the various taxa. In this region, there are 359 known species, grouped in two
tribes, 12 genera and 18 subgenera. The description of each taxon includes a list of the
available literature as well as the most relevant information. The book is addressed to
systematists, limnologists, ecologists, veterinarians and biologists in general.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 7

ADDRESSES

Authors:
DR. SIXTO COSCARÓN M.Sc. GUILLERMO RUEDA-DELGADO
Museo de Ciencias Naturales de La Plata Grupo de Investigación en Cuencas y
Paseo del Bosque S/N Humedales Tropicales GICHT
1900 La Plata, Argentina UNIMAGADALENA
e-mail: sixtocos@netverk.com.ar Universidad de Bogotá Jorge Tadeo Lozano
Laboratorio de Limnología
DR. CECILIA L. COSCARÓN ARIAS Carrera 4 No. 22-61
LIBIQUIMA (Laborartorio de Bogotá, D.C.
Investigaciones Bioquímicas y de Medio Colombia
Ambiente) e-mail: guillermo.ruedadelgado@utadeo.edu.co
Universidad Nacional del Comahue
Buenos Aires 1400, DR. KARL MATTHIAS WANTZEN
8300 Neuquen, Argentina Institute of Limnology
e-mail: ariascoscaron@hotmail.com University of Konstanz
Postfach M 659
Series Editors: 78457 Konstanz, Germany
PROF. DR. JOACHIM ADIS e-mail: matthias.wantzen@uni-konstanz.de
Max-Planck-Institute for Limnology
Tropical Ecology Working Group Scientific Reader to the Editors:
Postfach 165 PROF. DR. PETER H. ADLER
24302 Plön, Germany Division of Entomology
e-mail: adis@mpil-ploen.mpg.de Clemson University
P.O. Box 340315, 114 Long Hall
DR. JORGE R. ARIAS Clemson, SC 29634-0315, USA
5870 Colfax Avenue e-mail: padler@clemson.edu
Alexandria, Virginia 22311, USA
e-mail: jaria2@fairfaxcounty.gov
8 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

FOREWORD

Neotropical Simuliidae (Diptera: Insecta)


by Sixto Coscarón & Cecilia L. Coscarón Arias

Research on the Simuliidae in the Neotropical area has been ongoing for over a century. It
largely consisted of sporadic accounts of species by various authors with little coordination
of effort until the 1930s. Enderlein then produced many names of new species and genera
and a classification, which through various convoluted pathways helped to form the
taxonomic framework in use today. The driving force behind the subsequent increase in
taxonomic studies has been largely medical. Simuliid flies in Latin America are responsible
for two problems. Firstly, the presence of enormous biting populations in certain areas that
can preclude development of land for agriculture and deter tourism. Secondly, the
incrimination of several species in transmitting two filarial worms to man, Mansonella ozzardi
and Onchocerca volvulus, responsible for the diseases mansonelliasis and onchocerciasis. The
major impetus to these studies was the need to incriminate the vectors of human
onchocerciasis in the 1940s in Guatemala and Mexico. This led to the pioneering work on
vector incrimination and simuliid taxonomy by Luis Vargas and his colleague Alfonso Díaz
Nájera in Mexico. These authors provided a good taxonomic foundation for the Central
American fauna. In the 1950s Rachou’s work in the Amazon basin of Brazil on
mansonelliasis, a disease already suspected for some time to be associated with Simuliidae
in Argentina, inspired continuing studies on Simuliidae of the region. At about this time
Pedro Wygodzinsky started work on the family in Argentina, soon to be joined by Sixto
Coscarón. Their classic work on the family, with methodical collection of species and
further development of the taxonomic framework culminated in Coscarón’s overview of
the Neotropical Simuliidae in 1987. This publication, in conjunction with Crosskey’s world
inventory of Simuliidae produced at about the same time, has had monumental significance
in Latin America. For the first time simuliid researchers in Latin America were able to
access information on valid names for simuliid species, their classification and references
that supply data on simuliid biology and medical importance.
This latest Neotropical overview of Coscarón & Coscarón-Arias brings us up to date
with the considerable advances emanating from the discovery of human onchocerciasis in
Amazonia in the 1970s. Its publication in English now provides ready access for simuliid
specialists at an international level. This new book has brought together the relevant
voluminous literature in Portuguese, Spanish, French and English on simuliids from a vast
area of the planet. It provides a systematic analysis that largely corresponds to the scheme
in Crosskey’s world annotated catalogue. Main characters of species are covered, with ample
references to descriptions by these and other authors and very comprehensive illustrations
and species distribution maps. Methods used in collecting and preparing specimens for
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 9

subsequent examination and the terminology used in their description are described.
Synopses of the biology, medical importance and control of simuliids in Latin America are
also provided. The authors regard this work as a “snapshot view” of the complicated
Neotropical simuliid fauna for which much taxonomic study is still necessary. It provides a
sound basis for future in depth morphological studies of certain species, where warranted,
on economic or medical grounds, and for studies integrating morphological, cytological
and molecular methods for phylogenetic purposes.

TONY SHELLEY
The Natural History Museum
London, U.K.
10 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

PREFACE

Simuliidae are present in most freshwater streams where the larval stages develop and the
adults are well known to the nearby human population by the nuisance of the female’s bite.
The immature stages are present in currents of Neotropical creeks, streams and rivers, in
slow to torrential waters. During the larval and pupal stages, they live attached to various
substrates in waters of variable temperatures, salinity and turbidity, showing great adaptability
to different habitats. Larvae feed on algae, bacteria, organic particles of dead organisms
and inorganic material, showing a great capacity to metabolize different materials. This
aspect of their life history, added to the fact that they serve as food for fishes, birds and
other arthropods, gives an idea of the important role they play in the energy cycle of
continental freshwater ecosystems.
Adults are strong fliers with high dispersal qualities, and they have variable feeding
habits, ranging from plant nectar to vertebrate blood. This great adaptability is represented
by about 350 known species in the Neotropical region. Certainly this number will increase
since many areas are unstudied or have been scarcely studied.
This volume is a compilation of the known Neotropical black flies found from south of
the United States to the south of Tierra del Fuego, Argentina. The biogeography does not
coincide with political limits, but in this case it approximates the extension of this region.
The aim of this volume is to make a compilation of the recognizable species, arranged in
supraspecific taxa and arranged in accordance with the relationships obtained by cladistic studies.
Here we consider l2 genera and 284 species with keys, diagnoses, illustrations of morphology
and distribution maps. We do not include 65 species of Gigantodax and Pedrowygomyia, because
they were treated in a previous review (WYGODZINSKY & COSCARÓN, l989).
An introductory section on general morphology is included to understand the
anatomical names used in the descriptions. Also, we add information on biotic conditions
related to the species, as well as their public health importance and control.
The present volume was planned to help less experienced people that desire to
investigate this interesting family of Diptera. In order to address the Latin American
audience, keys and illustration legends are also offered in Spanish.
Carmencita (Carmen ENRIQUETA ANNUASI) is heartly thanked for her great patience
and continuous support.
We are very grateful to Nélida CALIGARIS for her help in typing and arranging most of the
illustrations, which are based on her illustrations made during many years of our publications.
Also, we are indebted to ALICIA GARCIA for her help in typing part of this manuscript.
We are indebted to all the Neotropical simuliidologists that have provided information
from their publications used here or have provided study material, namely Victor PY-DANIEL,
Anthony SHELLLEY, Sergio IBÁÑEZ-BERNAL, Marilza MAIA-HERZOG, Milton STRIEDER, Neusa
HAMADA, Gilson MOREIRA, Paulino LUNA-DIAS, Peter ADLER, Leonardo Gil AZEVEDO, Marcia
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 11

COSTA, Jaime RAMÍREZ PÉREZ, Paulina MUÑOZ DE HOYOS, and specially to Rafael MIRANDA
ESQUIVEL for the cladistic studies and his preoccupation to solve the phylogeny of the
Simuliidae. We are also grateful to several institutions that have provided study material,
including the American Museum of Natural History (New York), the United States National
Museum, (Smithsonian Institution, Washington, D.C.), The Natural History Museum (
London), the Museum für Naturkunde (Berlin), INDRE, (Mexico) and the Museu de
Zoologia (São Paulo). In the first institution we were also allowed to use information from
articles published in the Bulletin and Novitates. Our special acknowledgment goes to Zootaxa
for permission to use information published in volume 396 (2004), and to the Memoirs on
Entomology International for the information extracted from volume 14 (1999).
The authors are much indebted to CONICET for financial support.
We thank the editors and the scientific reader of the book for their valuable help
received while preparing the manuscript for publication.
We extend special acknowledgment to our late master, Pedro WYGODZISNKY, who gave
all his knowledge and enthusiasm to improve the study of black flies with modern methods
in morphology and phylogenetic analysis, and especially for encouraging us to explore the
interesting Simuliidae world.

THE AUTHORS
12 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Table of Contents

1I NTRODUCTION
Bionomics
15
15
Behavior 16
Public health and socioeconomic importance 17
Control 18

2M ORPHOLOGY AND CHARACTERS OF TAXONOMIC VALUE 22

3C OLLECTION AND PREPARATION OF MATERIAL 34

4S TUDY OF MATERIAL 36

5 S Keys to the genera of Neotropical Simuliidae


YSTEMATICS 38
39
Claves de géneros de simúlidos de la región Neotropical 45
Mayacnephia WYGODZINSKY & COSCARÓN 52
Tlalocomyia WYGODZINSKY & DÍAZ NÁJERA 63
Araucnephia WYGODZINSKY & COSCARÓN 66
Araucnephiodes WYGODZINSKY & COSCARÓN 76
Cnesia ENDERLEIN 81
Gigantodax ENDERLEIN 90
Pedrowygomyia COSCARÓN & MIRANDA ESQUIVEL 91
Cnesiamima WYGODZINSKY & COSCARÓN 93
Paraustrosimulium WYGODZINSKY & COSCARÓN 99
Lutzsimulium D’ANDRETTA & D’ANDRETTA 102
Kempfsimulium PY-DANIEL & NUNES DE MELLO 110
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 13

Simulium LATREILLE 114


Keys to Simulium subgenera 115
Claves de subgéneros de Simulium 125
Simulium (Eusimulium) ROUBAUD 137
Simulium (Nevermannia) ENDERLEIN 137
Simulium (Byssodon) ENDERLEIN 138
Simulium (Psilozia) ENDERLEIN 138
Simulium (Pternaspatha) ENDERLEIN 139
NIGRISTRIGATUM species group 159
NEMORALE species group 188
Simulium (Notolepria) ENDERLEIN 224
Simulium (Chirostilbia) ENDERLEIN 238
PERTINAX species group 247
SUBPALLIDUM species group 259
Simulium (Inaequalium) COSCARÓN & WYGODZINSKY 266
INAEQUALE species group 273
BOTULIBRANCHIUM species group 289
Simulium (Psaroniocompsa) ENDERLEIN 295
INCRUSTATUM species group 302
SIOLII species group 319
Simulium (Cerqueirellum) PY-DANIEL 325
Simulium (Coscaroniellum) PY-DANIEL 349
QUADRIFIDUM species group 354
QUADRIVITTATUM species group 364
Simulium (Ectemnaspis) ENDERLEIN 366
BICOLORATUM species group 385
ROMANAI species group 403
PERFLAVUM species group 409
DINELLII species group 414
Simulium (Psilopelmia) ENDERLEIN 419
ESCOMELI species group 437
BLANCASI species group 463
Simulium (Aspathia) ENDERLEIN 471
14 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Simulium (Hemicnetha) ENDERLEIN 517


PAYNEI species group 531
BRACHYCLADUM species group 540
MEXICANUM species group 547
OVIEDOI species group 556
Simulium (Hearlea) VARGAS, MARTÍNEZ PALACIOS &
DÍAZ NÁJERA 561
JUAREZI species group 571
CAROLINAE species group 586
Simulium (Trichodagmia) ENDERLEIN 597
Simulium (Thyrsopelma) ENDERLEIN 612

6M APS OF DISTRIBUTION 627

7R EFERENCES 648

8S UBJECT INDEX 671

9A DDENDUM 682
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 15

1
INTRODUCTION
Bionomics

Black flies can be found in highly variable environments from very cold water (about
0 °C) to warm water exceeding 25 °C, from sea level to an altitude of 5000 m, from
freshwater to water with a certain degree of salinity, from clear to turbid water, and from
oxygen-saturated to oxygen-poor water. The only distinctive condition that is necessary
for their development is continuously running water. With this extensive range of
possibilities, they can be found practically everywhere flowing water exists, from tiny
streams to large rivers (COSCARÓN ARIAS, 2001).
Larval black flies inhabit aquatic environments, constituting an important link in the
trophic chain of lotic biotopes. Generally, they choose places with continuous and fast-
flowing water. They are found near the surface where higher oxygen tension exists and
leaves, branches, stones or plastic substrates, to which they attach, are present and are free
of algae and fungus.
Physical and chemical factors regulate the presence, richness and abundance in the
micro- and macrodistribution of the species (COSCARÓN ARIAS, 2001). The more important
factors are the following:
• Turbidity: Larvae prefer clear water but they sometimes tolerate water with some degree
of turbidity during certain periods or permanently. Also, they require water that receives
direct light. Watercourses that run long distances through poorly lighted areas usually
have no larvae or pupae.
• Salinity: They live in oligohaline waters, which have a solid residue lower than 5 g/l.
Most of them live in waters with less than 1 g/l, but others, such as Simulium wolffhuegelii,
can breed with a concentration close to 4 g/l.
• pH: The habitat with a value near neutral is typical; however, some species can live in
water with pH values above or below neutral, ranging from 6.56 to 8.90.
• Oxygen: The high content of dissolved oxygen in micro-bubbles that elevate the O2
tension to saturation in torrential waters is a factor that benefits most black flies. Most
of the dissolved oxygen comes from the photosynthetic process that results in sunny
places.
• Temperature: The larvae are eurythermic and they can live in water with temperatures
close to 0 °C, such as the species found in the Puna (Argentina), high mountain ranges
or in Tierra del Fuego (Argentina) where they are found in streams covered by snow.
They can also be found in watercourses with temperatures of 25 °C such as those in
16 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

tropical areas. The great majority of species prefer water temperatures ranging from
7 °C to 15 °C that are more frequently found in springs.
• Altitude: They can be found from sea level up to 4700 m. The altitude is correlated
with temperature. For example, in Argentina, the highest percentage of species collected
(41%) is found between 500 and 1000 m. This is followed by those species that are
found between sea level and 500 m.
• Support strata: The larvae use a large variety of substrates to adhere to, but they
always require a solid support. They avoid places with many unicellular algae. The
most frequent substrates used are stones, leaves, sticks, roots, branches or tree trunks;
even though they rarely adhere to dead vegetation, 53% of the Argentinean species
adhere to vegetation. Some species have a predilection for a particular type of substrate
but there are others that are indifferent and use any kind of support, including pieces
of iron, plastic, and wood.
• Depth: The majority of larvae and pupae live on trailing vegetation in water that is not
deeper than 50 cm. Species are only observed in deeper water when there is no turbidity.
• Current speed: Simuliid larvae and pupae are positively reophilic. Since they filter the
water to obtain food and dissolved oxygen, the greater the volume of water they filter,
the more beneficial it is to them. Current speed is an important factor to define their
habitat because they need to maintain attachment to the substrate, keep their cephalic
fans erect and allow their muscles to contract when folding the fans to take the food
to the mouth. The species that live in the plains will survive in water with minimum
speeds of around 2 cm/s; however, there are species that can live in water with a
lesser speed. Most of the species prefer water speeds between 10 and 50 cm/s, although
there are some species that only live in torrential courses of 100 cm/s.

More information about the environmental factors in relation to the different species
of simuliids of the Neotropics can be found in COSCARÓN et al. (1996b; 2000), COSCARÓN
& COSCARÓN ARIAS (2000 b), GRILLET & BARRERA (1997), HAMADA & MCCREADI (1999),
MOREIRA et al. (1994).

Behavior

Simuliids are primarily diurnal insects. Both sexes feed on nectar from plants or flowers;
only the females of some species are haematophagous. Although some species can be
autogenous (i.e., they don’t suck blood), others require blood of vertebrates to lay viable
eggs. Depending on their haematophagic preferences, they can be recognized as zoophilic
(feeding on animals), ornithophilic (on birds), mammalophilic (on mammals) and
anthropophilic (on humans). Immature states (eggs, larvae and pupae) are aquatic.
Copulation occurs at the site where the adults emerge from their pupal casings. The
females produce 200 to 500 eggs in a single gonotrophic cycle and some are able to
distribute their eggs freely on the surface of the water (during flight) or in mass
ovipositing on leaves or branches near the water. Some females will crawl under the water
to deposit their eggs (PETERSON, 1981). The incubation period of the eggs varies from 4 to
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 17

30 days, depending on the species and the water temperature, being much longer in those
species in which the eggs undergo diapause (COLBO, pers. commun.). The eggs develop
under humid conditions or below water. Hatching occurs a few days after oviposition and
the larvae subsequently adhere to a substrate by the disk at the posterior end of their body
(abdominal proleg). Sometimes they will also use the thoracic proleg, as well as a fine silk
thread that is secreted through the mouth from their salivary glands. The larvae are non-
selective filter feeders, trapping small organisms (zooplankton and phytoplankton) and
organic particles of other insects circulating in the water column. The larvae occasionally
drift downstream, being carried by the water currents.
The final larval instars build a cocoon of silk secreted from the salivary glands. This
serves as a refuge for pupation and transformation to the adult stage. The pupae have a
slipper or shoe shape, with a series of filaments, which are the branches of gills (breathing
organs), anteriorly. Simuliids remain in the pupal stage from 7 to 20 days, depending on
the water temperature. During this period, they undergo a complicated metamorphosis to
the adult stage, from which the adult fly emerges wrapped in a bubble of air that pops to
the surface, where the fly becomes airborne to restart the cycle. At the beginning of the
adult stage, the female flies look for their host. With few exceptions, the females show no
host specificity. After taking a bloodmeal, the females look for a place to rest and digest
the meal for egg maturation. Once the eggs have matured, the females select the
appropriate site for oviposition. Simuliids generally feed during daylight hours, although
some species prefer less sun and will feed either at dawn or during twilight hours, as in
other haematophagous Diptera. Some species exhibit bimodal patterns of haematopha-
gous activity while other species remain active throughout the day. It is believed that solar
light stimulates haematophagous activity, but this is probably not the only factor, as
feeding activity has been observed in dark places or on cloudy days. On the other hand,
wind decreases feeding behavior, reducing the incidence of bites as wind velocity
increases. The adults generally remain close to the streams in which they breed, generally
in nearby humid microclimates, such as trees. The largest densities are observed within
100 m of the breeding sites, in shaded areas.
Simuliids can disperse by intrinsic and extrinsic flight mechanisms to distances as
great as 15 km or even as far as 40 km from their breeding sites, with the aid of wind
(COSCARÓN, 1981b) and in Africa up to 400 km (CROSSKEY, 1990).
Simuliid longevity is not well understood under field conditions but it is believed to
vary around two months in temperate areas.
The distribution of black flies is a consequence of their adaptive capacity to the
environment and ancestral origin; a significant correlation is evident between distribution
and endemism in the Neotropical realm (COSCARÓN & COSCARÓN ARIAS, 1995), as also
evidenced in intercontinental relationships (MIRANDA ESQUIVEL & COSCARÓN, 2004).

Public health and socioeconomic importance

Black flies are not only annoying bloodsucking flies, but they are also vectors of important
infectious diseases. Knowledge of this group will allow the reader to appreciate their
18 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

importance as entomo-epidemiological agents. Black flies are known vectors of patho-


gens, such as filarial worms, viruses (myxomatosis and encephalitis) and protozoans (e.g.,
Leucocytozoon in birds). Of the parasites transmitted by this family, the most dangerous one
to humans is Onchocerca volvulus, the etiological agent of onchocerciasis, where in extreme
case it can produce blindness. Other species of Onchocerca parasitize cattle and wild
animals (e.g., primates, birds) (Onchocerca lienalis, O. reticulata, O. gutturosa and O. cervicalis).
Another filarial worm that can be transmitted to humans by simuliids is Mansonella ozzardi,
although its resultant pathogenic action is slight.
Leucocytozoon smithi (a protozoan parasite) has caused serious problems and numerous
deaths on turkey farms, and L. simondi does the same in ducks in the USA. Leucocytozoon
spp. also parasitize wild birds, particularly raptors, but they also have been found in doves
(L. bonasae), chickens (L. andrewai), and even in penguins in New Zealand (L. tawaki). To
date, no studies dealing with this parasite exist in Latin America. Simuliids are also vectors
of arboviuses. They have been identified as being responsible for the transmission of
Venezuelan equine encephalitis (VEE) and eastern equine encephalitis (EEE). In
Australia, they are incriminated as vectors of the myxoma virus among others.
As well as the annoyance that they produce, the saliva that is excreted during biting
causes severe allergic reactions and may be responsible for secondary infections in people
when they scratch. Black flies also mechanically transmit tularemia and other diseases
(causing local reactions with itch and edema). Simuliids also cause economic losses at a
regional level by affecting agriculture. The flies inhibit people from working normally
outdoors, thereby reducing agricultural production. They also impact the cattle industry
because the irritation caused by the simuliids to the cattle inhibit them from feeding
properly, reducing the production of meat and milk. Losses to tourism must also be added
to the list of socioeconomic problems; the annoyance caused by these insects can be so
intense as to suspend recreational activities.

Control

There are different methods of black fly control: chemical, mechanical and biological; a
general review of this topic for the Neotropical realm can be found in CAMPOS &
ANDRADE (2001).

Types of Control
Chemicals are the best-known methods for insect control; synthetic insecticides have
been especially effective. However, their use, particularly the wide-spectrum, residual
insecticides, have altered ecosystems.
Pesticides have been used against both the adult flies that attack us and the larvae,
interrupting their life cycle; yet the methods to control the adult stage are not
recommended because the insecticides that have to be applied to large areas result in
adverse effects on non-target organisms, which include beneficial insects such as
pollinators, predators, fish, birds, and amphibians. Chemicals used in the control of adult
flies have included products such as DDT, methoxyclor, malathion, naled (commercially
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 19

sold as Dibron) (ROMAÑA & AVALOS, 1948; ANDRADE, 1989) or diverse pyrethroids (SMITH,
1973; ELLIOT & POTTER, 1978); extreme caution should be taken so that these products do
not contaminate rivers, streams or ponds.
Adulticides should be used only in the case of an epidemic, to stop nuisance fly
populations in restricted areas and only for short periods, as in the case of social events.
For these reasons, the larvae are targeted in control strategies because they are limited to
more defined areas, i.e. watercourses.
There are extensive lists of products that have been used efficiently in the control of
black flies, but for diverse reasons, such as high residual action, adverse effects on non-target
organisms and resistance, many products have been replaced by newer less persistent
insecticides. One of the latest insecticides being used has been temephos, mainly due to its
low toxicity to vertebrates. However, resistance and toxicity to other insects has led to its
replacement, especially by biological larvicides that have a more selective action. As yet there
is no commercial chemical product available that is specific against simuliids that can be
applied on a large scale that will not be harmful to the other organisms.
The most advisable technique for black fly control is the use of biological control
agents, including the use of live organisms or products that are derived from the black
flies themselves and that are aimed at the reduction or elimination of the damages caused
by species that are harmful to humans, domestic animals or agriculture.
There are several biological control methods, but those with better results are the
ones derived from bacteria, fungi and viruses. Natural enemies of simuliids in watercours-
es include organisms such as predators (e.g., fish, coleopteran larvae), parasites (Mermith-
idae), protozoa (microsporidia), fungi (especially Coelomycidium), viruses and bacteria. In
regions where natural enemies exist and do not reach sufficient levels to be effective as a
control measures, it is necessary to establish artificial rearing facilities to produce
significant densities of the pathogens and parasites. This would not be easy to implement
because they usually have complex cycles with diverse intermediary hosts and would be
difficult to produce in sufficient quantities to achieve a viable level of control.
Furthermore, there is still insufficient knowledge about the biological cycle of microspo-
ridia and fungi to say whether they could offer better options.
The search for a practical larvicide has led to the entomopathogen Bacillus thuringiensis
var. israelensis (Bti), which was found to be the ideal solution. Bacillus thuringiensis is a
bacterium that forms spores and produces protoxin crystals (delta-endotoxin) activated
inside the alkaline intestine of the insects, and which become toxic, causing the
degeneration of the epithelial lining of the intestine, paralysis and rapid death.
Another control method is the mechanical cleaning and elimination of substrates,
especially vegetation, to which the simuliid larvae and pupae can adhere. These methods
could be used in conjunction with the application of the B.t.i. as a form of integrated
control. Abundant subaquatic vegetation helps the development of black fly larvae. It has
also been shown that this vegetation notably reduces the flow and speed of the water
current that negatively impacts the effect of the biocide (LACEY & UNDEEN, 1988). The
reduced flow not only limits the spread of the biocide, but also facilitates its adsorption by
the vegetation and the periphyton. Consequently, it requires a higher dosage and increased
frequency of application if vegetation is present. It was estimated that it would be
20 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

necessary to apply the product at shorter intervals, requiring a 15- to 20-fold higher rate
of application in certain areas.
In large-scale control programs, the impact on the environment is of great
importance, particularly when the control measures are used in recreational areas or in
areas that are a source of potable water. For this reason, regulations exist in numerous
countries concerning the use of chemical or biological products in watercourses in order
to prevent the contamination or adverse effect on the quality of the water. Bacillus
thuringiensis var. israelensis has been registered as a larvicide that does not produce adverse
effects among the 10 most important groups of invertebrate aquatic organisms associated
with black flies (W.H.O., 1982).
To obtain the parameters to define a control strategy using Bti in the field, bioassays have to
be carried out under laboratory conditions, keeping in mind the susceptibility of local species to
different dosages, times of exposure, distances and the possible timing of application.

Laboratory trials
The system and procedure for testing B.t.i. are as follows. A rearing tank consists of
a system of six flasks (5 for trials and 1 for control), with capacity of 2000 ml
(approximately) each filled with 1000 ml of water collected from the same stream where
the larvae were collected (COSCARÓN ARIAS, 1998a). The use of water originating from the
appropriate streams assures that there is sufficient food for the larvae during the
experiments. A system of aquarium air pumps oxygenates the water with diffusion stones
to create a turbulence that maintains a constant agitation in the water. One hundred larvae
of the different instars, preferably not of the last instar, are placed in each flask, using
flexible forceps to avoid damaging the larvae. The suggested dosages of B.t.i. are 5.0, 7.5,
10.0, 15.0 and 25.0 ppm, with exposure times of 1, 5, 10, 30 and 60 min, respectively.
Each of the five experimental flasks, along with the control flask, has to be labeled
according to the time the larvae are to be exposed. The appropriate quantity of B.t.i.
suspension is introduced into the flasks by means of a pipette, while maintaining uniform
agitation in the flask. Once the pipette has been completely emptied, the time is recorded.
After the specified exposure time period has elapsed, the water with the B.t.i. is filtered
carefully to avoid the loss of larvae; next, the water is replaced with 1000 ml of water
from the collection site.
After 24 hours, the larvae, along with the possible pupae that formed in each flask,
are counted. Finally, the percentage relationship of larvae that survived is calculated.
The experiments are repeated three times with the same five dosages. Mortality data
should be discarded for the trials in which the larvae from the control flasks exceed 10%.
Results of the tests that have a mortality rate between 5 and 10% can be corrected by
means of Abbott’s formula (COSCARÓN ARIAS, 1998a).

Field trials
The laboratory trials provide a basis for determining the concentration of B.t.i. and
exposure time necessary during the field studies. The effect that the B.t.i. will have on the
larvae will depend on the environmental conditions that determine the maximum
dispersal distance and settling rates along the watercourses.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 21

Techniques, such as the one used by GRAY et al. (1999), are recommended. These
techniques entail the application of the B.t.i. in concentrations according to the laboratory
results. Prior to the application of the B.t.i., the velocity and flow of the water and the
temperature must be recorded. To determine the distance that the B.t.i. carries downstream
and the effect on the larval population, samples should be collected between three and five
hours after the application at different distances downstream from the site of application (to
be determined according to the characteristics of the watercourse). The samples will be
taken 500 m beyond where the last larvae are observed to have been affected by the B.t.i. At
each observation point, a sample of the substrate with larvae stuck to it will be placed in a
container with one liter of water and placed in a cooler and refrigerated for transport to the
laboratory. Once in the laboratory, aerators are placed in each container, and after 24 hrs the
larvae are recounted and the percentage mortality is calculated.
Adults are collected in the field at the time of application, as well as at various
intervals after treatment, using the same technique, in order to compare the effects on the
population after the application of B.t.i. The dates and times are recorded so that the trials
can be repeated in the same area in the future.
Based on the results obtained from the trials and the observations made, the trials must
be repeated to determine the time taken for repopulation between the controls events.
22 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

2
MORPHOLOGY AND CHARACTERS
OF TAXONOMIC VALUE
The description of the various characters will be presented in the order used in the species
diagnoses, beginning with the female and followed by the male, pupa and larva.

Adults: The body size is relatively small (2.2-6.0 mm long), but the wing length generally is
used and does not change when the specimen dries out. The color is generally blackish gray
to yellowish brown; commonly the female shows lighter coloration than the male. The
head is well developed with compound eyes (Fig. 1 B: o) occupying a large portion of the
cephalic capsule; they are composed of numerous ommatidia. Males have proportionally
larger eyes (Fig. 1F), with upper macrofacets (mc) and lower microfacets (mi). Female
eyes are separated by the frons and basally by the clypeus (Fig. 1B: c). At the base of the
frons are the insertions of the antennae, where the eyes have an area without ommatidia,
the fronto-ocular triangle (fo) (nudi-ocular area) of different depths. The male is holoptic,
with the frons highly reduced. Some males are dichoptic, such as Gigantodax femineum, having
an obvious frons. Males with different frons widths generally are intersexes (= specimens
with male and female characters), produced by parasites, especially mermithids. Intersexes
can be observed in different species, the genitalia showing parts of both sexes. Antennae
(Figs. 1 A: a, B) are relatively short, composed of 11 segments (exceptionally 10, as in
Paraustrosimulium anthracinum) of similar shape, although the two basal segments, scape (s)
and pedicel (p), can be more elongate and in some species living at high altitudes (over
3000 m asl), they show long hairs (Figs. 38 P, Q). The apex of the head is the proboscis
(Fig. 1 B) with relatively short mouthparts. Mandibles (md) and laciniae (lc) have serrations
on the margin of one (Fig. 26 D) or both sides (Fig. 13 D) in females when they are
functional. These serrations are absent or vestigial in males and non-haematophagous
females. The lacinia is connected with the maxillary palp (mp) (Fig. 1 B: mp), which has
five segments or palpomeres; the third palpomere has the sensory vesicle (sv), an organ
with tubercles and which opens to the outside. The sensory vesicle is bigger in females and
is of a different size in each species. Mandibles and laciniae are covered dorsally by a
subtriangular plate, the labrum (lr). The clypeus is dorsal and the labium is basal and
lateral, presenting the flexible labella (la) apically. Internally are the cibarium and
hypopharynx. The basal portion of the cibarium (Fig. 1C) has two well-sclerotized branches
laterally, the cornuae (co), and a median portion that can be smooth (Fig. 15 F) or have
denticles in different positions and with different shapes, showing great variation in
Neotropical species (Figs. 76 X, N; 80 D, P, Y; 84 I-W).
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 23

M1 F
Rs
M2
mc ma

A R1
Cu1
R+M mi
M
sb
Sc
C Cu2
Bas An1
st
An2
pr abdt
ast G

an m tr
mp k ps cx h f t
pl
E
ba tm
ca cl

pe

D
fl fo
p o
ca
s
c
mp co
lr sv
lc md

la
B C
Figure 1. A: Schematic black fly female in lateral view (a = antenna, abd t = abdominal tergites,
an = anepisternum, ba = basitarsus, cl = claw, cx = coxa, f = femur, h = halter, k = katepisternum,
m = mesoepimeron, mp = maxillary palp, pl = pleural sulcus, pr = pronotum, ps = posterior
spiracle, st = scutum, t = tibia, tm = tarsomeres 1, 2, 3, 4, tr = trochanter; wing veins: An1 = anal 1,
An2 = anal 2, Bas = basal cell, C = costa, Cu1 = cubitus 1, Cu2 = cubitus 2, M = media, M1 = media
1, M2 = media 2, mc = medial-cubital fold, R1 = radius 1, Rs = secondary radius (radius 2), R+M =
radial-medial, sb = basal sector of radius, Sc = subcosta). B: Schematic female head in frontal view
(c = clypeus, fl = flagellum, fo = fronto-ocular triangle, la = labellum, lc = lacinia, lr = labrum,
md = mandible, mp = maxillary palp, o = compound eyes, p = pedicel, s = scape, sv = sensory
vesicle). C: Basal portion of cibarium with denticles, S. (In.) subnigrum (co = cornuae). D: Basitarsus
apex and tarsomere 1 (ca = calcipala, pe = pedisulcus). E: claw with small subbasal tooth. F-G.
Schematic male head; F: lateral view (ma = macrofacets, mi = microfacets). G: frontal view.
24 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

The thorax is well developed with great variation in prescutum (pr) and scutum (st)
coloration as well as hair microstructure and arrangement (Figs. 5 A, B). The scutellum
and metanotum (postnotum) have uniform morphology but variable coloration. The
pleurae are generally without hair, except in Tlalocomyia, which presents some on the upper
third of the pleural membrane (Fig. 10 B). The anepisternum (an) and katepisternum
(k) can show some variation in sheen. The pleurae have anterior stigmata (= spiracles) (ast)
and posterior stigmata (ps) on the upper area. The wings (Fig. 1 A) are wide, with a hyaline
membrane covered by microtrichia, and along the anterior margin they have well developed
veins with hairs and setae. The costa (c), subcosta (Sc) and radius (R), all end before the
wing apex. The radius is bifurcated; the anterior branch is called R1 and the posterior is the
radial sector (Rs ); the portion before the bifurcation is the basal sector of the radius
(sb). The following veins are not readily evident. The media is bifurcated (M1 and M2).
The median and cubital fold (mc) represent the false vein. The cubital vein is also bifurcated
(Cu1 and Cu2) and two anal veins (An1 and An2) are present. Cu2 is generally curved but in
Gigantodax and Pedrowygomyia it is straight. A basal cell (Bas) at the base of M can be
present or not. Halteres (h) do not usually show variation in shape or coloration and are
not used to differentiate species. Legs are relatively short and robust, with variable
pigmentation and thickness, which is more accentuated in males, especially on the tibia
and basitarsus. The legs have the following segments: coxa (cx), trochanter (tr), femur
(f), tibia (t), basitarsus (ba), four tarsomeres (tm) and claws (cl). The legs are covered
with hair with variable morphology, from filiform to flattened (Figs. 5 C, D). The hind
basitarsus can have a flattened expansion on the apex, the calcipala (Fig. 1 D: ca), and
tarsomere I can have a dorsal incision, the pedisulcus (Fig. 1 D: pe). The male claws have
different morphology than those in the females and are always very uniform. Female claws
vary greatly in shape among different taxa, and may or may not have a basal or subbasal
tooth of various sizes (Figs. 1 E; 5 E, F).
The abdomen is robust and can have different colors, similar to the thorax, particularly
on the tergal area that also is well sclerotized; the pleurae have extensive membranous
areas. The sternites are less developed than the tergites. The abdomen has only nine evident
segments; the other two are transformed into genitalic organs. The first abdominal segment
is short, scale-like and with long hairs.
The female terminalia (Figs. 2 A, B) are composed of the eighth sternite (VIII st),
a wide sclerite with abundant hairs laterally, and which project medially as a pair of foliaceous
gonapophyses (go); the cerci (c); and paraprocts (p) (= anal lobes), all with various
morphologies. Internally are the genital fork (g) and the spermathecal duct (sd). The
genital fork has three branches and can have anterior apodemes on lateral branches, as in
Cnesia, Pedrowygomyia and Gigantodax (Figs. 17 H, Q; 20 I, J). The spermatheca (Fig. 2 A: sp)
is well sclerotized and subovoidal, with a neck at the union with the spermathecal duct that
can be sclerotized or not.
The male terminalia are composed of the gonopods and the aedeagus with associated
parts (Fig. 2 C). The gonopods are comprised of two pieces; the basal one is the gonocoxite
(gc) (= basimere or basistyle) and the distal one is the gonostylus (gs) (= distimere or
dististyle). These show distinct interspecific morphological variations. The gonostylus
generally has one apical or subapical spur (= spine) or several (2-5 or more). The aedeagus
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 25

sd

c
p
go
VIII st
g
sp

gc
vp
C

ms gs
a

Figure 2. A: Schematic female terminalia, lateral view, Simulium sp. (c = cercus, g = genital fork,
go = gonapophysis, p = paraproct, sd = spermathecal duct, sp = spermatheca, VIII st = eighth
sternite). B: Female genitalia, ventral view (left cercus and paraproct) mounted on slide position, S.
(E.) romanai. C: Male genitalia, ventral view, S. (E) mayuchuspi (a = aedeagal membrane, gc =
gonocoxite, gs = gonostylus, e = endoparamere, ms = median sclerite, vp = ventral plate).

can be associated with several sclerotized structures such as the ventral plate (vp)
transversely positioned and with variable morphology; the median sclerite (ms) on the
middle of the genitalia, showing a distal, deep incision; and the endoparameres (e) (=
parameres), composed of 1+1 well sclerotized branches, frequently with robust teeth (=
hooks or spines) on the distal portion. Bordering the muscular aedeagus is the aedeagal
membrane (a), with abundant minute spiculae.
26 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Pupa: Size variable (2.0-8.0 mm); the measures that are generally used are the cocoon
length dorsally and basally and the gill length. The pupal body and frequently the gills are
protected by a cocoon, which is secreted by the larva. The cocoon weave can be compact,
loose, or spongy,, sometimes not showing the threads. The common shape is that of a
slipper or shoe, with great variations such as ventral or dorsal projections, dorsal prominences
(e.g., a carina), or reduction to only a posterior covering of the pupal abdomen. The pupal
body shows great uniformity. It has a resistant integument, generally well sclerotized, with
yellowish to dark grayish brown tones. It has tubercles, trichomes, and spines, with great
variety of shape, size, number and position. The trichomes can be single or multibranched,
generally filiform (Figs. 3 B, C), but sometimes flattened (Figs. 33 S, T) or star shaped (Figs.

g
A
c

fr

B
C

rf
g

Figure 3. Black fly pupa, S. (Pt.) limay. A: lateral view (c = cocoon, g = gills). B: frontoclypeus with
tubercles and trichomes (f = facial trichome, fr = frontal trichomes). C: thoracic plate with gill (g),
respiratory filaments (rf) and trichomes. D: abdominal chaetotaxy.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 27

53 S, T); tubercles can be flattened or elevated with smooth or wrinkled surfaces or apically
acuminate; the spines can be isolated or grouped like a comb, with one or several branches,
straight or curved like a hook (Fig. 3 D). The frontoclypeus, the antennae and eye sheaths
are evident on the cephalic plate. The frontoclypeus can show great variation according to
the number, type and disposition of tubercles. The trichomes are generally the frontal (fr)
and facial (f) trichomes but can also include epicranial trichomes. For the thorax, special
attention is given to the dorsal exposed portion, with the presence or absence of tubercles
and different types of trichomes. The abdomen has nine evident segments, ending with a
pair of spines (Figs. 6 E, F). The tergal plates and sternal, pleural or intersegmental
membranes have different integumentary processes such as trichomes, spines and hooks
that are of taxonomic value (Fig. 3 D). On the anterior portion of the pupal body, there are
two prothoracic expansions (with a respiratory function), the gills (g) (Figs. 3 A, C). The
gills have great morphological variations and are very important for taxonomy. Generally,
the gills have a basal trunk from which emerge several filamentous branches with arborescent
disposition (Fig. 6 A). These filaments are sclerotized with varied porosity and are hollow
internally; externally they are smooth (Figs. 6 B, C) or with furrows (Fig. 6 D); the gill apex
can be acute (Fig. 6 B) or blunt (Fig. 6 C). The gill also can show thick and asymmetrical
branches with varied morphology (Figs. 66 L, M; 70 A-C, G-I, P-R; 122-124). The number
of filaments varies from one to more than two hundred; the numbering of branches is
from the dorsal to ventral position, following a clockwise direction. The branches emerging
from the primary branches are the secondary branches, and those branching from the
secondary branches are the tertiary branches, and so on.

Larva: Maximum length in the last larval stage varies from 4.0 to 12.0 mm. The larvae have
a subcylindrical shape with a well differentiated head; the posterior third of the body is
more expanded (Fig. 4 A). Coloration is brown, gray, yellow to greenish with distinct intensity
and tones from light to dark, especially on the dorsum and head. The lateral thoracic area
can have spots from light to dark brown corresponding to the appendage histoblasts and
especially to the gill histoblasts. On the ventral part of the body, it is often possible to see
the ventral nerve cord and the prothoracic ganglion at the base of the head. The head has
a hard consistency due to the sclerotized cephalic capsule. Generally it has a constant
coloration, although it is lighter when ecdysis has been recent. Unique coloration is shown
dorsally on the cephalic apotome (= frontoclypeal apotome) (Fig. 4 B: ca), a median plate
delimited by the ecdysial line (el). Generally the cephalic apotome has several spots arranged
in groups that can be positive or negative. According to their position, they are referred to
as anteromedial (as), posteromedial (pm), anterolateral (al) and posterolateral spots
(pl). Lateral to the ecdysial line on each side is the postgena (pg), with ocelli (o) in the mid
area. Anteroventrally, there is a projection, the hypostomium (Fig. 4 C: h) with well-
sclerotized teeth on the anterior margin (Fig. 4 D). These teeth are the median (md) (one
tooth), intermediate (i) (= sublateral) (generally 3 teeth on each side), lateral (l) (one
tooth on each side), and paralateral (pl) (several teeth) followed sometimes by serrations.
Anterolaterally, the hypostomium has a row of variably numbered setae in a diagonal
arrangement. Separating the hypostomium from the postgena is the hypostomial groove
and the space between this sulcus and postgenal cleft (pc) is the postgenal bridge. The
28 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

gh rp
a
ap
A
vt B
tp md i
l
a
pl m

s
D
cs
G
lm as el
o pa al o
in
ap pm pg
ms
pl ca F
po
H ce
r

l
m

mp
d mx
lh
me h sf

E pb
p
C
pc
t cf

Figure 4. Black fly larva. A: schematic general view in lateral position (a = anal sclerite, ap = abdominal
proleg, gh = pupa gill histoblast, rp = rectal papillae, tp = thoracic proleg, vt = ventral tubercles). B:
schematic head, dorsal view (a = antenna, al = anterolateral spots, as = anteromedial spot, ca = cephalic
apotome, ce = cervical sclerite, cs = cephalic fan stem, el = ecdysis line, m = mandible, o = ocellus, pg =
postgena, pl = posterolateral spots, pm = posteromedial spots, po = postocciput). C: schematic head,
ventral view (cf = primary cephalic fan, h = hypostomium, lh = labiohypopharynx, m = mandible,
mp = maxillary palp, mx = maxilla, pb = postgenal bridge, pc = postgenal cleft, r = cephalic fan rays ,
sf = secondary cephalic fan, t = tentorial pit). D: hypostomium, anterior border (i = intermediate teeth,
l = lateral tooth, md = median tooth, pl = paralateral teeth, s = lateral serrations). E: antenna, S. (Ch.)
subpallidum (d = distal, me = medial, p = proximal). F: maxillary palp. G: mandible apex, S. (Ec.) albanense
(in = inner teeth, lm = latero-mandibular process, ms = mandibular serrulations, o = outer teeth, ap=
apical tooth, pa = preapical teeth). H: anal sclerite with abundant hair, S. (Ec.) albanense.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 29

A B

C D

E F
Figure 5. Female. A-B, D-E: S. (N.) exiguum; A: scutum hair disposed in groups; B: scutum scale-
like seta; D: filiform and scale-like setae of leg; E: claw without subbasal tooth. C: S. (Pt.) stelliferum,
hind leg with filiform setae; F: Gigantodax clandestinus, claw with subbasal tooth.
30 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

A B

C D

E F
Figure 6. Pupa. A-B: Gigantodax igniculus; A: gill; B: apex of gill branch. C: G. marginalis, apex of gill
branch. D: S. (Tr.) lahillei, apex of gill branch. E-F: G. igniculus, abdomen apical portion in lateral and
dorsal view showing terminal spur, pleural membrane, tubercles, spine combs and hooks.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 31

postgenal cleft shows variation in shape and depth and has taxonomic importance. At the
postgenal base, ventrally on each side, is the posterior tentorial pit (p), and all around the
posterior margin is the well sclerotized postocciput (po). The postocciput, over the median
cephalic portion of the apotome, is thinner, and has the cervical sclerites (ce), which
meet the postocciput. Below the epidermis and near the postgenal cleft is the subesophageal
ganglion, which is sometimes darkly pigmented.
Dorsolaterally on the anterior portion of the head are the prominent cephalic fans
(= labral fans) (Fig. 4 C: cf), with a robust cephalic fan stalk (Fig. 4 B: cs) and variable
number of rays (r). Cephalic fan rays have thinner teeth arranged like a comb, showing
variable length and position (Fig. 7 B). The largest fan rays are the primary rays which
number from 20 to 80; there are other smaller ones know as secondary rays that are fewer
in number and located more basally (Fig. 4 C: sf); a third type consists of short, flat, straight
blades, the median rays, that form the scale fan (sensu CROSSKEY, 1990: 146). Behind the
cephalic fan are the antennae (Fig. 4 B: a), composed of three segments (Fig. 4 E): proximal
(p), medial (me) and distal (d) ending in a short and acute sensillum. Relative proportions
of segment lengths are useful in taxonomy.
Ventrally there are several mouthparts. Apicoventrally, the mandibles (Fig. 4 C: m) have
several brushes and teeth (Figs. 4 G; 7 E). The teeth are more important taxonomically,
showing variation in number and size. The outer teeth (o) are the first 2 or 3 large teeth.
Following these is the largest one, the apical tooth (ap) and it is followed by the preapical
teeth (pa) and the inner teeth (in) that can be present in several rows, and lastly the marginal
serrations (ms) which are also variable in number. Some species can have a latero-mandibular
process (lm) that can be single or double and cylindrical or a little flattened. In some
supraspecific taxa, a row of supramarginal setae (Fig. 79 B) can be present.
The larvae have maxillae with abundant long hairs (Fig. 4 C: mx); the maxillary palp
(mp) is cylindrical (Fig. 4 F), with apical sensillae of variable morphology (Fig. 7 A). The
labrohypopharynx is setose and has a uniform appearance (Fig. 4 C: lh). The labrum is in
front of the head and mostly covered by setae having little taxonomic importance. The
body is slightly curved ventrally but it can be strongly curved while moving from one place
to another. It shows little segmentation particularly on the thorax. The body integument is
generally covered by microscopic trichomes but can show different epithelial processes of
varied morphology, some of them being of larger size (Figs. 4 D; 135 G, L, S). Trichomes
and scales are frequently found around the anal sclerite (Fig. 4 H). On the thorax are
whitish histoblasts that correspond to wings, legs and pupal gills (Fig. 4 A: gh). Ventrally
there is an elongated cylindrical appendage, the thoracic proleg (= prothoracic proleg)
(Fig. 4 A: tp). On its apex, it has a row of hooks and subapically 1+1 sclerotized, subtriangular
to subtrapezoidal lateral sclerites, with a comb-like structure of elongated teeth frequently
arranged in groups (Figs. 7 F; 8 X; 12 M).
The larval abdomen has external pseudoannulations, sometimes evident as thin, pale
rings. Laterally, there are minute rounded dark spots corresponding to nonfunctional
spiracles. Posteriorly, on the ninth segment there are 1+1 ventral tubercles (= ventral
papillae) (Fig. 4 A: vt) with variable presentation in different taxa. On the posterior apex is
the abdominal proleg (ap) (= posterior proleg), which looks like a sucker, bordered by
hooks arranged in rows and disposed around the proleg as the anal ring (= posterior
32 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

A B

C D

E F
Figure 7. Larva. A: Gigantodax herreri, apical sensillae of maxillary palp. B: Gigantodax sp., microtrichia on
cephalic fan of primary ray. C: S. (Ps.) auripellitum, terminal portion of abdomen showing rectal papillae,
abdominal proleg, ring of hooks and cuticular hair. D: Simulium sp., cuticular scale-like hair. E-F: G.
igniculus. E: mandible portion showing teeth. F: apex of thoracic proleg showing lateral sclerite.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 33

circlet) (Fig. 7 C). The number of rows of hooks is variable (40 to more than 300) and is
very useful in taxonomy. Anterior to the abdominal proleg is a well-sclerotized structure,
the anal sclerite (a) (Fig. 4 A: a). In Neotropical species, it has an X shape with ventral
branches longer than the dorsal ones (Fig. 4 H). In some taxa, such as Gigantodax,
Pedrowygomyia and partially in Hearlea, the ventral branches are very elongated and joined
ventrally to form a circle (Fig. 20 S) around the anal ring. In some species of Hearlea, there
also are well-sclerotized accessory plates with different shapes near the anal sclerite (Figs.
127 B-J). Dorsal to the abdominal proleg is the anus. From the distal portion of the rectum,
larvae can extrude a membranous, elongate organ, the rectal papillae (Fig. 4 A: rp). Basically,
the rectal papillae is composed of three cylindrical lobes commonly present in the primitive
simuliid groups; as an apparent apomorphic variation each lobe can have one or more
diverticula (= lobules) on each lobe (Fig. 7 C). There can be over 30 diverticula on each
lobe. The rectal lobes frequently remain inside the larva after fixation and it is not possible
(without dissection) to see this character, which has some taxonomic value.

Eggs: The eggs are ovoid to subtriangular and uniform in color, which varies from light
yellowish gray to dark gray, depending on the time of ovoposition. There are no published
studies on Neotropical simuliid eggs for species identification. It is possible to find eggs
when they are deposited in large masses where several females gather to oviposit. The
curved larva and its dark head with the ruptor ovi (= egg buster) dorsally positioned in the
center of the head can be seen in the eggs before eclosion.
34 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

3
COLLECTION AND PREPARATION
OF MATERIAL
Adults: The easier of the two sexes to obtain is the female because of the haematopha-
gous habits, which attract them to humans or domestic animals. Females can be collected
by nets or traps such as Malaise, Shannon or Manitoba traps, which are commonly used to
capture other blood feeding insects. When using a trap, it is useful to have an attractant
such as a person, an animal or CO2. To capture biting females, it is best to put a small vial
over the fly when it is feeding and not use forceps so as to avoid damaging the specimen.
For taxonomic studies, it is preferable to kill the flies and maintain part of the material
dry to be pinned afterwards in the lab. To kill adults, ethyl acetate, ether, chloroform,
cyanide, etc. can be used, but not applied over the specimen. It is desirable to place
pinned material in a freezer for at least five weeks to avoid having the abdomen retract.
The remaining material should be fixed in 80% ethyl alcohol and maintained as long as
it is necessary for further studies. It is better to see coloration in dry material; however,
if fixed material is to be used for the observation of color, it should be placed in ethyl
acetate for at least ten minutes and dried on filter paper to evaporate all the liquid. After
this, the hairs will become more erect and better show the characteristics. To collect
males, it is necessary to use a hand net and try to find a swarm or to catch them as they
fly above the aforementioned traps.

Larval stages: Larvae and pupae are found together in different types of freshwater
streams and on various substrates. Generally, they are found near the surface, rarely below
one meter in depth. The best place to find them is where the current is fast. The material
can be collected with forceps and maintained alive or can be fixed directly in alcohol or
other fixative. It is convenient to keep material alive in a vial placed in a cooler while
transporting it to the lab where the flies can be reared or fixed for future studies. For
taxonomic studies, it is best to keep them in 80% ethanol. Carnoy’s fixative (100% ethanol:
glacial acetic acid = 3:1) should be used for cytological studies or 100% alcohol for
molecular studies. For the last two studies, it is convenient to change the fixative after 30-
60 minutes and then keep the vials in a refrigerator. Color information should be obtained
from fresh material (before fixation). Pupae can be fixed and stored in 80% alcohol.
It is possible to rear black flies under laboratory conditions to obtain different stages.
Larvae can be reared if water current is assured. There are different methods, some of
which are more sophisticated (EDMAN & SIMMONS 1985 a, b), and others less so. One of
the easier ways to obtain pupae is to maintain the larvae in a vial with an air flux at the
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 35

base and with a magnetic stirrer. Pupae can also be reared to obtain imagoes. They can be
maintained in a vial on humid paper with a cotton stopper, avoiding water condensation
on the walls. It is convenient to rear pupae individually in order to identify the adults and
have associated pupal exuviae.
36 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

4
STUDY OF MATERIAL
Taxonomic studies start with the observation of external characters such as size,
coloration and morphology. It is necessary to use a stereomicroscope that can also be
used for dissections and specimen assembly on slides. Anatomical parts used for the
identification of specimens are chitinous. To clarify them adequately, it is useful to
digest the accompanying tissue with NaOH or KOH at about 5% to 10%. This can be
done by boiling the material for 30-60 seconds, but since this procedure can destroy
delicate structures, it is better to leave the material in the solution for 12-18 hours at
room temperature instead of boiling it. In adults, the wings should be removed first, to
avoid damage. In larvae, this treatment needs to be done in less time, and some delicate
parts, such as the rectal gill and antennae, are better seen without the use of these
treatments. After this procedure, it is necessary to neutralize the material with acetic
acid or phenol and dissection. The parts can be stored in glycerin or mounted on glass
slides in Canada balsam or Euparal. Before mounting, it is desirable to study the
different pieces with a compound microscope to see the different structures in diverse
positions and to gain better knowledge of the morphology. There are different methods
used to clear the pieces before they are slide mounted. A fast method is to transfer from
NaOH or KOH directly to phenol (= phenic acid) for 10-15 minutes for clarifying. It is
also possible to dissect the material at this time, with histological needles. Materials
treated with phenol can be observed with a compound microscope in temporary
preparations, or can be assembled on a slide for a permanent preparation. Phenol can
crystallize at low temperatures (in air conditioning). In this case, it must be warmed and
several drops of 100% alcohol added to the remaining liquid.
When the material needs to be cleared further after passing through phenol, it is
convenient to leave it for an hour in creosote. It is also possible to leave the material for
more than one day in the phenol or creosote without causing any damage. If the
material is in phenol for several hours, exposed to ambient conditions, it will be
necessary to replace it with new phenol, because it can hydrolyze, and the preparation
may be not clear enough. When the structures are too clear, they can be dyed with a
coloring solution. Leaving the material for 30 minutes or more in “lignin pink”,
dissolved in creosote gives good coloration.
Gill histoblasts can be seen best if they are transferred to acetic acid diluted at 10-
20%. This helps to uncurl the filaments and, therefore, makes it is easier to count them.
Phase-contrast microscopes and electron microscopes are useful for seeing some
structures. For electron microscopy, it is best not to use material that has been preserved
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 37

for a long time. To clean the adhered particles, ultrasonic equipment can be used with
some detergent. It is important to control time in order not to lose some parts such as
hairs or to break others. It is important to check the material after 10-15 seconds and
repeat the procedure if necessary. After using the detergent, the structures must be rinsed
with water, alcohol and acetone. Before affixing the specimen to the support for metal
coating, it is necessary to dry the material carefully with filter paper. During the analysis of
the material it is useful to make drawings for future comparisons.
38 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

5
SYSTEMATICS
Neotropical Simuliidae are all included in the subfamily Simuliinae, according to the
general classification used by ADLER et al. (2004). In the present volume, it comprises 12
genera and 359 species, following the arrangement given by WYGODZINSKY & COSCARÓN
(1973a). The first ten genera were previously considered in the tribe Prosimuliini, as
described by CROSSKEY & HOWARD (1997, 2004). The validity of Prosimuliini was
questioned by PY-DANIEL (l994). Presently, the most representative of these ten genera are
included in the tribe Simuliini (ADLER et al., 2004). The genus Simulium is the most
abundant with l8 subgenera, as described in COSCARÓN (1987). The CROSSKEY & HOWARD
(2004) taxonomic arrangement is very close, yet with some small differences in
Kempfsimulium, Psaroniocompsa, Ectemnaspis and Trichodagmia. These Simulium subgenera are
considered genera by PY-DANIEL & MOREIRA SAMPAIO (1994, 1995). Here, we will not deal
with this supraspecific category, considering that a study of the world black flies would be
the best forum to address this issue.
In this compilation, the species of each taxon are arranged based on the arrangement
given in previous cladistic studies, particularly those in COSCARÓN (1987); COSCARÓN &
COSCARÓN ARIAS (1996b, 1997); COSCARÓN et al. (1996, 1999, 2004); MIRANDA-ESQUIVEL &
COSCARÓN (2001, 2004); and STRIEDER & PY-DANIEL (2002).
Generic and specific diagnoses are restricted to those morphological characters that
permit taxon identification, and are accompanied by figures facilitating their identifica-
tion. Diagnosis is accompanied by geographic distribution, including states or provinces
of each country and their corresponding maps. Succinct bionomic information is
included with each taxon. We have attempted to compile most of the available
information, but recognizing the vast extent of this realm, with areas yet to be explored or
collected, we consider that the information presented corresponds only to a snapshot
view of the Neotropical black flies.
With this volume we intend to aid researchers that are immersed in the complex
world of limnology and who are trying to understand the interrelationships of these
systems while trying to protect them from the damage “progress” has caused. Also, we
aim to help those that try to control nuisance or disease-bearing black flies. If rational
control is planned, it is necessary to know which species are pests and which are
innocuous species in order to save the latter which play an important role in competing
with the more dangerous species.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 39

KEYS TO THE GENERA OF NEOTROPICAL SIMULIIDAE


Adults
1 Pedisulcus well developed (Fig. 1 D: pe); basal cell invariably absent; length of
basal section of R (as measured from humeral cross-vein) less than 1/3 distance
from base of Rs to wing tip ................................................................................ Simulium
– Pedisulcus absent (Figs. 8 G, N); basal cell absent or present; length of basal
section of R about 1/3 distance mentioned or more ................................................ 2
2(1) Cu2 and A1 straight (Fig. 20 C); basal cell absent ........................................................ 3
– Cu2 and A1 curved (Fig. 8 E); basal cell absent (Fig. 26 F) or present (Figs. 8 E; 15 G)
................................................................................................................................................ 4
3(2) First flagellomere as long as wide; hair of basal flagellomeres and palpomeres
shorter than segment; clypeus longer than wide (Fig. 20 B); female mouthparts
normal, functional; calcipala present (Fig. 20 E); subbasal tooth on claw well
developed (Fig. 20 G) ....................................................................................... Gigantodax
– First flagellomere as long as wide; hairs of basal flagellomeres and palpomeres
longer than segment; clypeus as long as wide (Fig. 20 A); female mouthparts
reduced, non functional; calcipala absent (Fig. 20 D), subbasal tooth on claw
obsolescent (Fig. 20 F) .................................................................................Pedrowygomyia
4(2) Upper portion of pleural membrane hairy (Fig. 10 B); apical segment of maxillary
palp more than twice as long as penultimate segment .............................. Tlalocomyia
– Pleural membrane entirely bare; apical segment of maxillary palp generally less
than twice as long as penultimate segment (Figs. 13 C; 15 C; 21 B) ...................... 5
5(4) Antennae with 10 segments each; arms of furcasternum with conspicuous
projections (Fig. 23 A); male genitalia with median sclerite geniculate, composed
of slender basal Y-shaped portion and 2 subparallel elongate apical arms (Fig. 23
H); apical portion of endoparameres with numerous elongate denticles; genital
fork of female almost entirely unpigmented, its stem very stout, not longer than
arms (Fig. 23 E) .......................................................................................Paraustrosimulium
– Antennae with 11 segments each (Fig. 19 A); arms of furcasternum lacking
conspicuous projections (Figs. 8 D; 13 F) [except Lutzsimulium (Fig. 24 L)]; median
sclerite of male not as above, or apical portion of endoparameres obsolescent,
lacking denticles (Figs. 8 Q; 24 I); genital fork distinctly pigmented, with stem not
as above (Figs. 8 L; 10 E; 11F; 13 M; 17 H, Q) .......................................................... 6
6(5) R1 only with hair-like setae, spiniform setae not developed (Fig. 21 F); basal tooth
of claw of female large (Fig. 21 J) ................................................................ Cnesiamima
– R1 with hair-like and spiniform setae (Figs. 13 H, I); if spiniform setae not very
distinct, then basal tooth of claw of female small (Fig. 24 D) ................................ 7
7(6) Basal cell absent (Fig. 26 F); mandible of female toothed only on internal margin
(similar to Fig. 26 D); claws of female with small subbasal tooth (Figs. 24 D; 26
K); male genitalia with teeth of endoparameral organ not perceptible (Figs. 24 I;
27 A) ..................................................................................................................................... 8
– Basal cell present (Fig. 8 E); mandible of female as above or with well-developed
teeth on internal and external margins (Fig. 13 D); claws of female with larger
40 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

basal tooth (Figs. 13 G; 15 I); male genitalia with teeth of endoparameral organ
distinct (Figs.11K; 13T; 17K: ed) .................................................................................... 9
8(7) Frons longer than wide with median sulcus relatively short and without basal
bifurcated branches (Fig. 24 A); Sc with about 50 setae (Fig. 24 H); furcasternal
branches with large projections (Fig. 24 L) .............................................. Lutzsimulium
– Frons about as long as wide with median sulcus elongated and bifurcated, with
long basal branches (Fig. 26 A); Sc without setae (Fig. 26 F); furcasternal branches
with very small projections (Fig. 26 L) .................................................... Kempfsimulium
9(7) Wings with slight curvature of Cu2; A1 almost attaining wing margin (Fig. 17 B);
male genitalia with basal portion of endoparameres obsolescent, denticles of
distal portion well developed but few in number, forming tight group of
characteristic arrangement (Fig. 17 K); female with arms of genital fork bearing 2
forwardly directed apodemes (Figs. 17 H, Q) ......................................................Cnesia
– Wings with pronounced curvature of Cu2; A1 ending well before wing margin (Fig.
8 E); endoparameres of male not as above; genital fork of female without
apodemes (Figs. 8 L; 11 F) ............................................................................................. 10
10(9) Males ................................................................................................................................... 11
– Females ............................................................................................................................... 13
11(10) Median sclerite with basal transverse satellite plate continuous distally with deep
cleft and bifurcated apically (Fig. 8 R); endoparameres lacking teeth (similar to Fig.
8 Q) .................................................................................................................... Mayacnephia
– Median sclerite of complex geniculate shape, with distal half divided into 2 wide
but short and divergent arms (Figs. 11 J, K; 13 R, S); endoparameres with
numerous teeth (Fig. 11 K) ............................................................................................ 12
12(11) Genitalia with ventral plate subtriangular and basal arms narrow, straight (Figs. 11
I; 13 P); arms of median sclerite parallel (Fig. 13 S); endoparameres with basal
portion large and denticles of apical portion well developed (Figs. 11 K; 13 T) ..
.............................................................................................................................. Araucnephia
– Genitalia with ventral plate subrectangular and basal arms stout, rounded at apex
and curved (Figs. 15 P, Q); arms of median sclerite sharply diverging apically (Fig.
15 U); endoparameres with basal portion small and denticles of apical portion
obsolescent .................................................................................................. Araucnephioides
13(10) Basal tooth of claws narrow, hook like (Fig. 15 I) .............................. Araucnephioides
– Basal tooth of claws larger, subtriangular or spatulate (Figs. 8 H, J; 13 G) ........ 14
14(13) Frons very narrow (Fig. 8 A), frontal angle approximately 50°; inner surface of
spermatheca without spiculae (Mesoamerica) .......................................... Mayacnephia
– Frons wider (Fig. 15 A), frontal angle approximately 85°; inner surface of
spermatheca with scattered spiculae (southern South America) ........... Araucnephia

Females, based on genitalia


1 Arms of genital fork with long cephalad-directed apodemes (Figs. 17 H, Q; 20 I, J)
................................................................................................................................................ 2
– Arms of genital fork lacking long apodemes .............................................................. 3
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 41

2(1) Anterior branch of genital fork of uniform width (Figs. 17 H, Q; 20 J) ................


.................................................................................................................. Cnesia, Gigantodax
– Anterior branch of genital fork expanded on basal 1/3 (Fig. 20 I ) ....... Pedrowygomyia
3(1) Genital fork almost entirely unpigmented, its stem wide and very short, not longer
than arms (Fig. 23 E) .............................................................................Paraustrosimulium
– Genital fork not as above ................................................................................................. 4
4(3) Sclerotization of spermatheca continuing onto base of spermathecal duct (Figs.
26 P, Q) ................................................................................................................................ 5
– Spermathecal duct inserted on small membranous area of spermatheca (Figs. 8 J;
21L) ....................................................................................................................................... 8
5(4) Area of insertion of spermathecal duct on spermatheca with network-like
structure extending to base of duct (Fig. 24 G) ......................................................... 6
– Not as above ....................................................................................................................... 7
6(5) Sclerotized area of spermathecal duct also covering spermathecal neck and portion
of spermathecal duct with meshwork (Fig. 24 G); spermatheca internally with
spiculae; eighth sternite with sclerotized area reduced to medial area (Fig. 24 E) ....
............................................................................................................................ Lutzsimulium
– Sclerotized area of spermathecal duct reduced to neck, without meshwork on
spermatheca union (Figs. 26 P, Q); spermatheca internally without spiculae; eighth
sternite with sclerotized area covering all area from one side to other (Fig. 26 M)
.......................................................................................................................... Kempfsimulium
7(5) Paraproct very small, triangular; surface of spermatheca conspicuously reticulate,
its inner surface without spiculae; sclerotization of spermathecal duct ending
abruptly (Fig. 10 D) .......................................................................................... Tlalocomyia
– Paraproct of normal size, rounded (Fig. 21 M); surface of spermatheca smooth,
its inner surface with few spiculae; sclerotized portion of spermathecal duct
gradually fading into unpigmented portion (Fig. 21 L) ............................ Cnesiamima
8(4) Inner surface of spermatheca without spiculae ....................................... Mayacnephia
– Inner surface of spermatheca with scattered spiculae ............................................... 9
9(8) Paraproct about as long as wide (Fig. 13 L); inner surface of spermatheca not
rugose ................................................................................................................. Araucnephia
– Paraproct about one half shorter than wide (Fig. 15 L); inner surface of
spermatheca rugose ................................................................................... Araucnephioides

Males, based on genitalia


1 Endoparameral teeth not discernible; median sclerite comparatively simple, wide
basally, elongate lamella, apically divided into 2 short arms connected by
membrane (Figs. 24 I; 27 A) ............................................................................................ 2
– At least some portion of endoparameral organ discernible; median sclerite
different, generally of complex structure ..................................................................... 3
2(1) Gonocoxite about 2 times longer than gonostylus; gonostylus gradually acuminate
distally, apex relatively blunt, with 3 (2) terminal teeth (Figs. 24 I, M); ventral plate
distally with soft rounded curvature; median sclerite very elongated (Fig. 24 J);
pedisulcus absent ............................................................................................ Lutzsimulium
42 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

– Gonocoxite about 3 times longer than gonostylus; gonostylus strongly acuminate


distally with 2 terminal teeth (Figs. 27 A-C); ventral plate showing irregular border
distally (Figs. 27 A, D); median sclerite wider, not elongated and distally gradually
acuminate; pedisulcus absent to well developed ................................... Kempfsimulium
3(1) Basal portion of endoparameral organ not discernible or only very weakly
developed; denticulate portion composed of 1 or 2 large and several very closely
associated, often oppositely directed smaller denticles (Fig. 17 K) ........................ 4
– Basal portion of endoparameral organ distinct, plate like; denticles of distal
portion, varying from absent to well developed, but not as described above ...... 6
4(3) Ventral plate subpentagonal, about as long as wide, not conspicuously emarginated
apically (Fig. 17 N) ......................................................................................................Cnesia
– Ventral plate in most cases elongate-rhomboid, with distinct apical emargination
................................................................................................................................................ 5
5(4) Ventral plate short, with lateral margins at midlength concave and apical margin
bilobed (Fig. 20 L); endoparameral teeth elongated .................................. Gigantodax
– Ventral plate long, about 2 times longer than wide, with lateral margins at
midlength straight and apical margin straight or convex (Fig. 20 K);
endoparameral teeth short ..........................................................................Pedrowygomyia
6(3) Median sclerite consisting of transverse basal flange and deeply incised median
portion perpendicular to flange (Figs. 8 R; 10 G); endoparameres reduced to basal
plate-like portion, and without denticles (Fig. 8 Q) ........... Mayacnephia, Tlalocomyia
– Median sclerite different; endoparameres with or without denticles ...................... 7
7(6) Median sclerite strongly geniculate in side view (Fig. 21 U), roughly M-shaped
when seen in full view (Figs. 21 V; 23 H), with inner arms of M partly fused and
connected to basal flange ................................................................................................. 8
– Median sclerite not as above, its shape complex and occupying 3 different planes
................................................................................................................................................ 9
8(7) Endoparameral organ with free plate-like basal portion and numerous elongate
subparallel apical denticles not connected to basal portion (Fig. 23 H) ..................
.....................................................................................................................Paraustrosimulium
– Endoparameral organ reduced to subquadrate plate-like structure; denticles not
developed (Fig. 21 U) ....................................................................................... Cnesiamima
9(7) Ventral plate subpentagonal or subtriangular (Fig. 11 I), its basal arms in shape of
elongate slender rods (Fig. 13 P); branches of median sclerite parallel (Fig. 13 S);
endoparameres with conspicuous denticles distally (Fig. 11K) .............. Araucnephia
– Ventral plate subrectangular, its basal arms stout, apically knobbed (Fig. 15 P);
branches of median sclerite diverging apically (Fig. 15 U); endoparameres with
denticles of distal portion obsolescent (Fig. 15 S) ............................. Araucnephioides

Pupae
1 Abdominal sterna VI and VII divided longitudinally along middle by membranous,
striate area (Figs. 14 D; 18 D); hooks of terga VI-VIII invariably simple; abdominal
segments VIII and IX in many cases with strongly curved, twisted, looped or
grapnel-like strong setae (Figs. 18 E; 23 J; 25 E) .......................................................... 2
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 43

– Abdominal sterna VI and VII entire; in many cases, some hooks on terga VI-VIII
bifid or trifid; abdominal terga VIII and IX without above-mentioned setae ........
................................................................................................................................... Simulium
2(1) Terminal processes of abdomen short and pointed (Fig. 23 J), or absent (Fig. 10 L)
................................................................................................................................................ 3
– Terminal processes of abdomen elongate (Figs. 12 E; 18 E; 25 E) ....................... 4
3(2) Cocoon reduced to small pad on which terminal abdominal segments are inserted
(Fig. 10 H); gills shaped like thick stem with few thread-like filaments (Fig. 10 I);
abdomen strongly sclerotized, terga and sterna with large number of
supernumerary spines or hooks (Figs. 10 J, K); apex of abdomen blunt; terminal
processes absent (Fig. 10 L) ............................................................................ Tlalocomyia
– Cocoon well developed, of definite shape, covering whole body of pupa except
gills, latter in shape of twisted pseudo-segmented lamella (Figs. 22C; 23 I); abdomen
less sclerotized than thorax; abdomen without supernumerary hooks, and with short
terminal processes (Fig. 23 J) ........................................... Cnesiamima, Paraustrosimulium
4(2) Abdominal terga with spine-combs (Fig. 14 D); tergum VIII without large hooks
................................................................................................................................................ 5
– Abdominal terga without spine-combs; tergum VIII with 4 large hooks ..... Cnesia
5(4) Facial trichomes (Figs. 12 B; 27 E), thoracic trichomes (Figs. 12 D; 27 F, G), and
setae of eighth and ninth abdominal segments tightly looped apically (Fig. 25 E)
................................................................................................................................................ 6
– At least facial and thoracic trichomes not looped apically........................................ 7
6(5) Gill with 12-22 branches; frontoclypeus and thorax with tubercles (Fig. 25 D) ....
............................................................................................................................ Lutzsimulium
– Gill with 12 branches; frontoclypeus and thorax without tubercles (Fig. 27 E) ....
.......................................................................................................................... Kempfsimulium
7(5) Head sclerite with frontal, facial, epicranial, lateral, and genal hair-like trichomes
(Figs. 12 B; 14 B) .............................................................................................. Araucnephia
– Head sclerite not with all trichomes mentioned; facial trichomes hair-like or spine-
like ......................................................................................................................................... 8
8(7) Thorax with tubercles arranged in circles (Fig. 16 E); basal portion of gills with
minute plate-like cuticular structures; thoracic trichomes hair-like .......................
........................................................................................................................ Araucnephioides
– Thorax with tubercles not arranged in circles; basal portion of gills without
aforementioned cuticular structures; thoracic trichomes hair-like or spine-like
................................................................................................................................................ 9
9(8) Cephalic sclerite with 2+2 or 3+3 frontal trichomes; branches of gills more or less
tubular (Figs. 9 A-J) ........................................................................................ Mayacnephia
– Cephalic sclerite with 1+1 or without frontal trichomes; gill branches varied in
shape ................................................................................................................................... 10
10(9) Frontal and labial-frontoclypeal trichomes absent (Fig. 20 N); clypeus
comparatively narrow (Fig. 20 B) .................................................................. Gigantodax
– Frontal and labial-frontoclypeal trichomes present (Fig. 20 M); clypeus
comparatively wide (Fig. 20 A) ...................................................................Pedrowygomyia
44 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Larvae
1 Anal sclerite with accessory sclerite forming complete ring around posterior end
of body (Fig. 20 S); mandibles each with 3 outer teeth (Fig. 20 P) ........................ 2
– Ring-shaped accessory sclerite not developed (Fig. 12 N); mandibles each with 2
(Fig. 12 J) or 4 (Fig. 18 J) outer teeth ............................................................................ 3
2(1) Hypostomium with lateral tooth, projecting beyond level of first intermediate
tooth (Fig. 20 R); cephalic apotome without basal spots ......................... Gigantodax
– Hypostomium with first intermediate tooth generally projecting beyond level of
lateral tooth (Fig. 20 Q); cephalic larval apotome with basal spots (Fig. 20 O) .....
...........................................................................................................................Pedrowygomyia
3(1) Teeth of hypostomium arranged in 3 conspicuous groups (Fig. 8 W); mandible
with marginal serrations numerous, more than 5 (Fig. 12 J) .................................... 4
– Teeth of hypostomium more evenly distributed, not arranged in conspicuous
groups (Figs. 18 H; 23 M; 25 I); mandible with marginal serrations less numerous,
not more than 5, generally only 2 (Fig. 25 K) ............................................................. 7
4(3) Antennae approximately as long as stem of cephalic fan (Fig. 16 I); hypostomium
with either 13 (Fig. 8 W) or 17 teeth (Fig. 16 N) ........................................................ 5
– Antennae much shorter than stem of cephalic fan (Fig. 14 H); hypostomium with
15 teeth (Fig. 14 K) .......................................................................................... Araucnephia
5(4) Cervical sclerite very small, isolated in membrane (Fig. 16 H); mandibles with
accessory teeth at level of inner teeth, with second preapical tooth minute (Fig. 16
L) and with short basal setae slender, only faintly dentate; hypostomium with 17
teeth (Fig. 16 N) ......................................................................................... Araucnephioides
– Cervical sclerite small or transverse and isolated in membrane (Fig. 8 S), or transverse
and fused with upper ends of postocciput (Fig. 10 O); mandibles without accessory
serrations at level of inner teeth, with second preapical tooth as long as first (Fig. 10
M) or only slightly shorter (Fig. 8 U) and with short basal setae bearing conspicuous
elongated denticles (Fig. 8 V); hypostomium with 13 teeth (Fig. 8 W) ........................ 6
6(5) Proximal antennal segment more than half as long as medial segment (Fig. 8 T)
............................................................................................................................. Mayacnephia
– Proximal antennal segment at most half as long as medial segment .......... Tlalocomyia
7(3) Antenna much longer than stem of cephalic fan; distal antennal segment distinctly longer
than proximal and medial segments combined (Figs. 22 I; 23 L); backwardly directed struts
underlying main body of anal sclerite (Fig. 23 N) ................ Cnesiamima, Paraustrosimulium
– Antenna not longer than stem of cephalic fan (Fig. 27 H); distal segment not or
only slightly longer than proximal and medial segments combined (Figs. 18 G; 25
J); anal sclerite without struts (Figs. 18 M; 25 L) ........................................................ 8
8(7) Postgenal cleft very shallow (Fig. 19 N); hypostomium with 17 teeth, the 4+4
intermediate ones similar to remaining teeth (Fig. 19 M); mandibles with 4 outer
teeth (Fig. 18 J) ...........................................................................................................Cnesia
– Postgenal cleft well developed (Fig. 25 H); hypostomium with at most 13 teeth
(Figs. 25 I; 27 J), the 1+1 lateral ones, when perceptible, somewhat similar to
lateral serrations; mandible with 2 outer teeth (Figs. 25 K; 27 K) .......................... 9
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 45

9(8) Preapical teeth of mandible subequal in size (Fig. 33 X), or decreasing in size from
first to third (Fig. 38 G) ....................................................................................... Simulium
– Preapical teeth of mandible with second smaller than either first or third (Figs. 25
K; 27 K) ............................................................................................................................. 10
10(9) Body integument with (Fig. 25 N) or without setae; without cylindrical
projections; rays of cephalic fan with longer setae alternately disposed among
smaller setae; antenna with basal segments not overly wide (Fig. 25 J); antenna
slightly shorter than or as long as stem of cephalic fan (Fig. 25 G); ninth
abdominal segment with 1+1 ventral tubercles (Fig. 25 F); anal ring with about 75
rows and 13 hooks each ............................................................................... Lutzsimulium
– Body integument without setae; cylindrical projections present (Fig. 27 L); rays of
cephalic fan with setae arranged homogeneously, without bigger setae; antenna
with basal segments very wide (Fig. 27 I); antenna shorter than stem of cephalic
fan (Fig. 27 H); ninth abdominal segment without ventral tubercles; anal ring with
110-130 rows and about 22 hooks each. ................................................. Kempfsimulium

CLAVES DE GÉNEROS DE SIMÚLIDOS DE LA REGIÓN NEOTROPICAL


Adultos
1 Pedisulco bien desarrollado (Fig. 1 D: pe); célula basal siempre ausente; largo del
sector basal de R (medido desde la nervadura transversal humeral) menos de 1/3
de la distancia desde la base de Rs al extremo del ala .................................. Simulium
– Pedisulco ausente (Figs. 8 G, N); célula basal ausente o presente; largo del sector
basal de R aproximadamente 1/3 de la distancia mencionada o mayor ................ 2
2(1) Cu2 y A1 rectas (Fig. 20 C); célula basal ausente ......................................................... 3
– Cu2 y A1 curvadas (Fig. 8 E); célula basal ausente (Fig. 26 F) o presente (Figs. 8 E;
15 G) ..................................................................................................................................... 4
3(2) Primer flagelómero tan largo como ancho; pelos del flagelómero basal y
palpómeros más cortos que el artejo; clípeo más largo que ancho (Fig. 20 B);
piezas bucales de la hembra normales; calcipala presente (Fig. 20 E); diente
subbasal de la uña bien desarrollado (20 G) ............................................... Gigantodax
– Primer flagelómero tan largo como ancho; pelos del flagelómero basal y
palpómeros más largos que el artejo; clípeo tan largo como ancho (Fig. 20 A);
piezas bucales de la hembra, reducidas, no funcionales; calcipala ausente (Fig. 20
D); diente subbasal de la uña reducido (Fig. 20 F) ................................ Pedrowygomyia
4(2) Porción superior de membrana pleural con pelos (Fig. 10 B); segmento apical del
palpo maxilar más del doble de largo que el penúltimo ........................... Tlalocomyia
– Membrana pleural totalmente desnuda; segmento apical del palpo maxilar
generalmente menos del doble de largo que el penúltimo (Figs. 13 C; 15 C; 21 B)
................................................................................................................................................ 5
5(4) Antenas con 10 artejos; ramas del furcasternum con conspicuas proyecciones
(Fig. 23 A); genitalia del macho con esclerito geniculado, compuesto de una
porción basal en forma de Y y 2 ramas subparalelas alargadas apicalmente (Fig. 23
H); porción apical de endoparámeros con numerosos dentículos alargados;
46 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

horquilla genital casi enteramente sin pigmento, su pedúnculo muy robusto, no


más largo que los brazos (Fig. 23 E) ..................................................Paraustrosimulium
– Antenas con 11 artejos (Fig. 19 A); ramas del furcasternum sin conspicuas
proyecciones (Figs. 8 D; 13 F) [excepto Lutzsimulium (Fig. 24 L)]; esclerito
mediano del macho no como arriba, o porción apical del los endoparámeros
reducida, faltando dentículos (Figs. 8 Q; 24 I); horquilla genital bien pigmentada,
con pedúnculo no como arriba (Figs. 8 L; 10 E; 11 F; 13 M; 17 H, Q) ................ 6
6(5) R1 solamente con pelos como cerdas, sin cerdas con forma de espina (Fig. 21 F);
diente basal de la uña de la hembra, grande (Fig. 21 J) ............................ Cnesiamima
– R1 con pelos como cerdas y en forma de espinas (Figs. 13 H, I); si las cerdas en
forma de espina son poco evidentes, el diente basal de la uña de la hembra es
pequeña (Fig. 24 D) ........................................................................................................... 7
7(6) Célula basal ausente (Fig. 26 F); mandíbula de hembra dentada solo en el margen
interno (similar a figura 26 D); uñas de la hembra con un pequeño diente subbasal
(Figs. 24 D; 26 K); genitalia del macho con dientes no perceptibles en
endoparámeros (Figs. 24 I; 27 A) ................................................................................... 8
– Célula basal presente (Fig. 8 E); mandíbula de la hembra como arriba o con
dientes bien desarrollados en margen interno y externo (Fig. 13 D); uñas de la
hembra con diente basal grande (Figs. 13 G; 15 I); genitalia del macho con dientes
perceptibles en endoparámeros (Figs. 11 K; 13 T; 17 K: ed) ................................... 9
8(7) Frente más larga que ancha con surco mediano relativamente corto y sin ramas
basales bifurcadas (Fig. 24 A); Sc con alrededor de 50 tricomas (Fig. 24 H); ramas
del furcasternum con grandes proyecciones (Fig. 24 L) ..................... Lutzsimuliumm
– Frente aproximadamente tan larga como ancha con surco mediano alargado y
bifurcado, con largas ramas (Fig. 26 A); Sc sin tricomas (Fig. 26 F); ramas del
furcasternum con muy pequeñas proyecciones (Fig. 26 L) ................. Kempfsimulium
9(7) Alas con leve curvatura de Cu2; A1 casi llegando al borde del ala (Fig. 17 B);
genitalia del macho con porción basal del endoparámero poco desarrollada, y con
dentículos de la porción distal bien desarrollados, pero pocos en número,
formando un apretado grupo de disposición característica (Fig. 17 K); hembra
con ramas de la horquilla genital con 2 apodemas dirigidos hacia delante (Figs. 17
H, Q) ............................................................................................................................. Cnesia
– Alas con curvatura de Cu2 pronunciada; A1 terminando antes del margen (Figs.
8E) ; endoparámeros de machos diferentes a dilema anterior; horquilla genital sin
apodemas (Figs. 8 L; 11 F) ............................................................................................. 10
10(9) Machos ............................................................................................................................... 11
– Hembras ............................................................................................................................. 13
11(10) Esclerito mediano con una placa saliente basal transversa continuada distalmente
con una profunda incisión y bifurcada apicalmente (Fig. 8 R); endoparámeros sin
dientes (similar a Fig. 8 Q) ............................................................................ Mayacnephia
– Esclerito mediano de forma compleja curvada con la mitad distal dividida en 2
ramas anchas y cortas (Figs. 11 J, K; 13 R, S); endoparámeros con numerosos
dientes (Fig. 11 K) ............................................................................................................ 12
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 47

12(11) Genitalia con placa ventral subtriangular; ramas basales angostas y rectas (Figs. 11
I; 13 P); ramas del esclerito mediano paralelos (Fig. 13 S); endoparámeros con
porción basal grande y dientes de la porción distal bien desarrollados (Figs. 11 K;
13 T) .................................................................................................................... Araucnephia
– Genitalia con placa ventral subrectangular, sus ramas basales robustas,
redondeadas en el ápice y curvadas (Figs. 15 P, Q); ramas del esclerito mediano
notoriamente divergentes apicalmente (Fig. 15 U); endoparámeros con porción
basal pequeña y dentículos de la porción apical reducidos ............... Araucnephioides
13(10) Diente basal de la uña en forma de gancho angosto (Fig. 15 I) ...... Araucnephioides
– Diente basal de la uña más grande, subtriangular ó espatulado (Figs. 8 H, J; 13 G)
.............................................................................................................................................. 14
14(13) Frente muy angosta (Fig. 8 A), ángulo frontal de 50° aproximadamente; superficie
interna de la espermateca sin espículas (Meso América) ....................... Mayacnephia
– Frente más ancha (Fig. 15 A); ángulo frontal de 85° aproximadamente; superficie
interna de la espermateca con espículas dispersas (Sur de Suramérica) ..................
.............................................................................................................................. Araucnephia

Hembras, basadas en la genitalia


1 Ramas de la horquilla genital con largos apodemas dirigidos hacia delante (Figs.
17 H, Q; 20 I, J) ................................................................................................................. 2
– Ramas de la horquilla genital sin largos apodemas ..................................................... 3

2(1) Rama anterior de horquilla genital de ancho uniforme (Figs. 17 H, Q; 20 J.) ........
.................................................................................................................. Cnesia, Gigantodax
– Rama anterior de horquilla genital expandida en 1/3 basal (Fig. 20 I) ....................
...........................................................................................................................Pedrowygomyia
3(1) Horquilla genital casi totalmente sin pigmento, su tallo ancho y muy corto, no más
largo que las ramas (Fig. 23 E) ............................................................. Paraustrosimulium
– Horquilla genital no como arriba ................................................................................... 4
4(3) Esclerotización de la espermateca continuando en la base del ducto espermático
(Figs. 26 P, Q) ..................................................................................................................... 5
– Ducto espermático inserto en una pequeña área membranosa de la espermateca
(Figs. 8 J; 21 L) ................................................................................................................... 8
5(4) Área de inserción del ducto espermático con una peculiar estructura como una
red, extendiéndose a la base del ducto (Fig. 24 G) ..................................................... 6
– No como arriba .................................................................................................................. 7
6(5) Área esclerotizada del ducto espermático también cubriendo el cuello de la
espermateca y porción del ducto espermático con aspecto de malla (Fig. 24 G);
espermateca internamente con espículas; octavo esternito con porción
esclerotizada reducida al área mediana (Fig. 24 E) ................................. Lutzsimulium
– Área esclerotizada del ducto espermático reducida al cuello, sin aspecto de malla
en la unión con espermateca (Figs. 26 P, Q); espermateca internamente sin
espículas; octavo esternito con porción esclerotizada cubriendo todo el área desde
un costado al otro (Fig. 26 M) ................................................................... Kempfsimulium
48 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

7(5) Paraprocto muy pequeño, triangular; superficie de la espermateca conspicuamente


reticulada; su superficie interna sin espículas; esclerotización del ducto
espermático terminando abruptamente (Fig. 10 D) .................................. Tlalocomyia
– Paraprocto de tamaño normal, redondeado (Fig. 21 M); superficie de la
espermateca lisa, su superficie interna con pocas espículas; porción esclerotizada
del ducto espermático uniéndose gradualmente con la porción no pigmentada
(Fig. 21 L) ........................................................................................................... Cnesiamima
8(4) Superficie interna de la espermateca sin espículas ................................... Mayacnephia
– Superficie interna de la espermateca con espículas dispersas .................................. 9
9(8) Paraprocto aproximadamente tan largo como ancho (Fig. 13 L); superficie interna
de la espermateca no rugosa ......................................................................... Araucnephia
– Paraprocto aproximadamente la mitad más corto que ancho (Fig.. 15 L); superficie
interna de la espermateca rugosa ........................................................... Araucnephioides

Machos, basados en la genitalia


1 Dientes de endoparámeros no perceptibles; esclerito mediano comparativamente
simple, ensanchado basalmente, lamela alargada, apicalmente dividida en 2 cortas
ramas conectadas por una membrana (Figs. 24 I; 27 A) ........................................... 2
– Por lo menos una porción del endoparámero distinguible; esclerito mediano
diferente, generalmente de estructura compleja .......................................................... 3
2(1) Gonocoxito alrededor de 2 veces más largo que el gonostilo; gonostilo
suavemente aguzado distalmente; ápice relativamente romo con 3 (2) dientes
terminales (Figs. 24 I, M); placa ventral distalmente con suave curvatura
redondeada; esclerito mediano muy alargado (Fig. 24 J); pedisulco ausente ..........
............................................................................................................................ Lutzsimulium
– Gonocoxito alrededor de 3 veces más largo que el gonostilo; gonostilo
fuertemente aguzado distalmente, con 2 dientes terminales (Figs. 27 A-C); placa
ventral presentando distalmente un borde irregular (Figs. 27 A, D); esclerito
mediano ensanchado, no alargardo, distal mente aguzado; pedisulco desde ausente
a bien insinuado ............................................................................................ Kempfsimulium
3(1) Porción basal de endoparámeros no visible o apenas muy poco desarrollado;
porción denticulada compuesta de 1 o 2 grandes dientes y de varios pequeños
dentículos muy estrechamente asociados, a veces dirigidos en opuesta dirección
(Fig. 17K) ............................................................................................................................. 4
– Porción basal de endoparámeros distinta, en forma de placa; dientes de la porción
distal variando desde ausentes a bien desarrollados, pero no ordenados como fue
descrito arriba ..................................................................................................................... 6
4(3) Placa ventral subpentagonal, aproximadamente tan larga como ancha, no con
conspicuo reborde apical (Fig. 17 N) ....................................................................Cnesia
– Placa ventral en la mayoría de los casos en forma romboide alargado, con notorio
reborde apical ...................................................................................................................... 5
5(4) Placa ventral corta, con márgenes laterales en la mitad de su largo cóncavos y
margen apical bilobulado (Fig. 20 L); dientes de endoparámeros alargados ..........
............................................................................................................................... Gigantodax
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 49

– Placa ventral larga alrededor de 2 veces más larga que ancha, con márgenes
laterales en la mitad de su largo rectas y margen apical recto y convexo (Fig. 20 K);
dientes de endoparámeros cortos ..............................................................Pedrowygomyia
6(3) Esclerito mediano compuesto de una placa basal saliente transversal continuada
distalmente por una profunda incisión bifurcada apicalmente (Figs. 8 R; 10 G);
endoparámeros reducidos a una porción basal en forma de placa y sin dentículos
(Fig. 8Q) ....................................................................................... Mayacnephia, Tlalocomyia
– Esclerito mediano diferente; endoparámeros con o sin dientes .............................. 7
7(6) Esclerito mediano fuertemente curvo en vista lateral (Fig. 21 U), en vista de
conjunto con forma aproximada de M (Figs. 21 V; 23 H), con ramas internas de la
M parcialmente fusionadas y conectadas al reborde basal ........................................ 8
– Esclerito mediano no como arriba, en forma de una estructura compleja
ocupando 3 diferentes planos .......................................................................................... 9
8(7) Endoparámeros compuestos de una porción basal en forma de placa libre y
apicalmente numerosos dentículos alargados subparalelos no conectados a la
porción basal (Fig. 23 H) .......................................................................Paraustrosimulium
– Endoparámeros reducidos a una estructura subcuadrada en forma de placa;
dentículos no desarrollados (Fig. 21 U) ....................................................... Cnesiamima
9(7) Placa ventral subpentagonal o subtriangular (Fig. 11 I), con ramas basales en
forma de delgadas varillas (Fig. 13 P); ramas de esclerito mediano paralelas (Fig.
13 S); endoparámeros con conspicuos dentículos en la porción distal (Fig. 11 K)
.............................................................................................................................. Araucnephia
– Placa ventral subrectangular con ramas basales robustas apicalmente,
redondeadas (Fig. 15 P); ramas del esclerito mediano divergiendo apicalmente
(Fig. 15 U); endoparámeros con dentículos de la porción distal no desarrollados
(Fig. 15 S) ..................................................................................................... Araucnephioides

Pupas
1 Esternitos abdominales VI y VII divididos longitudinalmente a lo largo del medio
por un área estriada, membranosa (Figs. 14 D; 18 D); ganchos de los tergitos VI-
VIII invariablemente simple; segmentos abdominales VIII y IX en muchos casos
con cerdas robustas curvas, enruladas bífidas o en forma de gancho (Figs. 18 E;
23 J; 25 E) ............................................................................................................................ 2
– Esternitos VI y VII del abdomen enteros; en muchos casos con cerdas bífidas o
trífidas sobre los tergitos VI y VIII; tergitos VIII y IX sin las setas mencionadas
................................................................................................................................... Simulium
2(1) Proceso terminal del abdomen corto y puntiagudo (Fig. 23 J), o ausente (Fig. 10 L)
................................................................................................................................................ 3
– Proceso terminal del abdomen alargado (Figs. 12 E; 18 E; 25 E) .......................... 4
3(2) Capullo reducido a una pequeña capa ubicada sobre los segmentos terminales del
abdomen (Fig. 10 H); branquias en forma de un grueso tallo con unos pocos filamentos
en forma de hebras (Fig. 10 I ); abdomen fuertemente esclerotizado, tergitos y esternitos
con un número elevado, supernumerario de espinas o ganchos (Figs. 10 J, K); ápice del
abdomen romo; procesos terminales ausentes (Fig. 10 L) .............................. Tlalocomyia
50 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

– Capullo bien desarrollado, de forma definida, cubriendo todo el cuerpo de la pupa


excepto branquias, las que son dos, en forma de lámina y pseudosegmentadas (Figs.
22 C; 23 I); abdomen menos esclerotizado que el tórax; abdomen sin espinas
supernumerarias y con cortos procesos terminales (Fig. 23 J) .....................................
............................................................................................... Cnesiamima, Paraustrosimulium
4(2) Tergitos abdominales con peines de espinas (Fig. 14 D); tergito VIII sin grandes
ganchos ................................................................................................................................. 5
– Tergitos abdominales sin peines de espinas; tergito VIII con 4 grandes ganchos
........................................................................................................................................ Cnesia
5(4) Tricomas faciales (Figs. 12 B; 27 E), tricomas torácicos (Figs. 12 D; 27 F, G) y
cerdas del octavo y noveno segmento abdominal notoriamente curvado
apicalmente (Fig. 25 E) ..................................................................................................... 6
– Al menos tricomas faciales y torácicos no curvado apicalmente ............................ 7
6(5) Branquias con 12-22 ramas; frontoclípeo y tórax con tubérculos (Fig. 25 D) .......
............................................................................................................................ Lutzsimulium
– Branquias con 12 ramas; frontoclípeo y tórax sin tubérculos (Fig. 27 E) ...............
.......................................................................................................................... Kempfsimulium
7(5) Esclerito cefálico con tricomas frontal, facial, epicranial, lateral y genal en forma
de pelo (Figs. 12 B; 14 B) ............................................................................... Araucnephia
– Esclerito cefálico no con todos los tricomas mencionados; tricomas faciales con
forma de pelo o de espina ................................................................................................ 8
8(7) Tórax con tubérculos ordenados en círculos (Fig. 16 E); porción basal de las
branquias con pequeñas estructuras cuticulares en forma de placa; tricomas
torácicos en forma de pelo ...................................................................... Araucnephioides
– Tórax con tubérculos no ordenados en círculos; porción basal de las branquias sin
las estructuras mencionadas; tricomas torácicos en forma de pelo o espina ........... 9
9(8) Esclerito cefálico con 2+2 o 3+3 tricomas frontales; ramas de las branquias más o
menos tubulares (Figs. 9 A-J) ....................................................................... Mayacnephia
– Esclerito cefálico 1+1 o sin tricomas faciales; ramas de las branquias de forma
variada ................................................................................................................................. 10
10(9) Frontoclípeo sin tricomas frontal y labial (Fig. 20 N); clípeo comparativamente
angosto (Fig. 20 B) ............................................................................................ Gigantodax
– Frontoclípeo con tricomas frontal y facial (Fig. 20 M); clípeo comparativamente
ancho (Fig. 20 A) ...........................................................................................Pedrowygomyia

Larvas
1 Esclerito anal con esclerito accesorio formando un anillo completo alrededor de
la porción posterior del cuerpo (Fig. 20 S); mandíbulas con 3 dientes externos
(Fig. 20 P) ............................................................................................................................ 2
– Esclerito accesorio en forma de anillo no desarrollado (Fig. 12 N); mandíbulas
con 2 (Fig. 12 J) o 4 dientes externos (Fig. 18 J) ........................................................ 3

2(1) Hipostomio con diente lateral proyectándose sobre el nivel del primer diente
intermedio (Fig. 20 R); apotoma cefálico sin manchas basales ............... Gigantodax
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 51

– Hipostomio con primer diente intermedio generalmente proyectado por sobre el


nivel del diente lateral (Fig. 20 Q); apotoma cefálico con manchas basales (Fig. 20
O) ......................................................................................................................Pedrowygomyia
3(1) Dientes del hipostomio ordenados en 3 grupos conspicuos (Fig. 8 W);
serrulaciones del margen de la mandíbula numerosas más de 5 (Fig. 12 J) .......... 4
– Dientes del hipostomio uniformemente distribuidos, no ordenados en grupos
conspicuos (Figs. 18 H; 23 M; 25 I); serrulaciones del margen de las mandíbulas
menos numerosas, no mayor de 5, generalmente solo 2 (Fig. 25 K) ...................... 7
4(3) Antena aproximadamente tan larga como el tallo del abanico cefálico (Fig. 16 I);
hipostomio con 13 (Fig. 8 W) o 17 dientes (Fig. 16 N) ............................................. 5
– Antena más corta que el tallo del abanico cefálico (Fig. 14 H); hipostomio con 15
dientes (Fig. 14 K) ............................................................................................ Araucnephia
5(4) Escleritos cervicales muy pequeños, aislados en la membrana del cuello (Fig.16
H); mandíbulas con dientes accesorios al nivel de los dientes internos, con el
segundo diente preapical pequeño (Fig. 16 L) y con cerdas basales cortas y finas;
dentición del hipostomio corta, con 17 dientes (Fig. 16 N) ............. Araucnephioides
– Escleritos cervicales pequeños o transversales y aislados en la membrana del
cuello (Fig. 8 S), o transversales y fusionados con los bordes superiores del occiput
(Fig. 10 O); mandíbulas sin dientes accesorios al nivel de los dientes internos, con
segundo diente preapical tan largo como el primero (Fig. 10 M) o ligeramente más
corto (Fig. 8 U ) y con cortas cerdas basales llevando conspicuos dentículos
alargados (Fig. 8 V); hipostomio con 13 dientes (Fig. 8 W) ..................................... 6
6(5) Artejo proximal de la antena más de la mitad del largo del mediano (Fig. 8 T) ....
............................................................................................................................. Mayacnephia
– Artejo proximal de la antena por lo menos tan largo como el mediano ....... Tlalocomyia
7(3) Antena más larga de que el tallo del abanico cefálico; artejo distal de la antena
notoriamente más largo que el proximal y mediano juntos (Figs. 22 I; 23 L); sostenes
dirigidos hacia atrás a los costados del cuerpo principal del esclerito anal (Fig. 23 N)
............................................................................................... Cnesiamima, Paraustrosimulium
– Antena no más larga que el tallo del abanico cefálico (Fig. 27 H); artejo distal no o
solamente ligeramente más largo que el proximal y mediano juntos (Figs. 18 G; 25
J); esclerito anal sin los mencionados sostenes (Figs. 18 M; 25 L) .......................... 8
8(7) Hendidura postgenal muy superficial (Fig. 19 N); hipostomio con 17 dientes, los
4+4 intermedios similares a los restantes dientes (Fig. 19 M); mandíbula con 4
dientes externos (Fig. 18 J) ......................................................................................Cnesia
– Hendidura postgenal bien desarrollada (Fig. 25 H); hipostomio cuando más 13
dientes (Figs. 25 I; 27 J), los 1+1 laterales, cuando son perceptibles, se parecen a
las serrulaciones laterales; mandíbula con 2 dientes externos (Figs. 25 K; 27 K)
................................................................................................................................................ 9
9(8) Dientes preapicales de mandíbula subiguales en tamaño (Fig. 33 X), o decreciendo
en tamaño del primero al tercero (Fig. 38 g) .................................................. Simulium
– Dientes preapicales de la mandíbula con el segundo diente más pequeño que el
primero o el tercero (Figs. 25 K; 27 K) ...................................................................... 10
52 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

10(9) Tegumento del cuerpo con (Fig. 25 N ) o sin cerdas; sin proyecciones epidérmicas
cilíndricas; rayos de los abanicos cefálicos con dientes más largos alternativamente
dispuestos entre otros menores; antena con artejo basales no notoriamente gruesos
(Fig. 25 J); antena ligeramente más corta que el largo del abanico cefálico (Fig. 25
G); noveno segmento abdominal con 1+1 tubérculo ventral (Fig. 25 F); anillo anal
con alrededor de 75 hileras con 13 ganchos cada una ............................. Lutzsimulium
– Tegumento del cuerpo sin cerdas; presencia de proyecciones epidérmicas
cilíndricas (Fig. 27 L); rayos del abanico cefálico con dientes ordenados
homogéneamente sin dientes más largos; antena con artejos basales notoriamente
más gruesos (Fig. 27 I); antena más corta que el tallo del abanico cefálico (Fig. 27
H); noveno segmento abdominal sin tubérculos ventrales; anillo anal con 110-130
hileras de alrededor de 22 ganchos cada una ......................................... Kempfsimulium

Mayacnephia WYGODZINSKY & COSCARÓN (Figs. 8, 136.)


Mayacnephia WYGODZINSKY & COSCARÓN, 1973a: 144-148.
Type-species: Simulium pachecolunai DE LEON, 1945.
Adults: Antennae with 11 segments; male holoptic. Female with frons narrow (Fig. 8
A), frontal angle 50 degrees; mandibles and maxillae of female normally developed,
mandibles with teeth on both margins; apical segment of maxillary palp in both sexes less
than twice as long as penultimate (Fig. 8 B); sensory vesicle well developed, with abundant
tubercles (Fig. 8 C); mesepisternal sulcus wide, shallow, evanescent anteriorly; katepister-
num glabrous, in profile about as long as high; pleural membrane glabrous; branches of
furcasternum lacking projections (Fig. 8 D); metanotum glabrous; wings with basal cell
present (Fig. 8 E); R1 with hair and spine like setae (Fig. 8 F); basal section of R hairy; Rs
hairy below and hairy above on its apical half, beginning at level of junction of R1 with C;
Rs not forked; M appearing duplicated; Cu2 conspicuously curved; A2 ending well before
wing margin; calcipala present, bluntly rounded apically (Fig. 8 G); hind basitarsus of male
strongly widened (Fig. 8 N); claws of female with large, bluntly subtriangular basal tooth
(Figs. 8 H, I). Female with spermatheca normally sclerotized, with small depression on
one side, reticulate and without spicules internally, with small circular membranous area at
insertion of spermathecal duct, the latter not sclerotized (Fig. 8 J); gonapophysis large,
subtriangular, apices rounded, medial borders approximated, surface with numerous
microtrichia and a few setae (Fig. 8 K); lateral arms of genital fork with small anterior
processes and expanded distally into subrectangular plate (Fig. 8 L); stem slender, at least
as long as arms; paraproct short with rounded rim (Fig. 8 M); male with gonostylus about
as long as gonocoxite, conical, not flattened with two apical spines (Fig. 8 O); ventral plate
somewhat wider than long, subsemicircular, setose (Fig. 8 P). Endoparameres reduced to
plate like sclerite; denticles absent or only weak (Fig. 8 Q); median sclerite with transverse
basal flange, deeply narrow incision and bifurcated apically (Fig. 8 R).
Pupa: Cocoon irregularly shaped, covering part or all of body; body of pupa
sclerotized throughout; head, and thorax with pterothecae, and abdominal segments
tuberculate; gills with 2-15 thickened, tubular, occasionally divided (or basally fused)
branches, arising from short common base, all with superficial but mostly conspicuous
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 53

bc

C
Sc
C Rs R1
A F
B
H
G

D
N

I
M

K
L

T
J O
P

R
V
U
Q
W X S
Figure 8. A-Q, S-X: Mayacnephia aguirrei; A-M: Female; A: frons; B: maxillary palp; C: sensory vesicle
of maxillary palp with high magnification; D: furcasternum; E: wing, general view (bc = basal cell); F:
portion of anterior wing veins, showing pilosity (C = costa, R1= radius 1, Rs = secondary radius
(radius 2), Sc = subcosta); G: calcipala and tarsomere I; H-I: claw with large basal tooth; J:
spermatheca; K: eighth sternite and gonapophyses; L: genital fork; M: cercus and paraproct; N-Q:
Male; N: calcipala and tarsomere I; O: gonocoxite and gonostylus; P: ventral plate, with portion of
median sclerite; Q: endoparameres with portion of median sclerite and aedeagus membrane; R:
median sclerite positioned on ventral plate; S-X: Larva; S: cervical sclerite; T: antenna; U: teeth of
mandible; V: short basal setae of mandible; W: teeth of hypostomium; X: lateral sclerite of proleg. R:
M. roblesi; ventral plate and median sclerite. (A-X from WYGODZINSKY & COSCARÓN, 1973a).
54 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

transverse annulations with spiculae; head with 2+2 or 3+3 frontal and 1+1 facial trichomes;
all hair like. Thorax with approximately five simple trichomes; abdomen pointed posteriorly.
Distinct pleural plate absent; abdominal segments without perceptible transversal ridges;
terga III and IV with 4+4 or 5+5 hooks posteriorly, in some cases 5+5 hooks also on tergum
II, and 1+1 hooks on V. Terga V-IX, or VI-IX, with continuous spine-combs anteriorly; faint
spine-combs also perceptible on III and IV; sternum IV with 1+1 strong setae or hooks;
sternum V with from 2+2 to 4+4 hooks; sternal plates VI and VII either each with 1+1
hooks accompanied by no or 1+1 hooks in pleural membrane, or sternal plates VI with 3+3
and sternal plates VII with 2+2 hooks, accompanied by 1+1, 2+2 or 3+3 hooks in pleural
membrane; all hooks simple. Terminal hooks well developed, relatively stout, not sinuous;
posterior segments with few irregularly formed hook- or grapnel-shaped setae.
Larva: Head spots positive; cephalic apotome broadest well before hind margin;
cervical sclerites transverse, small or large, either free or fused to upper ends of postocciput
(Fig. 8 S); postgenal cleft very shallow; antennae as long as stern of cephalic fans; third
segment slightly darker than remaining. First antennal segment over half as long as second
(Fig. 8 T); cephalic fans normal; mandibles with two outer and one apical tooth; third
preapical tooth largest, but not so large as apical tooth; inner teeth arranged in single row;
mandibular serrations very numerous (15-22) (Fig. 8 U); short basal setae stout, with
conspicuous denticles along almost their entire length (Fig. 8 V); hypostomium with 13 teeth
arranged in three conspicuous groups, with median tooth and lateral teeth by far largest; 2+2
lateral teeth situated only slightly below level of remaining teeth (Fig. 8 W); lateral sclerite of
thoracic proleg well developed, about as long as wide, subrectangular, lower border
subparallel to longitudinal axis of proleg; setae of lateral sclerite less than half as long as
sclerite, arranged in irregular double row, this row extending slightly beyond sclerite below
(Fig. 8 X); larval cuticle bare, except simple setae at base of anal sclerite, and few perianal
scales; anal sclerite normal, X-shaped. Terminal ventral tubercles absent or present.
Distribution: Canada: Alberta, British Columbia, Yukon; Highlands of Mexico to
Panama; USA: Arizona, California, Idaho, Oregon, Washington (Fig. 136).
Discussion: This genus comprises 15 species, 11 of which are found in the
Neotropical Region south of the United States. Mayacnephia is most closely related to
Tlalocomyia WYGODZINSKY & DÍAZ NÁJERA; the two genera form a monophyletic group,
Tlalocomyia being the more apomorphic component. ADLER et al. (2004) synonymized
Mayacnephia with Tlalocomyia. Here they are maintained separate. One Neotropical species,
Mayacnephia alticola (DYAR & SHANON), is recorded from Mexico, but we could not obtain
more information and do not consider it to be a recognizable species.

KEY TO MAYACNEPHIA SPECIES

We include 10 species from Mexico to Panama considered as Mayacnephia by the authors.


We present only a key to pupae, which have good characters, especially in the shape of the
respiratory organs that help to identify species; our key is in part based on that of RAMÍREZ
PÉREZ et al. (1988), and includes the known Mayacnephia species at this time.
We don’t include keys to females, males or larvae because of the lack of enough
specimens; based on the available descriptions, these have very similar characters. Females
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 55

are unknown in Mayacnephia atzopensis and M. mixensis, and males, in addition to these two
species, also are unknown in Mayacnephia muzquicensis and M. tadai. DALMAT (1955) gave keys
to larva, pupae, males and females of three species: Mayacnephia aguirrei, M. pachecolunai and
M. roblesi. DÍAZ NÁJERA (1962) made a larval key to five species, including the three species
presented by DALMAT, as well as Mayacnephia atzopensis and M. mixensis. The characters used
by DÍAZ NÁJERA show great variation and do not offer security in identification.
Only distinctive characters are considered in the species descriptions. Those
characters not mentioned are in accordance with the generic description.

Key to Mayacnephia pupae


1 Gill with 2 tubular and relatively swollen and elongated branches (Fig. 9 A) ........
................................................................................................................ Mayacnephia aguirrei
– Gill with variable shape, 3-15 branches ......................................................................... 2
2(1) Gill with 3-7 branches ....................................................................................................... 3
– Gill with 11-15 branches .................................................................................................. 8
3(2) Gill with 3 branches (Fig. 9 E), (based on Fig. 20 of DÍAZ NÁJERA, 1962) .............
...............................................................................................................Mayacnephia mixensis
– Gill with 4-7 branches ....................................................................................................... 4
4(3) Gill with 4 branches .......................................................................................................... 5
– Gill with 6-7 branches ....................................................................................................... 6
5(4) Gill tubular, subannulated, apically acuminate (Fig. 9 C) ........... Mayacnephia grenieri
– Gill swollen, smooth, apically rounded (Fig. 9 I) (based on Fig. 2 of RAMÍREZ
PÉREZ et al., 1988) ................................................................................. Mayacnephia salasi
6(4) Gill with 6 smooth tubular branches with short stalks, subequal in length and
thickness (Fig. 9 J), (based on Fig. 6 of OCHOA & TAKAOKA, 1993) .........................
.................................................................................................................... Mayacnephia tadai
– Gill with 7 terminal branches .......................................................................................... 7
7(6) Gill with terminal filamentous branches originating each in number of 1 or 2 on 4
thick tubular subannulated stalks (Fig. 9 H) ................................... Mayacnephia roblesi
– Gill with terminal clavate branches originating on 4 relatively thin tubular and
smooth stalks (sometimes with thumb-like process on anterior internal branch)
(Fig. 9 D), (based on Fig. 8 of PETERSEN, 1985) ..................... Mayacnephia fortunensis
8(2) Gill with 11 terminal filaments of different lengths, with thick base and gradually
becoming thin to apex, (Fig. 9 B) (based on Fig. 9 of DÍAZ NÁJERA, 1962) ...........
............................................................................................................ Mayacnephia atzopensis
– Gill with 14 to 15 filaments ............................................................................................. 9
9(8) Gill with 12 or 13 elongated filamentous branches and 1 medial internal short,
horizontally disposed (Fig. 9 G) ............................................... Mayacnephia pachecolunai
– Gill with 14 or 15 arborescent filamentous branches emerging from 4 thick stalks
(Fig. 9 F), (based on Fig. 18 of DÍAZ NÁJERA, 1971 ........... Mayacnephia muzquicensis

Clave de especies de Mayacnephia basada en pupas


1 Branquias tubulares y con 2 ramas relativamente infladas y alargadas (Fig. 9 A)
................................................................................................................ Mayacnephia aguirrei
56 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

– Branquias con 3-15 ramas ................................................................................................ 2


2(1) Branquias con 3-7 ramas .................................................................................................. 3
– Branquias con 11-15 ramas .............................................................................................. 8
3(2) Branquias con 3 ramas (Fig. 9 E) (basada en figura 20 de DÍAZ NÁJERA, 1962) ....
...............................................................................................................Mayacnephia mixensis
– Branquias con 4-7 ramas .................................................................................................. 4
4(3) Branquia con 4 ramas ........................................................................................................ 5
– Branquia con 6-7 ramas .................................................................................................... 6
5(4) Branquias tubulares, subanilladas, apicalmente aguzadas (Fig. 9 C) ..........................
................................................................................................................. Mayacnephia grenieri
– Branquias infladas, lisas, apicalmente redondeadas (Fig. 9 I) (basada en figura 2 de
RAMÍREZ PÉREZ et al., 1988) ................................................................. Mayacnephia salasi
6(4) Branquias con 6 ramas tubulares con corto tallo, casi iguales en longitud y grosor,
y con superficie lisa (Fig. 9 J) (basado en OCHOA & TAKAOKA, 1993) ......................
.................................................................................................................... Mayacnephia tadai
– Branquias con 7 ramas terminales .................................................................................. 7
7(6) Branquias con ramas terminales filamentosas originadas cada una en número de 1
o 2 sobre 4 tallos tubulares gruesos subanillados (Fig. 9 H) ...... Mayacnephia roblesi
– Branquias con ramas terminales en forma de clava, originadas sobre 4 tallos
relativamente finos, tubulares y lisos (Fig. 9 D) (basado en figura 8 de PETERSEN,
1985) ................................................................................................. Mayacnephia fortunensis
8(2) Branquias con 11 filamentos terminales de diferente longitud con gruesa base y
afinándose suavemente hacia el ápice (Fig. 9 B) (basado en figura 9 de DÍAZ
NÁJERA, 1962) .................................................................................. Mayacnephia atzopensis
– Branquia con 14-15 filamentos terminales ................................................................... 9
9(8) Branquia con 12-13 ramas filamentosas, alargadas y una mediana interna corta,
dispuesta horizontalmente (Fig. 9 G) ..................................... Mayacnephia pachecolunai
– Branquia con 14-15 ramas filamentosas, arborescentes, emergiendo de 4 tallos
gruesos (Fig. 9 F) (basado en DÍAZ NÁJERA, 1971) ............. Mayacnephia muzquicensis

Mayacnephia aguirrei (DALMAT) (Figs. 8 A-Q, S-W; 9 A.)


Simulium aguirrei DALMAT, 1949: 544.
Cnephia aguirrei DALMAT, 1951: 48; VARGAS & DÍAZ NÁJERA, 1952: 53; DALMAT, 1955: 89; VARGAS &
DÍAZ NÁJERA, 1957b: 268; DÍAZ NÁJERA, 1962: 273.
Mayacnephia aguirrei WYGODZINSKY & COSCARÓN, 1973a: 149-150; CROSSKEY & HOWARD, 1997: 16; 2004: 7.
(Based on DALMAT, 1955: 89).
Female: Wing length 3.4 mm; frons, clypeus and palpi blackish; scutum velvety black
with gray pruinosity, with appressed silvery and few yellow hairs, metanotum velvety dark
brown; legs mostly dark brown; abdominal tergites dark brown, with silvery white hair,
sternites yellowish brown; frons narrow (Fig. 8 A); maxillary palp as in figure 8 B, with
sensory vesicle (Fig. 8 C) about 1/3 of basal segment length; furcasternum as in figure 8
D; wing as in figure 8 E, with basal cell well developed; Sc completely pilose; hair and
spinelike setae of C, R1 and Rs as in figure 8 F; calcipala as in figure 8 G; claw subbasal
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 57

tooth elongate (Fig. 8 H-I); spermatheca as in figure 8 J, spermathecal duct ending in a


membranous area; eighth sternite with about 19 trichomes (Fig. 8 K); genital fork as in
figure 8 L; paraproct subovoidal distally (Fig. 8 M).
Male: Wing length 3.0 mm; clypeus and palpi blackish; scutum velvety black with
short black pilosity; metanotum velvety black; legs dark brown; abdominal segments I-II
blackish, others velvety dark brown to black, with posterior margin brown to white
pruinose; Sc pilose on basal third; length/width ratio of hind basitarsus III = 3.1;
gonocoxite and gonostylus as in figure 8 O; ventral plate, endoparameres and median
sclerite as in figures 8 P, Q, R.
Pupa: Cocoon length at base 2.0 mm covering one half of abdomen, weave composed
of loose threads; thorax and abdomen with abundant granulosity; gills composed of 2 long,
tubular, subannulated branches tapered at apex (Fig. 9 A); maximum length 2.5 mm.
Larva: Total length 6.3 mm; general color grayish, with very long dark patches on
segments 6 and 7; cephalic fan with 30-35 rays; cervical sclerite as in figure 8 S; antenna as in
figure 8 T; ratio of proximal, medial and distal segments of antenna = 1:1.2:1.7; mandible
with 23 marginal teeth (Fig. 8 U), basal setae of mandible as in figure 8 V; hypostomium with
13 apical teeth (Fig. 8 W), 3 hairs on each side and laterally with 6 serrations; anal ring with
72-75 rows of hooks; area between anal sclerite arms with few bifid and trifid scales; eighth
sternite with 2 tubercles; thoracic proleg sclerite with about 28 setae (Fig. 8 X).
Distribution: Highlands of Guatemala; Mexico: Chiapas.
Bionomics: DALMAT (1955) reported that this species was found between 900 and
1825 m, most frequently above 1525 m. The aquatic instars occurred in streams with a
temperatures ranging between 14 °C and 23 °C, with most specimens at 19 °C and below.
None of the streams where this species breeds was wider than 1.5 m, and almost all were
no deeper than 30 cm. The preferred current was between 2.5 and 25 cm per second, and
never over 50 cm per second. Pupae preferred stones or rocks as a substrate. Less than 10
percent were found on leaves and twigs.

Mayacnephia atzopensis (DÍAZ NÁJERA) (Fig. 9 B.)


Cnephia atzopensis DÍAZ NÁJERA, 1962: 273.
Mayacnephia atzopensis WYGODZINSKY & COSCARÓN, 1973a: 150.
(Based on original description).
Female, Male and Pupa: Unknown.
Larva: Body length 9.0 mm; general coloration yellowish; cephalic fan with 25 rays;
mandible with marginal serrations of about 21 teeth; hypostomium with 13 teeth, 6 or 7
lateral setae on each side; ratio of proximal, medial and distal segments of antenna =
1:1.3:2.1; pupal gill histoblast with 11 tubular branches of irregular length, with two pairs on
each side with well-developed stalk (Fig. 9 B); ninth abdominal segment without ventral
tubercles; anal ring with 78-80 rows of hooks; area of anal sclerite with scarce thin spines.
Distribution: Mexico: Morelos.
Bionomics: Larvae live in small creeks, in shaded and cold water.
Discussion: The peculiar shape of the gill histoblast allows this species to be
differentiated from other species.
58 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

C
A
B

E F

H
G

I J

Figure 9. Mayacnephia. A-J: Gills; A: M. aguirrei; B: M. atzopensis; C: M. grenieri; D: M. fortunensis E: M.


mixensis; F: M. muzquicensis; G: M. pachecolunai; H: M. roblesi; I: M. salasi; J: M. tadai.

Mayacnephia fortunensis PETERSEN (Fig. 9 D.)


Mayacnephia fortunensis PETERSEN, 1985: 80; CROSSKEY & HOWARD, 1997: 16; 2004: 7.
(Based on original description).
Female: Wing length 3.2 mm; antennae, frons and clypeus brown, palp dark brown,
scutum dark brown densely covered with yellowish hair, legs brown, with tarsi light
brown; Sc sparsely pilose.
Male: Wing length 2.5 mm; general color dark brown; clypeus brown with silver
pruinosity and pale yellowish to brown hair; antenna light brown with yellowish base; palp
dark brown; legs as in female; abdominal tergites dark brown with distal border light brown.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 59

Pupa: Cocoon roughly pocket-shaped, loosely woven; gill with 7 thick tubular
branches emerging from 4 primary branches (Fig. 9 D), sometimes with small internal
protuberance like a short process on frontal internal branch; head and thorax with small
tubercles in irregular distribution.
Larva: Body length about 7.5 mm; general color dull cream yellow mottled from
gray green to dark brown; hypostomium with 13 teeth; mandible with marginal serrations
of about 22 teeth.
Distribution: Panama: Chiriqui.
Bionomics: Larvae live in small streams less than 1 m wide, on roots, leaves or rocks,
in shaded areas at water temperatures of 14-20 °C.

Mayacnephia grenieri (VARGAS & DÍAZ NÁJERA) (Fig. 9 C.)


Cnephia grenieri VARGAS & DÍAZ NÁJERA, 1948b: 325; 1957b: 270; DÍAZ NÁJERA, 1962: 273.
Mayacnephia grenieri WYGODZINSKY & COSCARÓN, 1973a: 149; CROSSKEY & HOWARD, 1997: 16.
(Based on original description).
Female: Wing length 3.5 mm; general coloration blackish; frons black with golden
pilosity, clypeus and palp black; scutum black with light brown pilosity, pleura black, legs
blackish with black hair and tarsi whitish; abdomen black, tergites laterally with silvery
pilosity; Sc with hair on entire length.
Male: Wing length 3.0 mm; general coloration as in female; scutum uniformly black;
abdomen whitish with silver pilosity; tergites blackish elongated; sternites also with
blackish coloration; Sc without hair.
Pupa: Cocoon length 3.0 mm, conical in shape, covering only abdomen; body
integument granulose; gill with four tubular elongated branches disposed in two pairs with
primary branches very short and emerging from small stalk (Fig. 9 C).
Larva: Total length 8.0-9.0 mm; general coloration light yellowish and darkened
distally on ventral side; antenna as long as stem of cephalic fan; hypostomium shape
typical for genus but with 9 teeth; 2 or 3 lateral setae on each side, and marginal serrations
with 5 small teeth; anal ring with 76 rows of hooks.
Distribution: Mexico: Veracruz.
Bionomics: The aquatic instars of this species were found at an altitude of 1500 m,
in a small clear stream 60 cm below its origin from a spring.

Mayacnephia mixensis (DÍAZ NÁJERA) (Fig. 9 E.)


Cnephia mixensis DÍAZ NÁJERA, 1962: 284.
Mayacnephia mixensis WYGODZINSKY & COSCARÓN, 1973a: 150; CROSSKEY & HOWARD, 1997: 16; 2004: 7.
(Based on original description).
Female, Male and Pupa: Unknown.
Larva: Length 8.0 mm; general color yellowish; cephalic fan with 28 rays; mandible
with marginal serrations of about 16 teeth; hypostomium with 15 teeth, 3 or 4 lateral setae
on each side; ratio of proximal, medial and distal segments of antenna = 1:1.5:2.2; pupal
gill histoblast with 3 thick tubular branches emerging from same place on very short stalk
60 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

(Fig. 9 E); ninth sternite without ventral tubercles; anal ring with 94 rows of hooks; area
of anal sclerite without scales or spines.
Distribution: Mexico: Oaxaca.
Bionomics: Larval habitats unknown.
Discussion: The peculiar pupal gill shape with 3 branches allows us to separate
Mayacnephia mixensis from all the known Mayacnephia species.

Mayacnephia muzquicensis (DÍAZ NÁJERA) (Fig. 9 F.)


Cnephia (Cnephia) muzquicensis DÍAZ NÁJERA, 1971: 239.
Mayacnephia muzquicensis PY-DANIEL, 1982: 311; CROSSKEY & HOWARD, 1997: 16; 2004: 7.
(Based on DÍAZ NÁJERA, 1971).
Female: Wing length 4.1 mm; general coloration blackish; clypeus yellowish; palp
blackish; scutum black with coppery shine and golden pilosity; metanotum reddish; pleura
blackish; abdomen blackish; Sc mostly haired.
Male: Unknown.
Pupa: Cocoon length 4.0 mm, weave soft, without definite shape; thorax with scarce
granulosity; gill arborescent with 14 or 15 filamentous branches emerging from 7 bifid
primary branches from thick stalk (Fig. 9 F).
Larva: Body length 8.5 mm; color darkened dorsally and whitish ventrally; head
black; cephalic fan with 27 rays; mandible with about 14 marginal teeth; ratio of proximal,
medial and distal segments of antennae = 1:2.5:2.4; hypostomium with 13 teeth; 5 or 6
lateral setae on each side; without scales; ninth segment without ventral tubercles.
Distribution: Mexico: Coahuila.
Bionomics: Larvae live on stones in small creeks of clear and cold water, without
vegetation. Feeding habits of the females are unknown.

Mayacnephia pachecolunai (DE LEÓN) (Fig. 9 G.)


Simulium pachecolunai DE LEÓN, 1945: 67.
Cnephia pachecolunai DALMAT, 1951: 48; VARGAS & DÍAZ NÁJERA, 1954: 67; DALMAT, 1955: 43; DÍAZ
NÁJERA, 1962: 273.
Mayacnephia pachecolunai WYGODZINSKY & COSCARÓN, 1973a: 148; CROSSKEY & HOWARD, 1997: 16;
2004: 7.
(Based on DALMAT, 1955).
Female: Wing length 4.1 mm; frons, clypeus, palpi and scutum blackish with gray
pruinosity; pleura dark brown, legs brown and abdomen blackish with posterior margin
brown; Sc pilose along basal three fourths.
Male: Wing length 3.6 mm; clypeus dark brown, palpi black, scutum velvety black
with long yellow and black hair, pleura dark brown to black, legs dark brown to black on
tarsi; abdomen velvety brown to black with posterior margin white-pruinose with black
hair; length/width ratio of hind basitarsus = 3.1-3.4.
Pupa: Cocoon length at base 4.0 mm, weave very soft without regular shape and
appearing to cling to pupa, covering abdomen and one fourth of thorax; thorax and abdomen
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 61

covered by tubercles; gills with 12-13 filamentous branches with median interior shorter;
branches flattened in one plane arising a little behind anterior margin of thorax (Fig. 9 G).
Larva: Length 5.9 mm; general color yellowish brown; cephalic fan with 25
branches; mandible with 25 marginal teeth; hypostomium with 13 teeth and 3 or 4 setae,
lateral margins with 8 serrations; anal ring with 80-81 rows of hooks; ninth abdominal
segment with 2 ventral tubercles; area between anal sclerite arms without scales.
Distribution: Guatemala: Solola and Totonicapan; Mexico: Chiapas.
Bionomics: The following data are based on DALMAT (1955). One adult was found
to bite humans, but the real hosts of this species are not known. Normally, Mayacnephia
pachecolunai occurs only above 2100 m and breeds in cold streams of not more than 17 °C,
although the concentration of specimens is greatest at water temperatures below 11 °C.
The breeding grounds were “almost entirely minute trickles of water that pass over a
swampy area supporting large quantities of vegetation; such rivulets usually dried up
during the months of November through March.” These rivulets were most frequently
less than 15 cm wide and 13 cm or less deep; the current was generally very slow, less than
38 cm per second. About half of the pupae collected were attached to rocks, and the
remainder to leaves or some few to branches or twigs.

Mayacnephia roblesi (DE LEÓN) (Figs. 8 R, 9 H.)


Simulium roblesi DE LEÓN, 1943: 97; 1945: 67.
Cnephia roblesi DALMAT, 1951: 48; VARGAS & DÍAZ NÁJERA, 1954: 64; DALMAT, 1955: 97.
Mayacnephia roblesi WYGODZINSKY & COSCARÓN, 1973a: 149; CROSSKEY & HOWARD, 1997: 16; 2004: 7.
Female: Wing length 4.3 mm; frons and clypeus dark brown with gray pruinosity,
scutum black with golden colored hair, with 3 longitudinal stripes of gray pruinosity;
pleura brown with gray pruinosity, legs dark brown with femur a little lighter; abdomen
dark brown with gray pruinosity, tergites with black or brown hair; Sc completely pilose.
Male: Wing length 4.0 mm; palpi and clypeus black; scutum velvety black with
appressed golden hair; pleura brown with gray pruinosity; legs mostly dark brown;
abdominal tergites velvety dark brown to black; sternites yellowish brown. Ventral plate
and median sclerite as figure 8 R.
Pupa: Cocoon length of base 5.3 mm; loosely woven, cocoon covering abdomen
and one fourth of thorax; thorax and abdomen without granulosity; gill with seven
filamentous terminal branches emerging from 4 inflated saclike, subannulated basal
branches, 3 with 2 terminal branches and the frontal external with only one (Fig. 9 H).
Larva: Length 8.3 mm; general color yellowish brown; cephalic fan with 24 rays;
mandible with 29-32 marginal teeth; ratio of proximal, medial and distal segments of
antennae = 1:1.1:1.5; hypostomium with 13 apical teeth and 15 lateral serrations; anal ring
with 95-100 rows of hooks; area between anal sclerite arms with scarce bifid and trifid
scales; ninth sternite with 2 ventral tubercles.
Distribution: Highlands of Guatemala.
Bionomics: The feeding habits of the adults are not known. This is a highland
species breeding at altitudes above 2100 m. The aquatic instars were found in streams with
a water temperature below 13 °C, not infrequently between 10 °C and 11 °C. Mayacnephia
62 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

roblesi breeds in the same types of small rivulets in swampy areas as M. pachecolunai (see
above), and is restricted to rivulets of less than 15 cm wide, with depths of generally less
than 3 cm; the current is slow, with speeds of 25 cm per second, or less. Contrary to the
situation for M. pachecolunai, the pupae of M. roblesi are never found on rocks or stones;
they are in most cases attached to leaves, with only a few found on branches or twigs.

Mayacnephia salasi RAMÍREZ PÉREZ, PETERSON & VARGAS (Fig. 9 I.)


Mayacnephia salasi RAMÍREZ PÉREZ et al., 1988: 66; CROSSKEY & HOWARD, 1997: 16; 2004: 7.
(Based on original description).
Female: Wing length 3.4 mm; antenna and palp dark brown; scutum dark brown to
blackish with lateral margins paler with grayish polinosity, covered by yellowish pilosity;
pleura dark brown mottled with some darker areas; legs dark brown; abdomen yellowish
brown.
Male: Wing length 3.1-3.4 mm; general color velvety dark brown.
Pupa: Cocoon loosely woven; gill with four cylindrical, globose branches originating
from a common short base (Fig. 9 I); head and thorax without granulosity.
Larva: Length 6.5-8.5 mm; color pale creamy brown; ratio of proximal, medial and
distal segments of antenna = 1:7.7:2.6; labral fan with 25-33 rays; hypostoma with 13
teeth; 3-6 lateral serrations; mandible with 16-25 marginal serrations; anal ring with 62-65
rows and 8-10 hooks each; ninth segment with 2 ventral tubercles; area of anal sclerite
with 8-10 thin setae.
Distribution: Costa Rica: San José.
Bionomics: Mayacnephia salasi lives in small temporary streams at an elevation of
3150 m in an area heavily shaded by forest, and with temperatures of 10-14 °C and a pH
of 7.1-7.8.

Mayacnephia tadai OCHOA & TAKAOKA (Fig. 9 J.)


Mayacnephia tadai OCHOA & TAKAOKA, 1993: 5; CROSSKEY & HOWARD, 1997: 16; 2004: 7.
(Based on original description).
Female: Frons, clypeus and scutum dark brown with whitish pollinosity; wing and
legs lost; abdomen dorsally dark brown.
Male: Unknown.
Pupa: Cocoon small, short, slipper shaped, loosely woven, covering only abdomen;
body length 4.0 mm; integument moderately covered by minute tubercles; gill with 6
tubular filaments with tapered apex arranged in three pairs with a short stalk (Fig. 9 J).
Larva: Length 6.5 mm; color pale yellow; antenna as long as stem of cephalic fan;
ratio of proximal, medial and distal segments = 1:0.85:1.1; cephalic fan with 36 rays;
mandible with about 22 marginal serrations; hypostomium with 13 teeth, and 4 lateral
setae; ninth segment with 2 ventral tubercles; anal ring with 76 rows of 18 hooks each.
Distribution: Guatemala: Esquintla.
Bionomics: Larvae live on twigs in small temporary streams, at 1000 m of altitude,
together with M. aguirrei. Feeding habits of the female are unknown.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 63

Tlalocomyia WYGODZINSKY & DÍAZ NÁJERA (Figs. 10, 136.)


Tlalocomyia WYGODZINSKY & DÍAZ NÁJERA, 1970: 83-108.
Type-species: Tlalocomyia revelata WYGODZINSKY & DÍAZ NÁJERA, 1970: 88.
Mayacnephia WYGODZINSKY & COSCARÓN, 1973a: 144-150; ADLER et al., 2004: 290 (synonym).
Adults: Frons very narrow; frontal angle 30 degrees; apical segment of maxillary
palp more than twice as long as penultimate; pleural membrane hairy on upper third (Fig.
B); claw with subbasal tooth subovoidal spermatheca reticulate on outer surface, not
spiculate on inner surface; membranous area at insertion of spermathecal duct with
evident sclerotization, the duct sclerotized at base and this sclerotization terminating
abruptly (Fig. D); genital fork with median and lateral branches well sclerotized (Fig. E);
paraproct relatively short, subtriangular; ventral plate subovoidal distally; endoparameres
with weak basal plates, no spines; median sclerite with posterior portion divided in two
short wide and divergent branches (Fig. G).
Pupa: Cocoon obsolescent, composed of closely woven basal cushion and irregular
threads loosely covering, at most, posterior half of abdomen of pupa (Fig. H); body of
pupa very heavily sclerotized throughout; gills composed of elongate, basally thickened
trunk from which arise sequentially a few threadlike filaments (Fig. I); filaments smooth;
facial trichomes spinelike; frontoclypeal tubercles large, acute and few in number;
abdomen rounded-truncate posteriorly; spine-combs of abdominal terga absent (Fig. J);
abdominal terga and sterna each with median transverse ridge; ridges on terga II-VIII and
sterna III-V each with one row of very numerous simple spines or hooks; sternal plates of
segments VI-VII with several irregularly arranged hooks (Fig. K); terminal hooks not
developed; pleural membranes of posterior segments with 4-5 additional spines or hooks
per segment (Fig. L); all hooks simple.
Larva: Head with normal shape; cephalic apotome well pigmented before posterior
margin; cervical sclerites large, fused to terminal portion of postocciput (Fig. O); antenna
as long as stem of cephalic fan, third preapical teeth longer than others but shorter than
apical; mandible with abundant marginal teeth (about 20) (Fig. M); hypostomium with 13
teeth arranged in three groups (Fig. N); postgenal cleft not deep; lateral sclerite of proleg
well developed, subtriangular; body cuticle bare except scarce perianal scales; anal sclerite
X shaped; rectal papillae trilobate without additional diverticula.
Distribution: Mexico (Fig. 136).
Discussion: Tlalocomyia is very close to Mayacnephia. They share a combination of
characters such as the loosely woven cocoon covering only a part of the body (generally
abdomen), reticulated and one flattened side of the spermatheca, deeply incised median
sclerite of the male genitalia and arrangement of the hypostomial teeth, confirming a
monophyletic group. ADLER et al. (2004) synonymized Mayacnephia with Tlalocomyia and
subdivided Tlalocomyia into three species groups: PACHECOLUNAI which comprises all
Neotropical species including Tlalocomyia revelata and T. andersoni, T. osborni, T. ramifera and
T. stewarti that are from North America.
The following characters of Tlalocomyia help separate it from Mayacnephia: accentuat-
ed differences between sexes, viz. sensory vesicle of the maxillary palp which is very big in
females but small in males; scutum bright orange-yellow in the female, black in the male;
64 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

maxillary palp of last segment twice as long as the second; anepisternal membrane with
hairs on the upper area; spermatheca strongly reticulate; spermathecal duct sclerotized at
base, terminating abruptly; median sclerite with wide incision and thick branches;
subovoidal tooth of basal claw; pupal gill composed of a thick tube from which several
thick short filaments emerge without spiculae on the annulations; facial spine-like

B
A
C

G
D
F

I
H

J K L

N O

Figure 10. Tlalocomyia revelata. A-E: Female; A: sensory organ of maxillary palp; B: thorax, lateral
view; C: claw; D: spermatheca. E: genital fork; F-G: Male; F: hind basitarsus and tarsomere I; G:
median sclerite. H-L: Pupa; H: general view; I: gill; J: chaetotaxy of abdomen, dorsal view; K: idem,
ventral view; L: idem of apex, lateral view. M-O: Larva; M: teeth of mandible; N: teeth of
hypostomium; O: cervical sclerites (A-O from WYGODZINSKY & COSCARÓN, 1973a).
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 65

trichomes; abdominal segments II-VIII and ventral segments III-VIII with abundant
hooks or robust spines (described in detail by PY-DANIEL & MOREIRA SAMPAIO, 1994: 139);
no terminal hooks; larva with proximal antennal segment not over half as long as medial,
and scale fan not evident.

Tlalocomyia revelata WYGODZINSKY & DÍAZ NÁJERA (Fig. 10.)


Tlalocomyia revelata WYGODZINSKY & DÍAZ NÁJERA, 1970: 88; WYGODZINSKY & COSCARÓN, 1973a:
151; CROSSKEY & HOWARD, 1997: 24; 2004: 7.
Female: Wing length 4.5-5.0 mm; general color yellowish-orange; frons and clypeus
brownish to orange; palpi brownish; scutum, scutellum, metanotum, pleurae and legs
yellowish orange; scutum with golden pilosity; fronto-ocular triangle longer than wide;
cibarium smooth; lacinia with 42-46 teeth; last palpal segment longer than the two before;
palpal sensory organ as long as the diameter of the segment with abundant sensory vesicles
(Fig. A); wings with well-developed basal cell; Sc with basal sector of R hairy; length/width
ratio of hind basitarsus about 7; claw with wide subovoidal subbasal tooth (Fig. C).
Male: General coloration blackish; scutum black with golden appressed hair, pleura
black; abdomen with tergites blackish and posterior border lighter; hind basitarsus wide (Fig.
F); length/width ratio about 3; median sclerite deeply bifurcated with wide branches (Fig. G).
Pupa: Body length 4.2-4.5 mm; general aspect as in figure H; tubercles rounded on
head, acuminate on thorax; maximum gill length 4.0 mm; gill with 5 branches (Fig. I);
trichomes simple, with 3+3 frontal and 1+1 facial, thicker than others; thoracic trichomes
simple; number of spines on abdominal tergites as follows: II: 16-17; III: 17-18; IV: 18; V:
14; VI: 12; VII: 10-11 and VIII: 4.
Larva: Maximum length 9.0-9.5 mm; general color whitish; head dark brown; ratio
of proximal, medial and distal antennal segments = 1:1.9-2.3:1.8-2.1; cephalic fan with 30-
34 rays; mandible with 20-24 teeth; hypostomium with 13 teeth (Fig. N); 8-11 lateral setae,
irregularly disposed; anal sclerite with perianal scales; anal ring with 72-74 rows with 14-17
hooks each.
Distribution: Mexico: Distrito Federal.
Bionomics: Larvae live in small temporary creeks, attached to stones and vegetation.

SPECIES INCERTAE SEDIS


?Tlalocomyia alticola (DYAR & SHANNON), 1927
Eusimulium alticolum DYAR & SHANNON, 1927: 24; VARGAS, 1945b: 110; VARGAS & DÍAZ NÁJERA,
1957b: 230.
Mayacnephia alticola CROSSKEY & HOWARD, 1997: 16; 2004: 7.
Tlalocomyia alticola ADLER et al., 2004: 290.
According to the original female description, based especially on the genital fork and
the elongated basal tooth of the claw, T. alticola is closer to Mayacnephia and Tlalocomyia
than to the other Neotropical genera; it doesn’t have enough characters to include it in the
keys and to differentiate it from other species. More material would be necessary to
adequately differentiate this species.
66 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Araucnephia WYGODZINSKY & COSCARÓN (Figs. 11-14, 136.)


Araucnephia WYGODZINSKY & COSCARÓN, 1973a: 165.
Type-species: Simulium montanum PHILIPPI.
Adults: Antennae with 11 segments; male holoptic; female with frons comparatively
wide (Fig. 13 A); frontal angle 85 to 100 degrees; mandibles denticulate on both margins
(Fig. 13 D); apical segment of maxillary palp of both sexes less than twice as long as
penultimate (Fig. 13 C); mesepisternal sulcus shallow, evanescent anteriorly; katepisternum
glabrous, in profile higher than wide; pleural membrane glabrous; metanotum glabrous;
arms of furcasternum without projections (Fig. 13 F); wings with basal cell present; R1 with
spine-like setae, more delicate than those of C (Figs. 13 H-I); Sc setose; basal section of R
with hair arranged in several irregular rows; Rs entirely hairy below, hairy above only on
apical half; Rs generally simple, very rarely faintly forked apically; M appearing duplicated for
most of its length; Cu2 conspicuously curved in both sexes (Fig. 13 E); A2 ending well before
wing margin; calcipala present although comparatively small, bluntly rounded; claws of
female with medium-sized pointed subtriangular tooth (Figs. 11 D; 13 G); male with
sclerotized areas of abdominal terga increasing in size toward posterior segments; sclero-
tized areas of abdominal sterna small, transverse; spermatheca of normal size, well
sclerotized, with circular membranous area at insertion of spermathecal duct, the latter not
sclerotized at base; surface of spermatheca not sculptured; internal surface smooth, with
scattered spicules; gonapophyses short, subtriangular, apex rounded, their median borders
narrowly pigmented, approximated; surface of gonapophyses with microtrichia and few
setae (Figs. 11 E; 13 K); paraprocts subovoidal distally (Fig. 13 L); genital fork without
forwardly directed apodemes; arms narrow (Figs. 11 F; 13 M); male with gonostylus slightly
shorter than gonocoxite, broadly conical, with two apical spines; median surface of
gonostylus smooth, glabrous, delimited by two carinae (Figs. 11 G-H; 13 N, O); ventral plate
transverse to subtriangular, its basal arms in shape of elongate parallel-sided sclerotized rods
(Figs. 11 I; 13 P, Q); median sclerite in shape of complex geniculate plate; basal half entire,
fitting into concavity of ventral plate; distal half arising at sharp angle with basal half,
divided into two straight parallel arms narrowed toward their apex (Figs. 13 R-S);
endoparameres with large plate-like basal portion connected to numerous denticles forming
apical portion (Figs. 11 K; 13 T); median sclerite expanded basal and distally (Figs. 11 J-K).
Pupa: Cocoon large, shapeless, covering entire pupa except respiratory organs (Figs. 12
A; 14 A); body of pupa well sclerotized throughout; head and thorax with or without
tubercles; gills arborescent, divided very close to base into primary and secondary branches;
general structure of branches filamentous (Fig. 12 C); head sclerite with 1+1 frontal and
facial, and 2+2 epicranial, labral and genal trichomes, all hair like (Figs. 12 B; 14 B); thorax
with 6-8+6-8 simple hair like trichomes (Figs. 12 D; 14 C); abdomen pointed posteriorly;
distinct pleural areas not developed; onchotaxy of abdomen as follows: Terga III and IV
with 4+4 hooks posteriorly; terga VI-IX with continuous spine-combs anteriorly; sternum V,
or IV and V, with 2+2 hooks posteriorly; sternal plates VI and VII with 1+1 hooks, 1+1
hooks each also in pleural membranes of segments VI and VII (Fig. 14 D); terminal spines
well developed, from stout to slightly sinuous (Fig. 12 E); posterior segments with several
strong S-shaped or apically curved or looped setae, only very rarely branched.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 67

Larva: Head spots positive; cephalic apotome broadest well before hind margin
(Figs. 12 G; 14 F); cervical sclerites wide to faintly fused with upper ends of postocciput
(Fig. 14 J); postgenal cleft shallow but distinct, inverted U-shaped (Fig. 12 L); length of
hypostomium approximately equal to length of postgenal bridge; antennae much shorter
than stern of cephalic fan (Fig. 14 H); proximal segment of antennae very short, less than
half as long as medial; distal segment distinctly darker than remainder (Figs. 12 H; 14 G);
cephalic fans with rays unusually stout and heavily sclerotized (Fig. 12 I); mandibles with
two outer teeth and one apical tooth; third preapical tooth larger than remaining teeth, but
shorter than apical tooth; first and second preapical teeth subequal in size (Figs. 12 J; 14 I);
inner teeth arranged in several rows; one large mandibular serration followed by
numerous (10-15) very unequal smaller ones; short basal setae of mandibles very stout,
with conspicuous barbules; maxillary palp relatively elongated (Fig. 12 K); hypostomium
with 15 teeth arranged in three conspicuous groups, with median tooth and lateral teeth
by far the largest; 3+3 lateral teeth descending below level of remaining teeth (Fig. 14 K);
lateral sclerite of proleg about as long as wide, subtriangular, narrow at base, strongly
widened apically (Figs. 12 M; 14 L); setae of lateral sclerite less than half as long as
sclerite, inserted on distinct individual tubercles, arranged in irregular double row, this row
extending slightly beyond sclerite below; larval cuticle bare, except for perianal scales; anal
sclerite normal, X-shaped (Fig. 12 N); terminal ventral tubercles absent.
Distribution: Northeast of Argentina; south of Brazil and Mediterranean climate
area of central Chile (Fig. 136).
Discussion: Araucnephia differs from the Mexican Mayacnephia and Tlalocomyia by the
specialized median sclerite of the male genitalia and also by the presence of distinct
endoparameral spines; by the spiculae of the spermatheca; the single pair of frontal
trichomes and by the rarity of branched setae at the end of the abdomen of the pupa. In the
larva the first segment of the antenna is not more than half as long as the second, as it is in
Mayacnephia and Tlalocomyia. There are only two South American genera with comparable
hypostomial characters, Gigantodax and Araucnephioides. Gigantodax is easily distinguished
from Araucnephia by the absence of a basal cell and its specialized wing venation and the
different genitalia in both sexes, and the accessory sclerite forming a complete ring at the
posterior end of the body of the larvae. The pupae are not so easy to distinguish, but those
of Gigantodax do not have the full complement of cephalic trichomes, as described for
Araucnephia. Araucnephia is possibly closest to Araucnephioides; this is discussed under the
heading of the latter genus.
Cnesiamima differs from Araucnephia in the absence of spinelike setae on R1 and
specialized genitalic features, such as the structure of the median sclerite in the male and
that of the spermatheca in the female.

KEYS TO SPECIES OF ARAUCNEPHIA


Females
– Wing length 2.7-2.8 mm; mandible with about 40 (14+26) teeth; length/width
ratio of hind basitarsus = 8.5; spermatheca subovoidal with some superficial
tubercles .............................................................................................Araucnephia iberaensis
68 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

– Wing length 3.3-4.3 mm; mandible with about 55 (21+34) teeth; length/width
ratio of hind basitarsus = 6.5; spermatheca spheroidal without superficial
tubercles .............................................................................................. Araucnephia montana

Males
– Wing length 2.6-2.7 mm; scutum brown, lighter on posterolateral border;
abdomen brownish; gonostylus with small depression before subapical spurs
(Figs. 11 G, H) ..................................................................................Araucnephia iberaensis
– Wing length 3.5-3.6 mm; scutum velvety black; abdomen blackish; gonostylus
without depression before subapical spurs (Figs. 13 O, N) ..... Araucnephia montana

Pupae
– Cocoon length 3.6-4.2 mm; gill with 9 terminal branches; facial and thoracic
trichomes curved distally (Figs. 12 B, D); cephalothorax with abundant tubercles
..............................................................................................................Araucnephia iberaensis
– Cocoon length 7.0-8.3 mm; gill with about 14 terminal branches; facial and
thoracic trichomes not curved distally (Figs. 14 B, C); cephalothorax without
abundant tubercles ............................................................................ Araucnephia montana

Larvae
– Maximum length 6.6-7.6 mm; ratio of proximal, medial and distal antennal
segments = 1:1.2-1.5:2.4-2.8; cephalic fan with 48-53 rays; comb of thoracic
proleg sclerite with about 15-17 teeth (Fig. 12 M); anal ring with 64-66 rows with
14-16 hooks each. ............................................................................Araucnephia iberaensis
– Maximum length 11.0 mm; ratio of proximal, medial and distal antennal segments
= 1:2.7-3.0:2.7-3.2; cephalic fan with 23-26 rays; comb of thoracic proleg sclerite
with about 61 teeth (Fig. 14 L); anal ring with 80-100 rows with 18-22 hooks each.
............................................................................................................... Araucnephia montana

CLAVES DE ESPECIES DE ARAUCNEPHIA


Hembras
– Longitud ala 2,7-2,8 mm; mandíbula con alrededor de 40 (14+26) dientes;
relación largo/ancho de basitarso posterior = 8,5; espermateca subovoidal con
algunos tubérculos superficiales ....................................................Araucnephia iberaensis
– Longitud ala 3,3- 4,3 mm; mandíbula con alrededor de 55 (21+34) dientes;
relación largo/ancho de basitarso posterior = 6,5; espermateca esferoidal sin
tubérculos superficiales .................................................................... Araucnephia montana

Machos
– Longitud ala 2,6-2,7 mm; escudo castaño, más claro en el borde latero posterior;
abdomen castaño; gonostilo con una pequeña depresión delante de los espolones
subapicales (Figs. 11 G, H) ............................................................Araucnephia iberaensis
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 69

– Longitud ala 3,5-3,6 mm; escudo negro aterciopelado; abdomen negruzco;


gonostilo sin depresión delante de los espolones subapicales (Figs. 13 O, N) .......
............................................................................................................... Araucnephia montana

Pupas
– Longitud del capullo 3,6-4,2 mm; branquia con 9 ramas terminales; tricomas faciales
y torácicos curvados distalmente (Figs. 12 B, D); cefalotórax con abundantes
tubérculos ...........................................................................................Araucnephia iberaensis
– Longitud del capullo 7,0-8,3 mm; branquia con alrededor de 14 ramas terminales;
tricomas faciales y torácicos no curvados distalmente (Figs. 14 B, C); cefalotórax
sin abundantes tubérculos ............................................................... Araucnephia montana

Larvas
– Longitud máxima 6,6-7,6 mm; relación de los artejos antenales proximal mediano
y distal = 1:1,2-1,5:2,4-2,8; abanico cefálico con 48-53 rayos; peine del esclerito de
la propata torácica con alrededor de 15-17 dientes (Fig. 12 M); anillo anal con 64-
66 hileras con 14-16 ganchos cada una .......................................Araucnephia iberaensis
– Longitud máxima 11,0 mm; relación de los artejos antenales proximal mediano y
distal = 1:2,7-3,0:2,7-3,2; abanico cefálico con 23-26 rayos; peine del esclerito de
la propata torácica con alrededor de 61 dientes (Fig. 14 L); anillo anal con 80-100
hileras con 18-22 ganchos cada una .............................................. Araucnephia montana

Araucnephia iberaensis COSCARÓN & COSCARÓN ARIAS (Figs. 11 A-K; 12 A-N.)


Araucnephia iberaensis COSCARÓN & COSCARÓN ARIAS, 2002: 81-87.
This species is very close to Araucnephia montana and here we include only the
differences from A. montana.
Female: Wing length 2.7-2.8 mm; frons, clypeus, antenna, palp and occiput black
with grayish pollinosity; scutum and metanotum blackish, ornamented as in figure 11 A;
legs brownish darkened on tarsi; abdomen blackish with gray pollinosity; frontal angle =
100°; fronto-ocular triangle as in figure 11 B; last segment of maxillary palp about twice
length of penultimate segment; sensory vesicle small about 1/5 of third palpomere length
(Fig. 11 C); mandible denticulate on both sides with 14 (15)-26 (27) teeth; lacinia with
about 25 teeth; length/width ratio of hind basitarsus = 8.5; calcipala small subovoidal;
eighth sternite mostly black; gonapophyses with 6-7 hairs (Fig. 11 E); spermatheca
subovoidal, showing some superficial tubercles; genital fork with lateral branches distally
expanded (Fig. 11 F).
Male: Wing length 2.6-2.7 mm Scutum brown, lighter on posterolateral border;
abdomen brownish, a longitudinal blackish stripe composed of subtrapezoidal sclerotized
tergal plates on segments II-V and 1+1 sublateral spots on segments VI-VIII; ventrally
with subquadrate spots blackish on segments III-VIII; length/width ratio of hind
basitarsus = 4.4; gonostylus distally with depression before two apical spurs (Figs. 11 G-
H); ventral plate subtrapezoidal (Fig. 11 I); median sclerite and aedeagus as in figures 11 J-
K, endoparameres with some well-developed denticles (Fig. 11 K).
70 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

F
B

C
D

Figure 11. Araucnephia iberaensis. A-F: Female; A: head and thorax, dorsal view; B: fronto-ocular
triangle; C: sensory vesicle of maxillary palp; D: claw; E: eighth sternite and gonapophyses; F: Genital
fork. G-K: Male; G: general aspect of genitalia dorsal view, showing gonocoxite, gonostylus, ventral
plate, median sclerite and endoparameres; H: gonostylus; I: ventral plate; J-K: different view of
aedeagus membrane, base and scarcely evident hooks of endoparameres and median sclerite.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 71

B
A

C
E

o.t.
G H
I a.t. J
p.t.

i.t.

m.s.

L N
K

Figure 12. Araucnephia iberaensis. A-E: Pupa; A: general aspect; B: frontoclypeus, with trichomes and
tubercles magnified; C gill; D: thoracic trichomes; E: abdominal terminal area, showing curled
trichomes. F-N: Larva; F: head, lateral view; G: cephalic apotome; H: antenna; I: teeth of cephalic
fan rays; J: apical portion of mandible with teeth (a.t. = apical teeth, i.t. = inner teeth, m.s. =
marginal serrations, o.t. = outer teeth, p.t. = preapical teeth); K: maxillary palp; L: postgenal cleft
and hypostomium; M: lateral sclerite of thoracic proleg; N: anal sclerite with scales and hairs.
72 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Pupa: Cocoon total length on dorsum 3.6-4.2 mm; total pupa length 6.0-6.3 mm;
cocoon not compacted, generally covering pupa up to base of gill (Fig. 12 A).
frontoclypeus and thorax with abundant tubercles mostly verrucose (Fig. 12 B); abdomen
with abundant tubercles; gill with 9 terminal branches with 3 short primary branches (Fig.
12 C); trichomes of head and thorax single and mostly curved distally (Fig. 12 D) as for
terminal area of the abdomen (Fig. 12 E).
Larva: Maximum length 6.6-7.6 mm; head light brown, darkened around eyes (Fig.
12 F); cephalic apotome with positive spots blackish brown and very evident (Fig. 12 G);
antenna about same length as stem of cephalic fan; ratio of proximal, medial and distal
antennal segments = 1:1.2-1.5:2.4-2.8 (Fig. 12 H); cephalic fan with 48-54 rays; disposition
of ray setae as in figure 12 I; mandible with marginal serrations of 8-12 teeth (Fig. 12 J);
maxillary palp as in figure 12 K; hypostomium and postgenal cleft as in figure 12 L; comb
of thoracic proleg sclerite with 15-17 teeth (Fig. 12 M); anal ring with 64-66 rows with 14-
16 hooks each.
Distribution: Argentina: Corrientes; Brazil: Paraná (unpubl. information).
Bionomics: The species breeds in small creeks 0.5-3.0 m wide and 0.1-0.5 m deep in
crystalline water and strong but not torrential currents; larvae and pupae are found on
vegetation. Females are haematophagus and bite humans, producing a hard spot followed
by an allergic reaction.
Discussion: The morphological characters allow it to be differentiated from Arauc-
nephia montana, as seen in the key.

Araucnephia montana (PHILIPPI) (Figs. 13 A-T; 14 A-L.)


Simulium montanum PHILIPPI, 1865: 633.
Simulium (Cnephia) montanum: EDWARDS, 1931: 149.
Cnephia montanum SMART, 1945: 498.
Araucnephia montana WYGODZINSKY & COSCARÓN, 1973a: 168-179; COSCARÓN, 1991: 14-15; CROSSKEY
& HOWARD, 1997: 11; 2004: 2.
Female: Length of wing, 3.3-4.3 mm, generally almost 4.0 mm (type 3.9 mm); head
black; occiput, frons, and clypeus gray pruinose; antennae, palpi, and labrum brownish
black; pilosity of antenna very short, that of palpi, clypeus, frons, and occiput somewhat
longer, its color from silvery gray to black; thorax dull black, with silvery gray pollinosity;
metapleura and wing base brownish; hair of thorax silvery gray; wings hyaline; veins light
brown, hair and spines blackish; tufts at wing base from silvery to brownish yellow;
halteres light brown; legs from light brown with darkened apices of femora, tibiae and
tarsi, to entirely dark brown or black; most specimens with legs dark, concolorous; hair of
legs from silvery gray to black; abdomen from dark grayish brown to dull black; dorsal
surface always darkest; hair of abdomen from silvery gray to black; frons as shown in
figure 13 A; frontal angle 85-90 degrees; fronto-ocular triangle (Fig. 13 B) approximately
as long as wide; last segment of maxillary palp (Fig. 13 C) slightly less than twice as long as
penultimate; diameter of sensory vesicle smaller than half diameter of third palpomere;
lacinia with 30-35 teeth; mandible with about 75 teeth (Fig. 13 D); furcasternum as in
figure 13 F; wings as in generic description and figure 13 E; spine-like setae of R1 often
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 73

A
C
B
E

G
D
C

H R1 I

Rs

F
L
K

M J
Q

R T
N O S
Figure 13. Araucnephia montana. A-L: Female; A: frons; B: fronto-ocular triangle; C: maxillary palp;
D: mandible; E: wing, general view; F: furcasternum; G: claw; H-I: portion of wings showing part
of anterior veins with chaetotaxy in different specimens (c = costa, R1= radius 1, Rs= secondary
radius (radius 2) ); J: calcipala and tarsomere I; K: eighth sternite and gonapophyses; L: cercus and
paraproct; M: genital fork. N-T: Male; N: gonocoxite and gonostylus, dorsolateral view; O:
gonostylus, showing membranous inner surface; P: ventral plate, anteroventral view; Q: ventral
plate and median sclerite in lateral views; R-S: median sclerite in different position; T:
endoparameres. (A-T from WYGODZINSKY & COSCARÓN, 1973a).
74 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

A
D

E F

G
J I
Figure 14. Araucnephia montana. A-D: Pupa; A: general view; B: cephalic sclerite with trichomes; C:
portion of thorax with trichomes; D: abdominal chaetotaxy. E-L: Larva; E: general aspect in lateral
view; F: cephalic apotome; G: antenna; H: portion of head dorsal view showing short antenna; H:
portion of mandible with teeth; J: posterior region of head, with cervical sclerites; K: anterior portion
of hypostomium; L: sclerite of thoracic proleg. (A-L from WYGODZINSKY & COSCARÓN, 1973a).
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 75

distinctly more slender than those of C (Fig. 13 H-I); hind basitarsus 6.1-6.9 times as long
as wide; claws as in figure 13 G; second hind tarsomere with strong rugosity (Fig. 13 J);
eighth abdominal sternum as shown in figure 13 K; pigmentation variable, from only
central area darkened to entire sternum dark; gonapophysis with abundant microtrichia
and 10-14 short setae; cerci and paraprocts as illustrated in figure 13 L; cerci much wider
than high; paraprocts short, their free extremity somewhat truncate; genital fork with stem
and apical portion of arms well pigmented (Fig. 13 M).
Male: Length of wing, 3.5-3.6 mm; head black; clypeus dark gray pollinose;
antennae, palpi, and proboscis brownish black; antennae with short pubescence; setae of
basal segments longer than that of remaining segments; occiput, line between eyes,
clypeus and basal segments of maxillary palps with several long hairs; thorax velvety
black; pleura with silvery gray pollinosity; scutum with decumbent hair from light silvery
to light brassy; distal portion of scutum as well as scutellum and metanotum with black,
erect setae; color of wings, halteres and legs as in female; abdomen blackish, dark dorsally,
grayish laterally and below; hair of abdomen colored as on thorax; chaetotaxy of wings as
in female, but fewer setae on Sc; spiniform setae of R1 distinctly smaller than those of C;
in some specimens, Rs with short bifurcation apically; hind basitarsus 3.9-4.6 times as long
as wide; calcipala narrowly rounded apically; gonostylus without apical depression (Figs.
13 N, O); ventral plate with arms and basal margin distinctly sclerotized, disc uniformly
pigmented (Figs. 13 P-Q); median sclerite and endoparameres as in figures 13 R, S and T.
Pupa: Cocoon covering pupa completely or almost so (Fig. 14 A), in some cases also
including part or all of gills; cocoon closely woven, threads not perceptible under medium
magnification; color light grayish brown; length along dorsum, 7.0-8.3 mm; total length of
pupa 4.0-4.5 mm; cephalopterothecal length, 3.0 mm; gill composed of six short primary
branches arising from extremely short common trunk; the first two of three dorsal primary
branches generally several times as long as wide, the third much shorter than others; all
giving rise to two filaments; two median branches giving origin to three filaments each,
filaments arising at variable relative positions; ventral primary branch very short, with two
filaments delicately pitted; head and thorax of pupa light brown; trichomes of head as in
figure 14 B; epicranial, lateral and genal trichomes in pairs, others single; all trichomes
simple; head sclerite lacking tubercles; thorax without tubercles and with not more than
eight trichomes on each side on disc (Fig. 14 C); abdominal segments distinctly sclerotized
although somewhat less so than thorax, intensity of sclerotization diminishing toward
posterior segments; onchotaxy and accessory hair and setae of abdomen as in generic
description and figure 14 D; tubercles on abdomen absent or few in number.
Larva: Maximum length, 11.0 mm, maximum width of head capsule, 1.2 mm.
General shape as shown in figure 14 E; color of larva light grayish brown, head often very
dark; cephalic apotome (Fig. 14 F) yellowish brown, darker along middle, along hind
border, and with one or two sublateral spots on each side; antennae as in figure 14 G; ratio
of lengths of proximal, medial and distal segments = 1:2.75-3.0:2.75-3.2; cephalic fan
with 23-26 rays; mandibles as in figure 14 I, with 13-20 inner teeth and 7-15 marginal
serrations irregularly decreasing in size; cervical sclerite as in figure 14 J; hypostomium as
shown in figure 14 K, its anterior border strongly pigmented; lateral border of
hypostomium with 3-8 distinct serrations; hypostomial setae arranged in one, rarely in two
76 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

irregular series, with 5-7 setae in each group; disc of hypostomium with few scattered
short hairs; postgenal cleft about half as long as postgenal bridge; lateral sclerite of
thoracic proleg as in figure 14 L; anal sclerite with numerous perianal scales, two to five-
pronged; crochet ring with 80-100 rows with 18-22 hooks each.
Distribution: Chile: Aconcagua, Coquimbo, Curico, Santiago, Valparaiso.
Bionomics: We have collected the aquatic stages of Araucnephia montana from the
surface of stones and rocks exclusively in small, clear, temporary streams, toward the end
of the wet season. This is apparently a species that completes its cycle in late winter or
very early spring (September or very early October).
Discussion: The closest relative of Araucnephia may be Araucnephioides; both genera
are sympatric, and individuals are often difficult to distinguish. We, therefore, present a
detailed enumeration of differential characters to help to identify specimens.

Differences between Araucnephia and Araucnephioides


Araucnephia
Female: Basal tooth of claws subtriangular (Figs. 11 D; 13 G); calcipala broadly
rounded (Fig. 13 J}; inner surface of spermatheca not rugose.
Male: Genitalia with subtriangular ventral plate, its basal arms narrow (Figs. 11 I; 13 P),
straight; arms of median sclerite parallel (Fig. 13 S); endoparameral organ with basal portion
well developed, plate like, and teeth of distal portion conspicuous (Figs. 11 K; 13 T).
Pupa: Gill with 14-17 filaments; head sclerite with full complement of trichomes,
i.e., frontal, facial, epicranial, ocular, and genal (Figs. 12 B; 14 B).
Larva: Antenna much shorter than stem of cephalic fan (Figs. 12 F; 14 H); rays of
cephalic fan very heavily sclerotized (Fig. 12 I).

Araucnephioides
Female: Basal tooth of claws narrowly hook like (Fig. 15 I); calcipala subtriangular
(Fig. 15 H).
Male: Genitalia with ventral plate subrectangular, its basal arms stout, rounded at
apex and recurved (Fig. 15 P); arms of median sclerite sharply diverging apically (Fig. 15
U); endoparameral organ with basal portion small and denticles of apical portion faint.
Pupa: Gill with 10-13 filaments; head sclerite without or with poorly developed
frontal trichomes (Fig. 16 D).
Larva: Antenna almost as long as stem of cephalic fan (Fig. 16 I); sclerotization of
rays of cephalic fan normal.

Araucnephioides WYGODZINSKY & COSCARÓN (Figs. 15, 16, 136.)


Araucnephioides WYGODZINSKY & COSCARÓN, 1973a: 179.
Type-species: Araucnephioides schlingeri WYGODZINSKY & COSCARÓN, 1973a.
Adults: Female with frons medium wide (Fig. 15 A); frontal angle 60-70 degrees;
antenna with 11 segments (Fig. 15 B); male holoptic; mandibles of female denticulate
on both margins; apical segment of maxillary palp of both sexes less than twice as long
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 77

as penultimate segment (Fig. 15 C); cibarium with basal portion smooth and with strong
concavity (Fig. 15 F); mesepisternal sulcus shallow, evanescent anteriorly; katepisternum
glabrous, in profile higher than wide; pleural membrane bare; metanotum glabrous;
arms of furcasternum without projections (Fig. 17 J); wings with basal cell present (Fig.

C
B A
I
G

E
H

L
D
F K

S
O

P
Q

T U

J R

Figure 15. Araucnephioides schlingeri. A-M: Female; A: frons; B: Antenna; C: maxillary palp; D:
sensory vesicle of maxillary palp; E: fronto-ocular triangle; F: basal portion of cibarium; G: wing,
general view; H: calcipala and tarsomere I; I: claw; J: furcasternum; K: genital fork; L: cercus and
paraproct; M: eighth sternite and gonapophyses. N-U: Male; N: gonocoxite and gonostylus; O:
gonostylus, showing membranous inner area; P: ventral plate with median sclerite visible through it;
Q: idem side view; R: idem full view; S: idem, with median sclerite and endoparameres; T-U: median
sclerite in different positions. (A-U from WYGODZINSKY & COSCARÓN, 1973a).
78 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

15 G); R1 with spinelike setae more delicate than those of C; Sc setose; basal section of
R with hair arranged in several irregular rows; Rs entirely hairy below, hairy above only
on apical half; Rs not forked apically; M appearing duplicated for most of its length; Cu2
conspicuously curved in both sexes; A2 ending well before wing margin; calcipala small,
narrowly subtriangular (Fig. 15 H); claws of female with medium-sized, pointed, and
curved hook-like basal tooth (Fig. 15 I); male with sclerotized areas of abdominal
terga of subequal size on all segments; sclerotized areas of abdominal sterna
subquadrate; spermatheca globular, of normal size, well sclerotized, with circular
membranous area at insertion of spermathecal duct, the latter not sclerotized at base;
surface of spermatheca not sculptured; internal surface irregularly rugose and with
scattered spicules; gonapophyses short, subtriangular, apically rounded, their median
borders pigmented, approximated; surface of gonapophysis with microtrichia and
setae (Fig. 15 M); genital fork without forwardly directed apodemes; posterior arms
wide (Fig. 15 K); male with gonostylus shorter than gonocoxite, broadly conical, with
two or three apical spines; median surface of gonostylus smooth, glabrous, delimited
by two carinae (Figs. 15 N, O); ventral plate subquadrate, its anterior border truncate
or slightly emarginated; its basal arms with terminal knobbed sclerotization (Figs. 15
P-S); median sclerite in shape of complex geniculate plate; basal half entire,
transverse, fitting into concavity of ventral plate; distal half arising at sharp angle
with basal half, divided into two apically, outwardly bent strong arms (Figs. 15 T-U);
endoparameral organ with narrow plate-like basal portion tapering into very narrow
obsolescent apical portion; denticles weakly developed.
Pupa: Cocoon large but shapeless, covering entire pupa except gills (Fig. 16 A); head
and thorax with numerous tubercles (Figs. 16 D-E); body of pupa well sclerotized
throughout; gills arborescent, consisting of several primary branches and not very
numerous (approximately 10) narrow filaments arranged in two more or less distinct
groups (Fig. 16 B); head sclerite without or with 1-2+1-2 very short frontal and with 1+1
hair-like well-developed facial trichomes; thorax with 6-8+6-8 simple hair-like trichomes;
abdomen pointed posteriorly; its structure and onchotaxy as in Araucnephia.
Larva: Body slightly wider posteriorly on preapical portion (Figs. 16 F); head spots
positive, distinct (Fig. 16 G); cervical sclerites small, isolated in membrane (Figs. 16 H-
I); antenna almost as long as stern of cephalic fan (Fig. 16 I); proximal antennal segment
very short, less than half as long as medial segment (Fig. 16 J); distal segment only very
slightly darker than remaining; cephalic fans with rays normal, slender, not heavily
sclerotized; setae of all rays distinctly of two sizes; maxillary palp normal (Fig. 16 K);
mandibles with two external teeth and one apical tooth; third preapical tooth larger than
the two others, but shorter than apical tooth; second preapical tooth minute; inner teeth
arranged in several rows; one large basal mandibular serration followed by numerous
smaller ones; accessory mandibular serrations situated on border of mandible at level
of internal teeth (Fig. 16 L); short setae of mandible slender, only very faintly dentate
or not at all; postgenal cleft shallow but distinct, biarcate (Fig. 16 M); hypostomium
shorter than length of postgenal bridge; hypostomium with 17 teeth arranged in three
conspicuous groups, with median tooth and lateral teeth by far largest; 4+4 lateral teeth
descending below level of remaining teeth (Fig. 16 N); lateral sclerite of thoracic proleg
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 79

as in Araucnephia; larval cuticle bare except for perianal scales; anal sclerite normal, X-
shaped; ventral tubercles absent.
Distribution: Mediterranean climate area of central Chile.

D B

I
C

L
G H

P O

Figure 16. Araucnephioides schlingeri. A-E: Pupa; A: general view; B: gill; C: basal portion of gill; D:
frontoclypeus; E: arrangement of tubercles of exposed portion of thorax; F-P: Larva; F: dorsal and
lateral views; G: cephalic apotome; H: posterior region of head with reduced cervical sclerites; I:
portion of cephalic capsule showing antenna and stem of cephalic fan; J: antenna; K: maxillary
palp; L: portion apical mandible and teeth; M: postgenal cleft and hypostomium; N: anterior border
of hypostomium; O: anal sclerite; P: rectal papillae. (A-P from WYGODZINSKY & COSCARÓN, 1973a).
80 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Araucnephioides schlingeri WYGODZINSKY & COSCARÓN (Figs. 15 A-U; 16 A-P.)


Araucnephioides schlingeri WYGODZINSKY & COSCARÓN, 1973a: 181-184; COSCARÓN, 1991: 17; CROSSKEY
& HOWARD, 1997: 11.
Female: Length of wing, 3.2-4.0 mm, in most specimens more than 3.5 mm; head
blackish, eyes dark piceous; occiput, frons and clypeus grayish brown pollinose; antennae
and palpi dark brown, labrum light grayish brown; pilosity of antennae very short, except
on two basal segments, that of palpi, clypeus, frons, and occiput somewhat longer; hair
from brassy to black; scutum dull black, with gray pollinosity and decumbent hair of same
color as that of occiput; pleura brownish, with silvery gray pollinosity; base of wings light
brown; scutellum piceous, hair semi-erect, from light gray to dark brown; metanotum and
katepisternum brown to black, with silvery pollinosity; wings and veins hyaline, veins light
brown; halteres piceous; legs in most specimens dark brown, with apices of femora, tibiae,
and tarsi piceous; in some specimens legs lighter, with dark apical regions of leg segments
more conspicuous; hair from silvery gray to blackish; color of abdomen dark grayish
brown to blackish; frons as shown in figure 15 A; frontal angle 60-70 degrees; fronto-
ocular triangle slightly wider than high (Fig. 15 E); shape and proportions of antennal
segments as shown in figure 15 B; mandible with 36-40 teeth; last segment of maxillary
palp (Fig. 15 C) distinctly less than twice as long as penultimate segment; diameter of
sensory vesicle (Fig. 15 D) less than half the diameter of third segment; maxillae with
about 30 teeth; wing as in figure 15 G, calcipala about as long as wide at base (Fig. 15 H);
claws as in figure 15 I; hind basitarsus 6.3-7.6 times as long as wide; eighth abdominal
sternum with abundant setae; gonapophyses each with 10-14 medium-sized hairs and with
numerous microtrichia (Fig. 15 M); paraprocts short (Fig. 15 L); genital fork with wide
lateral branches (Fig. 15 K).
Male: Length of wing, 2.9-3.3 mm; color of head, body and appendages brown dark
to blackish; last segment of maxillary palp less than twice as long as penultimate segment;
diameter of sensory vesicle smaller than half diameter of third segment; hind basitarsus
3.7-4.6 times as long as wide; genitalia as in generic description (Figs. 15 N-U).
Pupa: Cocoon (Fig. 16 A) covering entire pupa, in some cases including gills and
often also containing larval exuviae; color of cocoon very light brown, loosely woven with
body of pupa discernible; threads visible under medium magnification; length of cocoon
along dorsum, 4.0-6.0 mm; total length of body of pupa 3.5-4.3 mm, cephalopterothecal
length 2.05-2.10 mm; gills (Figs. 16 B-C) about half as long as body of pupa, or slightly
longer, with 10-12, rarely 13, filaments; two primary branches arising from very short
common trunk; dorsal primary branch with very short secondary branch immediately
beyond base, divided into two branches; secondary branch in some cases arising directly
from base, thus constituting third primary branch; dorsal primary branch with four or five,
rarely six, additional filaments; ventral primary branch generally giving origin to four,
rarely five filaments, two originating very close to base, the others slightly above; surface
of filaments delicately pitted, and with irregularly shaped and distributed minute plate-like
sclerotized structures basally; head and thorax of pupa light brown; frontoclypeus with
numerous tubercles (Fig. 16 D); thorax with numerous tubercles, arranged in roughly
circular groups, composed of five to eight tubercles each (Fig.16 E); abdomen somewhat
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 81

lighter than thorax; onchotaxy much as in Araucnephia montana, but spine-combs on


tergum V often absent, and composed of fewer spines (about 15) on tergum VI; all terga
with numerous tubercles in addition to usual hair and spines; specialized setae of eighth
and ninth segments with strong curvature distally.
Larva: Maximum length, 8.0 mm; maximum width of head capsule, 0.75 mm; body
shape as shown in figure 16 F; color of larva light grayish brown; cephalic apotome
yellowish brown, darker along middle, and with several spots as shown in figure 16 G;
antennae as in figure 16 J; ratio of length of proximal, medial and distal segments =
1:2.2:2.9-3; cephalic fan with 37-42 rays; mandible as in figure 16 L, with 15-17 inner teeth
and 10-15 marginal serrations; hypostomial teeth as shown in figure 16 N; anterior area of
hypostomium strongly pigmented; lateral border of hypostomium with 6-9 serrations;
hypostomial setae arranged on each side in one irregular row composed of about four
setae; disc of hypostomium with few scattered, short setae; postgenal cleft short and
subquadrate (Fig. 16 M); anal sclerite with perianal scales as shown in figure 16 O; crochet
ring with 60-67 rows composed of 18-20 hooks each.
Distribution: Same as for genus.

Cnesia ENDERLEIN (Figs. 17-19, 136.)


Cnesia ENDERLEIN, 1934b: 273; STONE, 1963: 4; WYGODZINSKY & COSCARÓN, 1973a: 151; COSCARÓN,
1991: 18; CROSSKEY & HOWARD, 1997: 11.
Type-Species: Simulium (Cnephia) dissimile EDWARDS, 1931 (as Simulium (Cnephia) gynandrum
EDWARDS, 1931).
Adults: Antennae with 11 segments; male holoptic; female with frons medium wide;
frontal angle 50-67 degrees; mandibles with well-developed teeth only on internal margin,
external border weakly serrated (Fig. 17 A); apical segment of maxillary palp in both sexes
less than twice as long as penultimate segment; mesepisternal sulcus shallow, evanescent
anteriorly; katepisternum glabrous, in profile slightly longer than high; pleural membrane
glabrous; metanotum with hair in some but not all species; arms of furcasternum lacking
conspicuous projections (Fig. 17 E); wings with basal cell present (Fig. 17 B); R1 with hair
and spine-like setae, the latter not more delicate on R1 than on C (Fig. 17 I); Sc and basal
section of R with hair arranged in several irregular series; Rs entirely hairy below, hairy
above only on apical half; Rs not forked apically; M not appearing double; Cu2 only weakly
curved in both sexes; A2 almost attaining wing margin; calcipala present, large, bluntly
rounded apically (Fig. 17 C); claws of female with large subtriangular basal tooth (Fig. 17
D); spermatheca of normal size, normally sclerotized, oval; internal surface minutely
wrinkled and with small irregularly scattered spicules; area of insertion of spermathecal
duct membranous, distinctly larger than diameter of duct; duct not sclerotized (Fig. 17 F);
gonapophyses subtriangular, transverse, their medial borders narrowly pigmented, ap-
proximated, their surface with microtrichia and few setae; paraproct short (Fig. 17 G);
posterior arms of genital fork wide, with small but distinct forwardly directed apodemes
(Figs. 17 H-Q); male with gonostylus distinctly shorter than gonocoxite, narrowly conical,
with two apical spines; medial surface of gonostylus smooth, delimited by two carinae
(Figs. 17 K-L); ventral plate subpentagonal, about as long as wide, setose, with very low
82 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

median longitudinal keel (Figs. 17 M-N; 19 C-E); median sclerite with flaring base, then
narrowed, apically widened again and with two wing-like pointed subapical flanges (Figs.
17 O-P; 19 F-G); endoparameral organs with basal portion strongly reduced, apical
portion composed of a group of one or two large and several oppositely directed
medium-sized denticles.
Pupa: Cocoon large, shapeless, covering pupa to base of respiratory organs (Fig. 18
A); body of pupa well sclerotized throughout; head and thorax with or without tubercles;
gills bushy, much shorter than pupa proper, composed of 35-40 slender filaments arising
from about 12 primary branches very close to their base (Figs. 18 B; 19 H); head with 1+1
prefrontal, 1+1 or 2+2 frontal, 1+1 facial, and occasionally 1+1 labial trichomes;
prefrontal and single or anterior frontal trichomes spinelike; posterior frontal, facial and
labial trichomes hair like (Figs. 18 C; 19 I); disc of thorax with approximately 5+5
trichomes, two or three dorsalmost ones in shape of strong spines, others hair like;
abdomen pointed posteriorly; distinct pleural areas not developed; terga with numerous
small tubercles, those near anterior margins of sclerite arranged in very conspicuous
transverse ridges; onchotaxy of abdomen as follows: terga III and IV with 4+4 short
hooks posteriorly; tergum VIII with 2+2 very elongate spines posteriorly; terga without
spines-comb; sterna IV and V with 3+3 hooks posteriorly, those of IV arranged in one
median and 1+1 submedian groups of two spines each, those on V arranged in 1+1
closely approximated groups of three spines each; sternal plates VI with 2+2 hooks, VII
with 1+1 hooks; 1+1 hooks each also in pleural membrane of segments VI and VII (Fig.
18 D); terminal spines well developed, stout, not sinuous; posterior segments with several
strong S-shaped setae, often apically curved, looped or spiraled, occasionally branched
near base or apex (Fig. 18 E).
Larva: Body shape with wider preapical region (Fig. 19 K); head spots positive;
cephalic apotome broadest well before hind margin (Fig. 18 F); cervical sclerites very small,
isolated in membrane; postgenal cleft very shallow, broadly U-shaped (Figs. 18 I; 19 N);
antennae about as long as stem of cephalic fans; medial antennal segment about two times
as long as proximal, distal segment darker than remaining segments (Figs. 18 G; 19 L);
mandibles with four outer teeth in addition to apical tooth; third preapical tooth longer and
more strongly sclerotized than other preapical teeth, as long as apical tooth; first inner tooth
as long and stout as third preapical tooth; inner teeth arranged in three or more series; two
large mandibular serrations followed by two or three small ones (Fig. 18 J); hypostomium
slightly bi-emarginate anteriorly, with 17 teeth, not arranged in conspicuous groups; median
tooth and lateral teeth somewhat larger than remaining teeth; 4+4 lateral teeth inserted
almost at same level as other teeth (Figs. 18 H; 19 M); lateral sclerite of thoracic proleg well
developed, rather narrow for most of length but abruptly flaring below and above at
anterior margin (Fig. 18 K); setae conspicuously decreasing in size from dorsal to ventral
portion of anterior border of sclerite; lowermost setae barbed, some others once or twice
branched on their apical half; setae arranged in one or two irregular rows, not arranged in
groups and not inserted upon distinct tubercles; larval cuticle bare, except perianal scales;
anal sclerite normal, X-shaped (Fig. 18 M); terminal ventral tubercles absent.
Distribution: Valdivia and Magellanic areas of southern Chile and western Patagonia,
with slight intrusion into Mediterranean climate area of central Chile (Fig. 136).
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 83

Discussion: Cnesia differs from Cnephia by the much smaller spermatheca that is
unusually large in Cnephia, from Stegopterna by the simplified hypostomial teeth of the
larva, and from both genera by the large basal tooth on the claw of the female. Among the
Neotropical Prosimuliini there are two others in which the teeth of the hypostomium of
the larva are not arranged in three prominent groups: Lutzsimulium and Paraustrosimulium.
Cnesia shares with Gigantodax the apomorphic structure of the genitalia in both sexes, and
on this basis alone the two genera are indistinguishable. Cnesia and Gigantodax differ
considerably in other ways (wing venation, onchotaxy of the abdomen of the pupa,
several characters of the larvae), but their synapomorphic genitalia suggest common
ancestry not shared by any other known genus.

KEYS TO SPECIES OF CNESIA


Females
1 Metanotum glabrous ...................................................................................... Cnesia ornata
– Metanotum with short hair posteriorly ......................................................................... 2
2(1) Scutum pale reddish brown, dark brown on disc in some specimens; wing length
more than 3.0 mm ..................................................................................... Cnesia dissimilis
– Scutum black, wing length 3.0 mm or less ............................................... Cnesia pusilla

Males
1 Scutum bright reddish brown; metanotum glabrous .............................. Cnesia ornata
– Scutum varying from dark brown to black; metanotum with short hair posteriorly
................................................................................................................................................ 2
2(1) Wing length more than 3.0 mm; scutum varying from dark reddish brown to black
........................................................................................................................ Cnesia dissimilis
– Wing length 3.0 mm or less; scutum black ............................................... Cnesia pusilla

Pupae
1 Head and thorax with numerous tubercles; those on thorax generally arranged in
circles (Fig. 19 J) ............................................................................................. Cnesia ornata
– Head and thorax without tubercles ................................................................................ 2
2(1) Cephalopterothecal length approximately 2.5 mm ............................. Cnesia dissimilis
– Cephalopterothecal length approximately 2.0 mm ................................. Cnesia pusilla

The larvae of the species of Cnesia are so similar that we have been unable to find
any differential characters, but larvae of Cnesia pusilla from Argentina can be distinguished
from those of the two other species by their smaller head width.

CLAVES DE ESPECIES DE CNESIA


Hembras
1 Metanoto glabro ............................................................................................. Cnesia ornata
– Metanoto posteriormente con cortos pelos ................................................................. 2
84 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

2(1) Escudo castaño rojizo claro, castaño oscuro sobre el disco en algunos
especimenes; longitud del ala mayor de 3,0 mm ................................. Cnesia dissimilis
– Escudo negro, longitud del ala 3,0 mm o menos .................................... Cnesia pusilla

Machos
1 Escudo castaño rojizo brillante; metanoto glabro .................................. Cnesia ornata
– Escudo variando desde castaño oscuro a negro; metanoto posteriormente con
cortos pelos ......................................................................................................................... 2
2(1) Longitud ala mayor de 3,0 mm; escudo variando desde castaño rojizo oscuro a
negro ............................................................................................................. Cnesia dissimilis
– Longitud ala 3,0 mm; escudo negro........................................................... Cnesia pusilla

Pupas
1 Cabeza y tórax con numerosos tubérculos, estando los del tórax generalmente
ordenados en círculos (Fig. 19 J) ................................................................ Cnesia ornata
– Cabeza y tórax sin tubérculos .......................................................................................... 2
2(1) Longitud de cefalopterotecas aproximadamente de 2,5 mm ............ Cnesia dissimilis
– Longitud de cefalopterotecas aproximadamente 2,0 mm ..................... Cnesia pusilla

Las larvas de las especies de Cnesia son tan similares que no hemos podido encontrar
caracteres diferenciables; pero la larva de Cnesia pusilla de Argentina puede ser separada de
las otras2especies por su cabeza que es menos ancha que los otros.

Cnesia dissimilis (EDWARDS) (Figs. 17 A-P; 18 A-M.)


Simulium (Cnesia) dissimilis EDWARDS, 1931: 145.
Cnephia dissimilis SMART, 1945: 498.
Cnesia dissimilis VARGAS & DÍAZ NÁJERA, 1951: 166; COSCARÓN & WYGODZINSKY, 1962: 310;
COSCARÓN, 1968: 66; 1991: 20-21; WYGODZINSKY & COSCARÓN, 1973a: 155-159; COSCARÓN,
1991: 20; CROSSKEY & HOWARD, 1997: 12; 2004: 3.
Stegopterna dissimilis ENDERLEIN, 1934b: 274.
Simulium (Cnephia) gynandrum EDWARDS, 1931: 147.
Cnephia gynandrum SMART, 1945: 498.
Cnesia gynandra ENDERLEIN, 1934b: 278.
Cnesia gynandrum VARGAS & DÍAZ NÁJERA, 1951: 166.
Cnephia (Stegopterna) gynandrum: RUBTZOV, 1940: 132.
Simulium gynandrum VARGAS, 1945b: 142.
Female: Wing length 4.2-5.3 mm; head brown; scutum light reddish brown to dark
brown in some specimens; scutellum light yellowish gray; metanotum brown; pleura
brown violaceous; hyaline, wing with veins light brown with hair and spiniform setae
black; legs light brown; abdomen dark grayish brown; mandible as in figure 17 A; calcipala
and claw in accordance with Figs. 17 C, D; furcasternum as in figure 17 E; eighth sternite
with abundant setae (Fig. 17 F); paraproct short (Fig. 17 G); genital fork with wide
branches (Fig. 17 H).
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 85

A
B
I

M N

O P
F

v.p.

G L H

K
e.d.

Q
E
C
R
Figure 17. Cnesia dissimilis. A-H : Female; A: mandible; B: wing, general view; C: calcipala and
tarsomere I; D: claw; E: furcasternum; F: eighth sternite and gonapophyses, genital fork visible
through them and spermatheca; G: cercus and paraproct; H: genital fork. I-P: Male; I: anterior
portion of wing, showing chaetotaxy of costa, radial 1 and radial secondary veins; J: hind basitarsus;
K: gonocoxite, gonostylus ventral plate (v.p.) and endoparameres (e.d.), ventral view; L: gonocoxite
and gonostylus, dorsal view; M-N: ventral plate in different position; O-P: median sclerite; O:
sublateral aspect; P: dorsal view. Q-R: C. pusilla; Q: Female, genital fork; R: Male, hind basitarsus.
(A-P from WYGODZINSKY & COSCARÓN, 1973a).
86 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

C F B

H
L D

J E M

Figure 18. Cnesia dissimilis. A-E: Pupa; A: general view; B: gill; C: frontoclypeus of female; D:
abdomen chaetotaxy; ventral view; E: terminal portion of abdomen, ventral view. F-M: Larva; F:
cephalic apotome; G: antenna; H: hypostomium; I: cephalic capsule, ventral view; J: apex of
mandible and teeth; K: sclerite of thoracic proleg; L: gill histoblast; M: anal sclerite. (A-L from
WYGODZINSKY & COSCARÓN, 1973a).
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 87

Male: Wing length 3.4-4.5 mm; color as in female but a little darker; hind basitarsus
wide (Fig. 17 J); ratio of length/width = 3.2-3.4; genitalia as in figures 17 K-P.
Pupa: Cocoon light brown, loosely woven; covering all of body (Fig. 18 A); length of
body including gills about 5.0 mm; gill length 1.7 mm; head and thorax without
granulosity; gills with about 38 branches emerging from numerous primary branches
implanted on short basal trunk (Fig. 18 B); frontoclypeus with prefrontal and anterior
frontal trichomes spinelike (Fig. 18 C); ventral abdominal chaetotaxy as in figures 18 D, E.
Larva: Length 7.5-8.0 mm; head dark brown and body yellowish brown to dark
greenish; cephalic apotome and cervical sclerite as in figure 18 F; postgenal cleft not deep
(Fig. 18 I); antenna as in figure 18 G; ratio of proximal, medial and distal segments =
1:1.2:1.5; cephalic fan with about 40 rays; hypostomial teeth, mandibular border, postgenal
cleft, thoracic proleg sclerite, gill histoblast and anal sclerite in accordance with figures 18
H-M; thoracic proleg sclerite with about 44 teeth on comb; anal ring with 115-120 rows
with 25-28 hooks each.
Distribution: Cnesia dissimilis extends through all the Valdivian and Magellanic area
of southern Chile and penetrates into the Mediterranean climate area of central Chile
north to the province of Valparaiso. It is also very common east of the Andes in
Argentina, at least from Chubut to southern Neuquen.
Bionomics: The aquatic stages of this species are restricted to clear, large streams
and rivers with considerable current; they are never found in trickles, small streams,
irrigation ditches or any slow-flowing watercourse. The preferred water temperature
varies from 10 to 12 °C. Larvae are generally found attached to branches, stems, and roots
of terrestrial plants trailing in the streams; the plant involved is frequently a bamboo,
Chusquea culeou DESVAUX. Pupae are found where larvae occur, but occasionally also on the
undersurface of large stones or rocks resting on the river bottom. Larvae and pupae
generally occur in large aggregations, with cocoons often forming irregular, multilayered
masses, as in certain Prosimulium in the Northern Hemisphere. Cnesia dissimilis is a spring
species, with abundant larvae and pupae found as early as October. In January, larvae
become very rare, and most pupae encountered are empty. Dissection of freshly emerged
females shows the species to be anautogenous. The feeding habits are unknown, but to
judge from the structure of the claws of the female with their large basal process, Cnesia
dissimilis is probably ornithophilic.
Discussion: Although this widespread species is surprisingly uniform, some varia-
tion was observed. The wing length fluctuates between 3.2 and 4.3 mm; most specimens
have wings close to 4.0 mm. Females with wing length close to 3.0 mm were observed
occasionally in Chilean material. None of the Argentinean females examined had wings
shorter than 3.5 mm; these variations are paralleled by variations in the measurements of
pupae and larvae. The cephalopterothecal length of Argentinean pupae remains very
close to 2.5 mm, but fluctuates in a Chilean population from Cautín between 2.25 and
2.50 mm. Likewise, the head width of last-instar larvae of Argentinean material varies
from 0.70 to 0.75 mm, with 0.75 mm being the most frequent, but in the Chilean
population mentioned it extends from 0.65 to 0.75 mm, with most specimens ranging
between 0.65 and 0.70 mm. Some variation was also observed in the color of the thorax
of the adults. In males, the scutum varies from dark brown to black. The scutum of the
88 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

females, which is normally dull orange brown, may become dark brown in some
specimens, especially at the center of the disc, but it is never black.

Cnesia pusilla WYGODZINSKY & COSCARÓN (Figs. 17 Q, R.)


Cnesia pusilla WYGODZINSKY & COSCARÓN, 1973a: 159-161; COSCARÓN, 1991: 21-22; CROSSKEY &
HOWARD, 1997: 11.
Female: Wing length 2.9 mm; head blackish; scutum dull black with yellowish
pronotal area; metanotum dark grayish brown; frontal angle narrow, 50 degrees; hind
basitarsus 7 times as long as wide; eighth sternite, gonapophysis, paraprocts and cerci as in
Cnesia dissimilis; genital fork as in figure 17 Q; terminal plates with conspicuous areas of
pigment differences.
Male: Length of wing 3.0 mm; overall color of head and its appendages as in female;
lateral portion of thorax colored as in female; scutum velvety black; shoulders silvery gray;
scutellum, metanotum, wings, and legs as in female; abdomen gray; tergal plates velvety
black; basal fringe silvery gray; hind basitarsus wide (Fig. 17 R) 2.4 times as long as wide;
genitalia as in Cnesia dissimilis.
Pupa: Cephalopterothecal length 1.8-2.1 mm, in most specimens between 1.9 and
2.0 mm; head and thorax without tubercles; structure and onchotaxy as in Cnesia dissimilis.
Larva: General structure and antenna as in Cnesia dissimilis, but specimens smaller;
head width only 0.65-0.70 mm.
Distribution: Argentina: Neuquen, Rio Negro.
Bionomics: Larvae adhere to stones in creeks of moderate volume, about 0.5 m deep.
Discussion: Cnesia pusilla seems to be very close to C. dissimilis, and may not always
be easy to recognize. It differs from C. dissimilis mainly in being smaller in size in all stages,
the black scutum of the female, and morphologically by the comparatively small
membranous area at the insertion of the spermathecal duct on the spermatheca. Cnesia
pusilla can be distinguished from C. ornata, by its smaller overall size, the presence of small
hairs on the metanotum of the adults, the black scutum of the male, and the absence of
tubercles on the pupa.

Cnesia ornata WYGODZINSKY & COSCARÓN (Figs. 19 A-N.)


Cnesia ornata WYGODZINSKY & COSCARÓN, 1973a: 161; COSCARÓN, 1991: 22-23; CROSSKEY &
HOWARD, 1997: 11; 2004: 11.
Female: (Pharate) Color unknown; characters of head, eyes, antennae, mouthparts, and
legs with claws as in Cnesia dissimilis, as far as discernible; thorax and abdomen not examined.
Male: Length of wing 4.0 mm; scutum bright reddish brown with golden decumbent
hair; scutellum pale yellow brown with long golden hair; metanotum light reddish brown,
with golden pruinescence; pleura light brown; katepisternum slightly darker; wings with
veins pale and hair golden brown, including basal tuft; legs light yellowish brown, their
hair from golden to brown; coxae, bases and apices of femora and tibiae, and all tarsi,
darkened; abdomen grayish brown, except posterior border of terga and pleural regions,
which are grayish; hair of abdomen, including that of basal fringe, brass-colored.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 89

F
B

A G
C

D
E

J
L

H
K

M N
Figure 19. Cnesia ornata. A-G: Male; A: antenna; B: calcipala and tarsomere I; C-E: ventral plate in
different position; F-G: median sclerite in different position. H-J: pupa; H: gill; I: frontoclypeus and
trichomes; J: trichomes and arrangement of tubercles on thorax. K-N: Larva; K: general aspect,
dorsal view; L: antenna; M: anterior border of hypostomium; N: postgenal cleft and hypostomium.
(A-N from WYGODZINSKY & COSCARÓN, 1973a).
90 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Antenna as in figure 19 A; last segment of maxillary palp less than twice as long as
penultimate segment; sensory vesicle slightly smaller than one-third diameter of third
palpomere; hind basitarsus 3.8 times as long as wide; calcipala as in figure 19 B; genitalia
much as in Cnesia dissimilis; ventral plate and median sclerite as in figures 19 C-G.
Pupa: Cocoon light brown, covering pupa almost completely, exposing only gills and
part of dorsal surface of thorax; cocoon with filaments of varying diameter, easily
perceptible; cocoon often covered with foreign organic matter; length of cocoon dorsally
along middle, 3.5 mm; length along base, 4.5 mm; length of body of pupa, 3.5-4.0 mm;
cephalopterothecal length 2.4-2.5 mm; maximum length of gill, 1.5 mm; head and thorax
of pupa light brown; gills (Fig. 19 H) very similar to those of Cnesia dissimilis; number of
filaments about 35, arising from about 12 primary branches inserted on very short
common trunk; some primary branches divided twice, thus giving rise to three filaments;
surface of filaments minutely granulose; their apex rounded; chaetotaxy of head much as
in C. dissimilis, but prefrontal and frontal anterior trichomes capitate; frontoclypeus with
tubercles arranged as in figure 19 I; chaetotaxy of thorax as in C. dissimilis, but strong
trichomes capitate (Fig. 19 J); exposed dorsal surface of thorax with numerous smooth
tubercles arranged in circles; abdomen with general onchotaxy as in C. dissimilis; stout
setae of tergum II blunt apically; tubercles of terga more numerous than in C. dissimilis.
Larva: Maximum length 6.5-7.2 mm; color of body light yellowish gray-brown; head
dark brown; body shape as in figure 19 K, with wider subapical area; cephalic apotome
brownish with negative spots; hypostomium with about 15-20 teeth (Fig. 19 M); sublateral
setae in number of 5-7 on each side; lateral serrations absent; cervical sclerites reduced
and isolated from postocciput sides; antennal shape as in figure 19 L; ratio of proximal,
medial and distal antennal segments = 1:1.0-1.7:1.2-1.6; postgenal cleft with small
rounded concavity (Fig. 19 N); head fan with 38 rays; mandible and proleg sclerite very
similar to those of Cnesia dissimilis; ninth segment without ventral tubercles; anal ring with
144-152 rows of 25-30 hooks each; anal integument without hair.
Distribution: Argentina: Rio Negro; Chile: Chillán, Curacautín, Talca.
Bionomics: Larvae live in torrential creeks, over “caña colihue” (Chusquea culeou).
Discussion: Cnesia ornata differs from the other two species of the genus mainly by
its bright reddish brown scutum in the male, the absence of setae on the metanotum, and
the presence of tubercles and capitate trichomes on the head and thorax of the pupa.

Gigantodax ENDERLEIN (Figs. 20 B, C, E, G, H, J, L, N, P, R, S.)


Gigantodax ENDERLEIN, 1925: 205.
Type-species: Gigantodax bolivianus ENDERLEIN, 1925: 205-206.
Archicnesia ENDERLEIN, 1934b: 273; Type-species Simulium (Gigantodax) femineum EDWARDS, 1931: 135.
Diagnosis: Antennae with 11 segments; scutum with recumbent hair; mesepistenal
sulcus shallow, evanescent anteriorly; katepisternum and pleural membrane glabrous; arms
of furcasternum without projections (Fig. 20 H); wing with basal cell not developed; basal
portion of R with hair arranged in several irregular series, R1 dorsally with or rarely without
spinelike setae and fine hair, Rs not forked apically, Cu-A2 straight (Fig. 20 C); calcipala well
developed (Fig. 20 E); female claws with large subbasal tooth of varied shapes (Fig. 20 G);
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 91

genital fork distinctly pigmented, lateral arms with 1+1 forwardly directed well-developed
sclerotized apodemes (Fig. 20 J); male gonostylus more than half as short as gonocoxite with
2-3 apical spines, ventral plate elongated with long basal arms connected by wide membrane,
endoparemeres scarcely developed basally and teeth reduced to a group of 1-4 medium-
sized denticles isolated on aedeagal membrane (Fig. 20 L).
Pupa: Cocoon generally large; shapeless and covering most of pupal body, which is
well sclerotized; frontal trichomes absent (Fig. 20 N); cephalic and thoracic trichomes
spinelike; gills of varied structure, from filamentous to arborescent, tubular or globular;
terminal spines thin and stout.
Larva: Cephalic apotome with positive spots; cervical sclerite generally transverse
and connected to upper ends of postocciput; postgenal cleft not developed; mandible
with three external teeth, preapical teeth small, marginal serrations with one or two large
teeth followed by several minute serrations (Fig. 20 P); hypostomial intermediate teeth
shorter than lateral tooth (Fig. 20 R); larval cuticle generally bare; anal sclerite with
posterior arms prolonged forming a complete ring and 1+1 backwardly directed struts
underlying the X-shaped portion (Fig. 20 S); rectal papillae with 3 single lobes; abdominal
segment IX with pair of ventral tubercles.
Distribution: Cool and temperate western South America, high lands of Mesoamer-
ica and one species from southwestern North America.
Discussion: Gigantodax and Pedrowygomyia are differentiated from all the known
Neotropical black flies by the combination of the straight Cu, An2 and structure of the
larval mandible, and also by the presence in larvae of a complete anal ring and backwardly
directed struts, although this anal ring is partially present in some Simulium (Hearlea)
species. Gigantodax is very close to Cnesia, as was shown by COSCARÓN & MIRANDA
ESQUIVEL (1998a) and reinforced by chromosomal similarity (COSCARÓN ARIAS, 1998); an
interesting cladistic revision of the Gigantodax species group, using morphological and
chromosomal information is presented by PINTO-SANCHEZ et al.2005. Gigantodax contains
65 species; 60 were treated in the revision of WYGODZINSKY & COSCARÓN (1989); the other
species of Gigantodax are G. adleri MOULTON, 1996; G. bierigi VARGAS & RAMÍREZ PÉREZ,
1988; G. conviti RAMÍREZ PÉREZ, 1980a; G. lazoi TAKAOKA et al., 1988 and G. willei VARGAS &
RAMÍREZ PÉREZ, 1988.

Pedrowygomyia COSCARÓN & MIRANDA ESQUIVEL (Figs. 20 A, D, F, I, K, M, O, Q.)


Pedrowygomyia COSCARÓN & MIRANDA ESQUIVEL, 1998a: 164.
Type-species: Gigantodax cortesi WYGODZINSKY & COSCARÓN, 1989: 22-27.
Diagnosis: Pedrowygomyia has a close relationship with Gigantodax and the species
were included by WYGODZINSKY & COSCARÓN (1989) in that genus as Gigantodax CORTESI
species group; COSCARÓN & MIRANDA ESQUIVEL (1998) through cladistic studies showed
the monophyly of Gigantodax species treated by the former authors, but considered that
the Gigantodax CORTESI species group has enough characters to designate it as an
independent genus. Here we consider only the characters that differ from Gigantodax.
Adults: Frons and clypeus relatively wide, frontal median sulcus scarcely evident
(Fig. 20 A); basal flagellomeres and palpomeres with long hair; scape and pedicel globose
92 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

E F

A B H

G
J

I M

K
N

P
Q

R
O

Figure 20. Gigantodax and Pedrowygomyia. A-B: head, frontal view; A: Pedrowygomyia cortesi, B: Gigantodax
brophyi. C: wing, G. dryadicaudicis; D-E: apex of basitarsus and tarsomere 1; D: P. punapi, E: G. igniculus. F-
G: claws; F: P. cortesi, G: G. igniculus. H: furcasternum, G. antarctiucs. I-J: genital fork; I: P. cortesi; J: G.
rufidulus. K-L: male genitalia, ventral view, K: P. jatunchuspi, showing gonocoxite, gonostylus, ventral plate
and median sclerite; L: G. igniculus, showing gonocoxite, gonostylus, ventral plate, median sclerite,
aedeagus membrane and teeth of endoparameres. M-N: Pupa, frontoclypeus; M: P. jatunchuspi; N: G.
marginalis. O-S: Larva; O: P. jatunchuspi, head dorsal view; P: G. igniculus, mandible teeth; Q-R: hypostomial
teeth, anterior margin; Q: P. cortesi, R: G. minor; S: anal sclerite, with scales and lateral struts, G. igniculus.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 93

and first flagellomere elongate, female mandible and maxilla obsolescent; scutum with
erect and recumbent hair; R1 without spines but with numerous placoid sensilla; calcipala
absent (Fig. 20 D) and claws without transverse sulcus and very short subbasal tooth (Fig.
20 F); genital fork with anterior branch wide basally, becoming thin toward apex (Fig. 20
I); male with hair on interocular space, scutum relatively flattened, ventral plate
comparatively elongated (Fig. 20 K) and lateral margins at midlength convex, endoparam-
eral teeth very short.
Pupa: Frontoclypeus with frontal and facial trichomes and clypeus comparatively
thinner (Fig. 20 M).
Larva: Large body size, up to 9.6 mm; cephalic apotome widest near base and with
basal spots (Fig. 20 O); hypostomium with first intermediate tooth longer than lateral
tooth; body cuticle with small single setae implanted in dark spot.
Distribution: Pedrowygomyia is restricted to the high plateau (Puna) of the Andes
from northern Chile to northwestern Argentina and central Peru. This genus contains
four species: Pedrowygomyia chacabamba, P. cortesi, P. jatunchuspi and P. punapi. Detailed
descriptions of these species can be found in WYGODZINSKY & COSCARÓN (1989).

Cnesiamima WYGODZINSKY & COSCARÓN (Figs. 21, 136.)


Cnesiamima WYGODZINSKY & COSCARÓN, 1973a: 184.
Type-Species: Simulium (Cnephia) atroparvum EDWARDS, 1931: 148.
Adults: Antennae with 11 segments; male holoptic; female with frons relatively wide
(Fig. 21 A); mandibles of female with well-developed denticles on both margins (Fig. 21 C);
apical segment of maxillary palp of both sexes less than twice as long as penultimate
segment (Fig. 21 B); mesepisternal sulcus shallow, although deeper than in Cnesia, somewhat
evanescent anteriorly; katepisternum glabrous, in profile slightly wider than high; pleural
membrane bare; metanotum glabrous; arms of furcasternum without projections (Fig. 21
G); wings with basal cell distinct (Fig. 21 E); Sc with numerous hairs arranged in single row;
basal section of R with hair arranged in several rows (Fig. 21 F); R1 with hair only, spiniform
setae absent, Rs entirely hairy below, hairy above on its apical half; Rs not forked apically; M
appearing thickened on most of its length; Cu2 rather weakly curved in female, more
strongly so in male; A2 almost attaining wing margin; calcipala present, large, bluntly
rounded apically (Fig. 21 O); claws of female with large basal tooth (Fig. 21 J); spermatheca
of normal size, about as large as gonapophyses; globular, its internal surface faintly wrinkled
and with few scattered spicules; whole spermatheca and base of spermathecal duct
sclerotized; sclerotized portion gradually fading into unpigmented portion; gonapophyses
widely subtriangular, somewhat transverse, their medial borders slightly pigmented, approx-
imated; their surface with microtrichia and few setae (Fig. 21 L); genital fork distinctly
pigmented, with stem slender, longer than arms, latter wide, lacking forwardly directed
apodemes; male with gonostylus slightly shorter than gonocoxite (Figs. 21 P, Q); gonocoxite
with semicircular membranous area opposite inner basal angle of gonostylus, the latter with
corresponding bulge at base; gonostylus only slightly narrowed toward apex, with two distal
spines (Figs. 21 Q, R); medial surface of gonostylus smooth, delimited by two carinae;
ventral plate subtriangular, about as log as wide, setose, with low median longitudinal keel
94 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

(Figs. 21 S, T); median sclerite geniculate in side view (Fig. 21 U), roughly M-shaped in full
view, with the inner arms of the M partly fused and connected to basal flange (Fig. 21 V);
endoparameres reduced to subquadrate plates connected to arms of ventral plate (Fig. 21
U); denticles not developed.
Pupa: Cocoon well defined, covering pupa except anterior extremity and gills; shape
broadly oval, flattened, shield like; borders of anterior opening slightly but distinctly
reinforced, especially at center, somewhat salient at middle (Fig. 22 A-B); cocoon without
ventral layer; gills cigar-shaped, inserted perpendicularly on extremely short trunk at
posterior third of body of gill, the gill projected forward and downward and somewhat
laterally; gills about as long as maximum width of body of pupa, with several irregularly
transverse septa; surface of gills rugose-reticulate; gills bearing delicate scattered respira-
tory filaments extending perpendicularly to longitudinal axis of gill, length of filaments
over twice that of diameter of gill, their surface minutely pitted, their apex slightly clavate
(Fig. 22 C); head only with 1+1 hair-like facial trichomes (Fig. 22 D); disc of thorax with
5-6+5-6 trichomes, the three dorsal ones in shape of apically bent strong spine-like setae,
the others hair like; abdomen pointed posteriorly; distinct pleural areas not developed;
onchotaxy of abdomen as follows: tergum I glabrous; tergum II with several fine hairs,
1+1 submedian ones slender, spinelike; terga III and IV with 3+3 short hooks posteriorly;
terga V-VIII with spine-combs; terminal spines short, sharply pointed; sternum V with
2+2 elongate spine-like hooks; sternal plates VI and VII with 1+1 similar hooks; 1+1
hooks also in pleural membrane of segments VI and VII; pleural area of terminal
segments with several extremely complex grapnel-shaped setae (Fig. 22 E).
Larva: Abdomen relatively elongated, with 1+1 ventral tubercles (Fig. 22 F); cephalic
apotome widened before posterior border; cervical sclerites of reduced size and isolated
from postocciput margin (Fig. 22 G); postgenal cleft obsolescent (Fig. 22 H); antenna
surpassing base of cephalic fan stem; distal antennal segment very thin and more than
twice length of basal segments and medial segment very short (Figs. 22 I, J); palp
elongated; rays of cephalic fan very thin showing larger setae alternately; mandible with
scarce marginal teeth (Figs. 22 K, L); hypostomium with anterior border surpassing the
teeth except the angle tooth (Fig. 22 M); thoracic proleg subquadrate; anal sclerite X-
shaped, without or only faint rods (Fig. 22 N); body cuticle nude.
Distribution: Valdivian and Magellanic areas of southern Chile, and western Patagonia.
Discussion: The adults of Cnesiamima are superficially very similar to those of
Paraustrosimulium, as already stated by EDWARDS (1931); this similarity extends to the
general shape of the median sclerite of the male genitalia, but the structure of the
endoparameres and of the female genitalia as found in the two genera is quite different,
and does not support a hypothesis of close relationship. The following enumeration
summarizes the significant differences between the adults of Cnesiamima atroparva and
Paraustrosimulium anthracinum. In regard to the remaining South American Prosimuliini, the
adults of Cnesiamima can be distinguished from the Chilean-Patagonian Cnesia, Arauc-
nephia, and Araucnephioides by, among other characters, the absence of spine-like setae on
R1, the sclerotized neck of the spermatheca, and certain features of the male genitalia,
such as the bulge at the base of the distimere, the absence of denticles on the
endoparameres, and the M-shaped median sclerite. Cnesiamima can be further distin-
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 95

E
B C

D
K
F
G

H
I J

O N

P
Q

R
M

T U
S V
Figure 21. Cnesiamima atroparva. A-M: Female; A: frons; B: maxillary palp; C: mandible; D: fronto-
ocular triangle; E: wing, general view; F: portion of wing with C and R1; G: furcasternum; H: sensory
vesicle of maxillary palp; I: hind leg; J: claw; K: genital fork; L: eighth sternite and gonapophyses,
genital fork and spermatheca; M: cercus and paraproct: right, ventral view, left mounted on slide. N-
V: Male; N: hind leg; O: calcipala and tarsomere I; P: gonocoxite, gonostylus, ventral plate and median
sclerite in ventral view; Q: gonocoxite and gonostylus, dorsal view; R: gonostylus, showing inner
membranous surface; S-T: ventral plate in different position; U: ventral plate, median sclerite and plate
of endoparameral organ; V: median sclerite. (A-V from WYGODZINSKY & COSCARÓN, 1973a).
96 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

guished from Araucnephia and Araucnephioides by A2 which almost attains the wing margin,
and from the latter genus also by the larger basal tooth of the claws of the female. The
differences between Cnesiamima and Gigantodax are obvious in wing venation and the anal
sclerite of the larva. Lutzsimulium differs by the absence of spiniform setae on R1, the
large tooth of the claw of the female, and by the peculiarities of the male and female
genitalia. Cnesiamima differs from Mayacnephia by the absence of spiniform setae on R1, the
katepisternum which is wider than high, the sclerotized neck of the spermatheca, and by
the quite different structure of the median sclerite of the male. Cnesiamima shares with
Tlalocomyia a spermatheca with a sclerotized neck, although there are differences in detail,
but the apical segment of the maxillary palp of Tlalocomyia is very long in both sexes, the
pleural membrane is hairy on its upper portion, spiniform setae are on R1, and the median
sclerite of the male is quite different.

Cnesiamima atroparva (EDWARDS) (Figs. 21 A-V; 22 A-O.)


Simulium (Cnephia) atroparvum EDWARDS, 1931: 148.
Cnesiamima atroparva WYGODZINSKY & COSCARÓN, 1973a: 187-189; 1973b: 5; COSCARÓN, 1985: 67;
1991: 74-75; CROSSKEY & HOWARD, 1997: 12; 2004: 3.
Cnephia atroparvum SMART, 1945: 498.
Cnesia atroparvum VARGAS & DÍAZ NÁJERA, 1951: 165.
Simulium atroparvum VARGAS, 1945a: 116.
Simulium (Astega) atroparvum RUBTSOV, 1940: 532.
Female: Wing length 3.1-3.5 mm; color of head and its appendages black; eyes,
antenna, palp, proboscis, and occiput dark grayish brown; frons and clypeus gray
pollinose; pilosity of antenna very short, that of palp, clypeus, frons, and occiput
somewhat longer; hair from brass-colored to black; scape and pedicel slightly lighter than
flagellum, and with longer hair; scutum dark gray, with silvery to nacreous pruinosity;
shoulders reddish; appressed setae of scutum brass-colored; scutellum reddish brown, its
hair from brass-colored to dark brown; metanotum black, silvery pollinose; wings hyaline,
veins medium brown; spines and hair from dark brown to black; hair on wing base from
yellowish to piceous; halteres from yellowish to pale brown; pleural tuft brass-colored;
pleura dark grayish brown, silvery pollinose; legs rather uniformly light brown to piceous,
their hair from brass-colored to black; abdomen brownish or grayish, tergal plates darker;
membranous areas light grayish brown; hair of abdomen, including basal fringe, brass-
colored; frons (Fig. 21 A) medium wide, frontal angle 75 degrees; fronto-ocular triangle
distinctly deeper than wide (Fig. 21 D); maxillary palp as shown in figure 21 B; third
segment somewhat swollen; structure of sensory vesicle as shown in figure 21 H, its
diameter about half that of third palpomere; lacinia with 25-28 denticles, mandible with 37-
41 denticles (Fig. 21 C); structure of wing as in generic description; shape and proportion of
segments of hind leg as shown in figure 21 I; calcipala shorter than width at base; claws as
shown in figure 21 J; tergal plates of abdomen wider than long, except on terga II-IV where
they are as wide as long; ventral plates not developed; eighth sternum with 1+1 groups of
16-20 long setae; disc of eighth sternite uniformly pigmented except darker subsemicircular
band anteriorly (Fig. 21 L); paraprocts short, rounded distally (Fig. 21 M).
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 97

Male: Wing length 2.9-3.2 mm; head black; eyes red in live specimens, mostly
black in dried material; antennae, palp, and clypeus black; occiput dark gray pollinose;
proboscis dark grayish brown; antennae with short silvery to black pubescence; scape
and pedicel, palpi, clypeus and occiput with long brown or black hair; scutum velvety
black, scutellum dark brown, with brown to black hair; metanotum black, with silvery
pruinosity; color of wings, legs, pleura, and pleural tuft as in female; abdomen velvety
black dorsally, grayish at sides and below; hair of abdomen from brass-colored to black;
third palpomere more slender than in female; sensory vesicle only slightly smaller than
in female; wings as in female, but Cu2 with stronger curvature; shape and proportions
of segments of hind leg as shown in figure 21 N; hind basitarsus 3.5-3.9 times as long as
wide; calcipala slightly shorter than width at base (Fig. 21 O); gonocoxite subtrapezoidal
(Figs. 21 P-Q), longer than wide, with membranous area apically on inner surface
projecting in some cases; gonostylus elongate, subcylindrical (Figs. 21 P-R); ventral
plate as shown in figures 21 S, T, U; lateral projections and basal margin conspicuously
sclerotized; median sclerite as shown in figures 21 V-U Endoparameral organ as in
generic description and partially in figure 21 U.
Pupa: Cocoon grayish brown, with distinct threads; shape as in figures 22 A-B;
length of cocoon 3.3-4.0 mm, its maximum width 2.7 mm; total length of pupa (gills
not included) 2.7-2.9 mm; tubercles weakly developed, perceptible only on thorax, even
there ill defined; gill length 1.2-1.4, its maximum width 0.25-0.30 mm; gills each with
13-17 septa, and with about 35 rather regularly spaced respiratory filaments; filaments
inserted on spaces between septa (Fig. 22 C); onchotaxy of head, thorax, and abdomen
as in generic description and figures 22 D-E; spine-combs of posterior terga with 12-22
widely spaced teeth.
Larva: Maximum length 6.0-6.5 mm; cephalic capsule width, 0.7 mm; general color
light grayish green, head light yellowish brown; shape as in figure 22 F; cephalic apotome
with positive dark brown spots, and darkened on posterior margin (Fig. 22 G); cephalic
fan with unknown number of rays; ratio of proximal, medial and distal antennal segments
= 1: 0.16-0.27:2.5-3.4; shape as in figures 22 I, J; mandible with 10-12 internal teeth and 3-
5 marginal denticles (Figs. 22 K, L); hypostomium with median tooth lower than lateral
teeth (Fig. 22 M); 3-5 lateral setae on each side (Fig. 22 H); thoracic proleg lateral sclerite
with 14-17 teeth; anal sclerite with hair, not scales (Fig. 22 N); anal ring with 75-80 rows of
13-15 hooks each; pupa gill histoblast as in figure 22 O.
Distribution: Argentina: Neuquén, Río Negro, Tierra del Fuego; Chile: Curicó,
Llanquihue, Magallanes, Talca, Valdivia.
Bionomics: The pupae were collected in crystal-clear, small mountain streams at
water temperatures of 0-6 °C. The larvae were found in Tierra del Fuego in a creek
covered by ice and snow; pupae were found singly, on the upper surface of stones or on
leaves of shrubs trailing in the water.
Discussion: Cnesiamima has a great similarity with Paraustrosimulium in characters of
male genitalia, pupa and larva. They are easy to separate by the pupal gills that are quite
different superficially, those of Paraustrosimulium being lamellate and glabrous and those
of Cnesiamima cigar-shaped and beset with numerous respiratory filaments.
98 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

C
E
D

G J

N
I

M O

Figure 22. Cnesiamima atroparva. A-E: Pupa; A-B: general aspect in dorsal and lateral views. C: gill.
D: frontoclypeus and trichome. E: apical portion of abdomen, ventral view, with trichomes and
terminal spur. F-O: Larva; F: general aspect, lateral view; G: cephalic apotome; H: postgenal cleft
and hypostomium; I: antenna, general view; J: portion of antenna showing medial and part of
proximal and distal articles; K: apical portion and teeth of mandible; L: marginal serrations of
mandible; M: anterior portion of hypostomium; N: anal sclerite and hair; O: gill histoblast. (A-E
from WYGODZINSKY & COSCARÓN, 1973b).
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 99

Paraustrosimulium WYGODZINSKY & COSCARÓN (Figs. 23, 136.)


Type-species: Simulium anthracinum BIGOT, 1888.
Austrosimulium (Paraustrosimulium) WYGODZINSKY & COSCARÓN, 1962: 244; STONE, 1963: 14.
Paraustrosimulium CROSSKEY, 1969: 17.
Adults: Antenna with 10 segments; male holoptic; female with frons relatively narrow
(frontal angle about 65 degrees); mandibles of female with well-developed denticles on both
sides; apical segment of maxillary palp of both sexes less than twice as long as penultimate
segment; mesepisternal sulcus deep, well defined anteriorly; katepisternum glabrous, in profile
much wider than high; metanotum glabrous; arms of furcasternum with conspicuous projections
(Fig. 23 A); pleural membrane bare; wings with basal cell distinct; Sc with numerous hairs
arranged in single row; basal section of R with hair arranged in several rows; R1 with hair and very
slender spine-like setae arranged in one or two irregular rows; Rs entirely hairy below, hair
arranged in single row at base and apex of vein, in several rows on central portion of vein; apical
portion of Rs dorsally on apical half with one series of hairs; Rs not forked apically; M appearing
thickened on most of its length; curvature of Cu2 normal, somewhat more pronounced in male
than in female; A2 falling distinctly short of wing margin; calcipala present, large, rounded apically;
claws of female with medium-sized subtriangular basal tooth (Fig. 23 B); spermatheca larger than
usual, larger than gonapophysis; its internal surface smooth, with very small number of scattered
spiculae; spermatheca fully sclerotized, except very small subcircular membranous area at
insertion of spermathecal duct; paraproct short with truncate apical border (Fig. 23 C);
gonapophyses widely subtriangular, somewhat transverse, and closely approximated, their surface
with microtrichia and few very short setae, medial border slightly pigmented (Fig. 23 D); genital
fork lacking conspicuous pigment; stem very short and wide, not longer than arms, the latter also
very wide; forwardly directed apodemes absent (Fig. 23 E); male with gonostylus slightly shorter
than gonocoxite, subtriangular, with two to four spines; medial surface of gonostylus smooth,
delimited by two carinae (Fig. 23 F); ventral plate subpentagonal, with distinct median carina (Fig.
23 G); median sclerite distinct, geniculate in lateral view; basal portion narrow proximally,
somewhat widened and deeply incised distally; apical portion composed of two simple divergent
arms (Fig. 23 H); endoparameres composed of distinct free plate-like basal portion and group of
elongate parallel denticles not connected by sclerotized area to plate-like portion.
Pupa: Cocoon well defined, wall-pocket shaped, viz., without anteroventral collar;
undersurface of cocoon reduced to few irregular strands near base of cocoon; body of pupa
distinctly sclerotized throughout, although not strongly so; respiratory organs in shape of twisted
pseudo-segmented lamella (Fig. 23 I); head sclerite with frontal and facial trichomes, all hair like;
disc of thorax with approximately 5+5 hair like trichomes, furthermore with 3+3 spine-like
trichomes dorsally near midline; abdomen pointed posteriorly; distinct pleural areas not
developed; terga III and IV with 4+4 hooks posteriorly; terga V to IX with continuous spine-
combs anteriorly; sternum V with 4+4 hooks posteriorly; sternal plates VI and VII with 1+1
hooks, and 1+1 hooks each also in pleural membrane of segments VI and VII; terminal spines in
shape of short, pointed tubercle; posterior segments with several grapnel-shaped setae (Fig. 23 J).
Larva: Head spots positive; cephalic apotome widest slightly before hind margin; cervical
sclerites composed of small subcircular sclerotized area and transverse less strongly sclerotized
area, the latter not fused to upper ends of postocciput (Fig. 23 K); postgenal cleft not developed;
100 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

B
D

C
J
H

L K

Figure 23. Paraustrosimulium anthracinum. A-E: Female; A: furcasternum; B: claw; C: cercus and
paraproct; D: eighth sternite and gonapophyses; E: genital fork. F-H: Male; F: gonocoxite and
gonostylus, dorsal view; G: ventral plate; H: aedeagus with median sclerite, endoparameres and ventral
plate visible through it. I-J: Pupa; I: general aspect dorsal view; J: apex of abdomen with trichomes,
spines and terminal spurs. K-N: Larva; K: cephalic apotome; L: antenna; M: hypostomium; N: anal
sclerite, with lateral rods, hair and spines. (A-N from WYGODZINSKY & COSCARÓN, 1973a).
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 101

hypostomium much shorter than postgenal bridge; antennae slightly longer than stem of cephalic
fans; their two basal segments very dark, the apical two very light-colored (Fig. 23 L); cephalic fans
with rays normal; mandibles with two external and one apical tooth; first and second preapical
teeth of identical size, rather wide; third preapical tooth larger than others, but much smaller than
apical tooth, and indistinguishable from internal teeth; the latter arranged in single row; one large
mandibular serration followed by from two to four smaller ones; short basal setae of mandible
very slender, glabrous; hypostomium with 13 teeth, not arranged in conspicuous groups, median
tooth and lateral teeth slightly larger than others; the 2+2 lateral teeth inserted almost at same
level as remaining teeth (Fig. 23 M); teeth of hypostomium underlain by lamellar extension of
hypostomium; lateral sclerite of thoracic proleg subtriangular, slightly wider than long rounded at
base, strongly widened apically; setae of lateral sclerite shorter than sclerite, arranged in two or
three irregular rows, inserted individually on barely perceptible tubercles; abdominal cuticle bare,
except for conspicuous perianal scales; anal sclerite X-shaped; 1+1 backwardly directed struts
under lying main body of sclerite (Fig. 23 N).
Distribution: Valdivian and Magellanic areas of southern Chile, and western Pata-
gonia (Fig. 136).
Bionomics: SILVA FIGUEROA (1917) mentioned that P. anthracinum (here Simulium moorei)
bites humans on the Taitao Peninsula of Chile, and can become very bothersome. EDWARDS
(1931) examined a specimen that “bit, causing painful swelling of hand.” We have not
recorded anthropophilic behaviour in this species. The aquatic instars are found in clear,
cold, small and large streams. Pupae and larvae were found attached to rocks, leaves, pieces
of wood, and dead branches trailing in the stream, invariably close to the surface.
Discussion: Paraustrosimulium presents an array of autapomorphic characters that
distinguish it from all other Western Hemisphere Simuliini genera, i.e., the 10-segmented
antennae, the peculiar endoparameral organ and the almost unpigmented genital fork with its
short and wide stem; the well-formed cocoon and the unique gills of the pupa. The extremely
elongate third antennal segment in the larva are shared with Cnesiamima; the backwardly
directed struts underlying the anal sclerite are also found in Gigantodax. Paraustrosimulium shares
with Lutzsimulium and Simulium the presence of conspicuous projections on the arms of the
furcasternum, an apomorphic character within the framework of the family.

Paraustrosimulium anthracinum (BIGOT) (Fig. 23.)


Simulium anthracinum BIGOT, 1888: 15.
Simulium (Austrosimulium) anthracinum EDWARDS, 1931: 143.
Austrosimulium anthracinum SMART, 1945: 499; WYGODZINSKY, 1953a: 293; DUMBLETON, 1960: 543.
Austrosimulium (Paraustrosimulium) anthracinum WYGODZINSKY & COSCARÓN, 1962: 242; COSCARÓN, 1968: 66.
Paraustrosimulium anthracinum CROSSKEY, 1969: 73; WYGODZINSKY & COSCARÓN, 1973a: 191-192;
COSCARÓN 1991: 76-78.
Simulium moorei SILVA FIGUEROA, 1917: 30.
Simulium (Austrosimulium) moorei EDWARDS, 1931: 144.
Austrosimulium moorei SMART, 1945: 499.
Female: Wing length 2.8-3.3 mm; general coloration blackish; scutum black; scutellum
grayish-brown, metanotum black with silvery pollinosity; hyaline wing with brown veins,
102 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

with hair and setae black; legs dark brown, abdomen grayish brown; palpal sensory organ
about 1/3 segment length; length/width ratio of hind basitarsus = 6.1; calcipala about 1/3
tarsomere I length; furcasternum (Fig. A), claw (Fig. B) and genitalia (Figs. C-E) as in generic
description.
Male: Wing length 3.0 mm; coloration as for female; palpal sensory organ about half
of segment length; length/width ratio of hind basitarsus = 4.5-4.8; calcipala about ¼
tarsomere length; abdominal tergal plates large, sternal plates small; genitalia (Figs. F-H)
as in generic description.
Pupa: Cocoon color light yellowish brown, weave compact; reinforced on anterior
margin, covering most of body but leaving gill free (Fig. I); length on dorsum 3.2-3.5 mm, on
base 3.5-4.4 mm; gill length 2.0-2.2 mm; gill foliaceus, annulated with 2 branches basally,
internal larger than external; cephalothorax with abundant small tubercles; chaetotaxy as in
generic description; thorax trichomes very strong and terminal spines very short (Fig. J).
Larva: Maximum length 6.5 mm; general coloration light grayish yellow; cephalic
apotome darkened basally and medially, with positive spots (Fig. K); antennal shape as in
figure L; ratio of proximal, medial and distal antennal segments = 1:0.8-0.9:2.1-2.2;
hypostomial teeth shorter than frontal margin of hypostomium; with 5 or 6 lateral setae
on each side (Fig. M); sclerite comb of pseudopod with about 33 teeth; rectal papillae with
70 rows of 15-18 hooks each; anal sclerite with abundant small scales (Fig. N); rectal
papillae with three single lobes.
Distribution: Neantarctic biogeographic region (Central Chile plus subAntarctic
realms); Argentina: Chubut, Neuquén, Tierra del Fuego; Chile: Aysen, Chiloe, Llan-
quihue, Magallanes, Malleco, Osorno, Valdivia (Fig. 136).
Bionomics: It lives on vegetation in creeks and small rivers, not too torrential.

Lutzsimulium D’ANDRETTA & D’ANDRETTA (Figs. 24, 25, 136.)


Lutzsimulium D’ANDRETTA & D’ANDRETTA, 1947: 402.
Type-Species: Simulium hirticosta LUTZ, 1909: 135 (as Lutzsimulium cruzi D’ANDRETTA & D’ANDRETTA).
Adults: Antennae 11-segmented; male holoptic; female with frons comparatively wide
(Fig. 24 A); frontal angle about 90 degrees; mandibles of female toothed only on inner
margin; apical segment of maxillary palp of both sexes only very slightly longer than
penultimate segment; mesepisternal sulcus rather deep, not evanescent anteriorly; katepister-
num glabrous, in profile about as high as wide; arms of furcasternum with conspicuous
projections (Fig. 24 L); metanotum glabrous; wings without basal cell; R1 with spine-like
setae (Fig. 24 H); Sc and basal section of R setose; Rs entirely hairy below, setose above on
distal half; Rs not forked apically; M appearing duplicated for most of its length; Cu2
conspicuously curved, in some specimens with rudimentary branch submedially; A2 ending
well before wing margin; calcipala large, widely rounded apically (Fig. 24 C); hind basitarsus
of male not strongly widened; claws of female with one very small, pointed, subbasal tooth
(Fig. 24 D); gonapophyses large, transversely subtriangular, their apex bluntly rounded;
medial borders approximated, slightly pigmented (Fig. 24 E); surface of gonapophysis with
microtrichia and few short setae; genital fork distinctly pigmented; stem slender, only slightly
longer than arms, the latter wide, without apodemes (Fig. 24 F); spermatheca of normal size,
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 103

fully sclerotized, not reticulate on either surface; internal surface with scattered spiculae;
subcircular area at insertion of spermathecal duct with peculiar network-like structure
extending to base of duct, there gradually fading (Fig. 24 G); male with gonostylus slightly
shorter than gonocoxite, subconical, with from two to four apical spines (Figs. 24 I, M);
ventral plate subquadrate or subrectangular, with conspicuous keel (Fig. 24 J); median
sclerite somewhat S-shaped in lateral view; in frontal view with proximal portion entire, wide
basally, narrowed distally (Fig. 24 J); apical portion in shape of two short, diverging arms
connected by membrane; short ends of arms free, backwardly curved; endoparameres
scarcely perceptible, reduced to basal plates (Fig. 24 I).
Pupa: Cocoon without definite shape, covering abdomen and only part of thorax of
pupa; body of pupa well sclerotized throughout (Fig. 25 A); head and thorax with or without
tubercles; many trichomes of head and thorax and setae of terminal abdominal segments
tightly looped apically (Fig. 25 E); gills arborescent, divided into primary and secondary
branches close to base; general structure of branches filamentous (Figs. 25 B, C); head
sclerite with 1+1 facial trichomes; frontal trichomes present or absent; thorax with 4+4 or
5+5 trichomes; abdomen pointed posteriorly; distinct pleural areas not developed; onchot-
axy: terga III and IV with 4+4 hooks posteriorly; terga VI-IX with continuous spine-combs
anteriorly; sternum IV without or with 1+1, V with 2+2 hooks posteriorly; sternal plates VI
and VII with 1+1 hooks; 1+1 hooks each also in pleural membranes of segments VI and
VII; terminal spines well developed, long and slender, strongly curved at base (Fig. 25 E);
setae of segments VIII and IX strong, looped apically, never branched.
Larva: Head spots positive; cephalic apotome widest well before hind margin (Fig.
25 G); cervical sclerites either wide, fused to upper end of postocciput, or small and free;
postgenal cleft large (Fig. 25 H); antennae as long or slightly shorter than stem of cephalic
fan (Fig. 25 G); third segment distinctly darker than preceding; cephalic fans normal with
large setae alternating among shorter setae; mandibles with two outer teeth in addition to
apical tooth; second preapical tooth much smaller than either first or third; third smaller
or larger than first, and distinctly larger than first inner tooth; inner teeth arranged in three
series; mandibular serrations numbering two, first much larger than second; short basal
setae slender, glabrous (Fig. 25 K); hypostomium with anterior border slightly emarginat-
ed; teeth numbering 13, subequal in size, not arranged in conspicuous groups; 2+2 lateral
teeth inserted much below level of lateral teeth (Fig. 25 I); lateral sclerite of proleg well
developed, subtriangular, about as long as wide (Fig. 25 M); setae of lateral sclerite less
than half as long as sclerite, arranged in irregular row extending slightly beyond main
body of sclerite below; larval cuticle bare or conspicuously hairy (Fig. 25 N); perianal
scales well developed; ninth segment with ventral tubercles; anal sclerite normal, X-
shaped (Fig. 25 L).
Distribution: This genus has been found in Misiones (Argentina) and in the coastal
ranges of southern Brazil, from the State of Rio de Janeiro south to Paraná (Fig. 136).
Bionomics: The aquatic instars have been found in fast- and slow-flowing mountain
streams, generally at elevations of 1000 m or above, although a few sites of lower
elevation are known. LUTZ (1910) collected females of L. pernigrum on horses.
Discussion: In many respects, Lutzsimulium fits the features normally associated with the
Prosimuliini, as understood by CROSSKEY (1969): the short apical segment of the maxillary palp
104 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

of the adults, the absence of a pedisulcus, multiple apical spines on the gonostylus, the
structure of the median sclerite of the genitalia of the male not unlike that found in
Prosimulium or Stegopterna, the shapeless loosely woven cocoon, the comparatively heavily
sclerotized pupal abdomen, the membranous area dividing sterna VI and VII of the pupa
along their middle, the long terminal spines of the pupal abdomen and the modified setae on
the terminal abdominal segments, and the dark third segment of the antenna of the larva. On
the other hand, Lutzsimulium shows an array of apomorphic characters generally associated
with the tribe Simuliini sensu CROSSKEY: the rather deep mesepisternal sulcus, the absence of a
basal cell in the wing, the conspicuous apodemes of the arms of the furcasternum, the
Simulium-like hypostomium of the larva, the small number of mandibular serrations, and the
extremely well-developed postgenal cleft. It is not possible to place Lutzsimulium, using
conventional criteria, unequivocally in one or the other tribe accepted by CROSSKEY (loc. cit.).
The combination of the peculiar autapomorphic characters of Lutzsimulium, such as the mesh-
like surface structure of part of the spermatheca, the reduced endoparameral organ in the
genitalia of the male, the apically looped trichomes of the thorax of the pupa, and the size
ratios of the preapical teeth of the mandible of the larvae are sufficient to distinguish
Lutzsimulium from any described genus-group in the Simuliidae.

KEYS TO LUTZSIMULIUM SPECIES

Adults
Lutzsimulium flavopubescens adults are not included due to lack of sufficient information.
– Female and male scutum reddish brown with coppery pilosity; female sensory
organ of maxillary palp big (about 0.5 of palpomere length, Fig. 24 B); third
segment of palp relatively thick; male median sclerite without strong constriction
on median area (Fig. 24 J) ............................................................ Lutzsimulium hirticosta
– Female and male scutum dark velvety brown with brown to reddish pilosity;
female sensory organ of maxillary palp small (about 0.3 of palpomere length, Fig.
24 K); third segment of palp relatively thin; male median sclerite with strong
constriction on median area .......................................................Lutzsimulium pernigrum

Pupae
1 Gill branches 20-22; frontoclypeus and thorax with abundant tubercles (Fig. 25 D)
............................................................................................................ Lutzsimulium hirticosta
– Gill branches 12-14; frontoclypeus and thorax with or without tubercles ............ 2
2(1) Gill branches 14; frontoclypeus and thorax without tubercles ..................................
...........................................................................................................Lutzsimulium pernigrum
– Gill branches 12; frontoclypeus and thorax with tubercles ........................................
.................................................................................................... Lutzsimulium flavopubescens

Larvae
– Postgenal cleft deep, with postgenal bridge half length of hypostomium; cervical
sclerites isolated from postocciput border; anal ring with 72-74 rows of 12-13
hooks each ....................................................................................... Lutzsimulium hirticosta
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 105

– Postgenal cleft not deep, with postgenal bridge as long as or longer than
hypostomium; cervical sclerites wide at postocciput border; anal ring with 96-110
rows of 18-25 hooks each .......... Lutzsimulium pernigrum, Lutzsimulium flavopubescens

CLAVE DE ESPECIES DE LUTZSIMULIUM


Adultos
No se incluye Lutzsimulium flavopubescens por ser escasa la información.
– Escudo de hembra y macho castaño rojizo con pilosidad cobriza; vesícula sensorial
del palpo maxilar de hembra grande (aproximadamente 0,5 de la longitud del
palpómero, Fig. 24 B); tercer artejo del palpo relativamente grueso; esclerito
mediano del macho sin fuerte constricción en área mediana (Fig. 24 J) ....................
............................................................................................................ Lutzsimulium hirticosta
– Escudo de hembra y macho castaño oscuro aterciopelado, con pilosidad castaño
rojiza; vesícula sensorial del palpo maxilar pequeña (aproximadamente 0,3 del
palpómero, Fig. 24 K); tercer artejo del palpo relativamente fino; esclerito
mediano del macho con fuerte constricción sobre el área mediana .........................
...........................................................................................................Lutzsimulium pernigrum

Pupas
1 Ramas de las branquias en número de 20-22; frontoclípeo y tórax con abundantes
tubérculos (Fig. 25 D) ................................................................... Lutzsimulium hirticosta
– Ramas de las branquias en número de 12-14; frontoclípeo y tórax con o sin
tubérculos ............................................................................................................................ 2
2(2) Ramas de las branquias en número de 14; frontoclípeo y tórax sin tubérculos .....
...........................................................................................................Lutzsimulium pernigrum
– Ramas de las branquias en número de 12; frontoclípeo y tórax sin tubérculos .....
.................................................................................................... Lutzsimulium flavopubescens

Larvas
–- Hendidura postgenal profunda, con puente postgenal la mitad del largo del
hipostomio; escleritos cervicales aislados del borde del postocciput; anillo anal
con 72-74 hileras de 12-13 ganchos cada una ......................... Lutzsimulium hirticosta
– Hendidura postgenal no profunda, con puente postgenal tan largo o más que el
hipostomio; escleritos cervicales anchos, llegando al borde del postocciput; anillo
anal con 96-110 hileras de 18-25 ganchos cada una .....................................................
.......................................................... Lutzsimulium pernigrum, Lutzsimulium flavopubescens

Lutzsimulium hirticosta (LUTZ) (Figs. 24 A-J; 25 A-N.)


Simulium hirticosta LUTZ, 1909: 135; 1910: 226; PINTO, 1931: 756.
Lutzsimulium hirticosta VULCANO, 1967: 4; WYGODZINSKY & COSCARÓN, 1973a: 197; PY-DANIEL, 1982:
305; COSCARÓN, 1991: 80-81.
Lutzsimulium cruzi D’ANDRETTA & D’ANDRETTA, 1947: 401; 1948: 639.
106 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

K
A

D
E

B C

L N

Figure 24. A-J: Lutzsimulium hirticosta. A-G: Female; A: frons; B: third palpomere and sensory
vesicle; C: calcipala and palpomere I; D: claw; E: portion of eighth sternite and gonapophyses; F:
genital fork; G: portion of spermatheca with insertion base of spermathecal duct. H-J: Male; H:
chaetotaxy of wing anterior veins; I: genitalia, ventral view showing gonocoxite and gonostylus; J:
ventral plate and portion of median sclerite. K-N: L. pernigrum. K-L: Female; K: third palpomere
with sensory vesicle; L: furcasternum. M-N: Male; M: gonocoxite and gonostylus; N: Pupa, portion
of thorax with gill and trichomes.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 107

G
F

K
B

C E
D

L
N
Figure 25. Lutzsimulium hirticosta. A-E: Pupa; A: general view; B: gill; C: basal portion of gill,
magnified; D: frontoclypeus; E: abdomen distal portion, ventral view, showing curled trichomes and
terminal spur. F-N: Larva; F: general aspect, lateral view; G: head, dorsal view; H: postgenal cleft and
hypostomium; I: hypostomium, anterior border; J: antenna; K: mandible apex showing teeth; L: anal
sclerite, with scales and hairs; M: lateral sclerite of thoracic proleg; N: body integument hair.
108 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Female: Wing length 2.4-2.7 mm; head blackish; antenna, palp and proboscis dark brown
to blackish; frons light grayish brown, scutum reddish brown with coppery pilosity and
abdomen dark grayish brown; legs brownish with darker areas blackish, hair and spines dark
brown to black; frons with faint median sulcus and abundant pilosity (Fig. 24 A); sensory organ
of maxillary palp about 2.5 times length of third palpomere (Fig. 24 B); Sc with about 50 hairs;
calcipala as figure 24 C; eighth sternite with about 27 hairs; gonapophyses with truncate border
(Fig. 24 E); genital fork with median branch well sclerotized (Fig. 24 F); spermatheca with
insertion base of spermathecal duct not totally sclerotized, mesh like(Fig. 24 G).
Male: Wing length 2.4-2.6 mm; coloration as in female but more darkened and with
more abundant blackish pilosity; distribution of wing vein hair and setae as in figure 24 H;
length/width ratio of hind basitarsus = 6; gonocoxite, gonostylus, endoparameres and
ventral plate as shown in figure 24 I; median sclerite expanded distally and without strong
constriction medially (Fig. 24 J).
Pupa: Cocoon leaving thorax and gills free (Fig. 25 A); length at base 3.0-3.5 mm,
dorsally 2.5-3.1 mm; gill length 2.0-2.5 mm; frontoclypeus and thorax dorsally with
abundant tubercles (Fig. 25 D); frontal trichomes only present in male; gill arborescent
with 20-22 branches arriving from short basal trunk with primary and secondary branches
relatively short (Figs. 25 B-C); abdomen with tubercles; distal portion of abdomen with
trichomes apically curved and terminal spines strong (Fig. 25 E).
Larva: Maximum length 5.0-5.5 mm; general aspect as figure 25 F; color grayish
yellow; cephalic apotome with abundant hair and positive spots well developed; cervical
sclerites small and isolated in postocciput (Fig. 25 G); postgenal bridge nearly half
hypostomium height and 1/4 of postgenal cleft (Fig. 25 H); ratio of length of proximal,
medial and distal antennal segments = 1:0.8:2.0; hypostomium with median tooth at same
height as lateral teeth (Fig. 25 I) with intermediate teeth well developed; antenna as figure
25 J; mandibular dentition as in figure 25 K; thoracic proleg sclerite with about 30 teeth
(Fig. 25 M); anal sclerite area with abundant hair and minute single scales (Fig. 25 L); anal
ring with 72-74 rows with 12-13 hooks each; larval cuticle with hair more abundant on
distal portion of body (Fig. 25 N).
Distribution: Argentina: Misiones; Brazil: Rio de Janeiro, Rio Grande do Sul, Santa
Catarina, São Paulo.
Bionomics: Lutzsimulium hirticosta breeds on vegetation in creeks with a small
volume of crystalline water.
Discussion: The closest species is L .pernigrum; differences are given in the keys.

Lutzsimulium pernigrum (LUTZ) (Figs. 24 K-N.)


Simulium pernigrum LUTZ, 1910: 263; PINTO, 1931: 708-756.
Simulium (Eusimulium) montanum LUTZ, 1909: 128 (nec PHILIPPI).
Simulium (Friesia) pernigrum RUBTZOV, 1940: 124.
Lutzsimulium pernigrum D’ANDRETTA & D’ANDRETTA, 1952: 312-324; WYGODZINSKY & COSCARÓN,
1973a: 197; PY-DANIEL, 1982: 304-305; COSCARÓN, 1991: 81-82.
Female: Wing length 2.5-3.1 mm; head blackish with dark brown appendages;
scutum dark brown with brown to reddish hair; legs dark brown; sensory organ of
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 109

maxillary palp small, about 1/5 of third palpomere length (Fig. 24 K); furcasternum as in
figure 24 L; genitalia similar to those of Lutzsimulium hirticosta.
Male: Wing length 2.8-3.0 mm; coloration velvety dark brown, homogeneous,
without special ornamentation and abundant coppery pilosity; length/width ratio of hind
basitarsus about 6.3; gonocoxite and gonostylus as in figure 24 M; median area of median
sclerite with strong constriction.
Pupa: Cocoon with irregular weave, leaving thorax free dorsally; body length 3.8-4.2 mm;
gill length 2.3-2.8 mm; frontoclypeus and thorax dorsally without tubercles; gill arborescent
with about 14 branches, with primary branches shorter than in Lutzsimulium hirticosta (Fig. 24
N); frontal trichomes in 2 pairs on male; abdominal trichomes abundant, curved distally.
Larva: Body length 7.5-8.0 mm; general color grayish brown; head yellowish brown;
cephalic apotome with positive spots not well pigmented; postgenal bridge as long as
hypostomium height and postgenal cleft smaller than either one; cervical sclerites
elongated and reaching occiput; cephalic fan with 35-41 rays; Ratio of proximal, medial
and distal antennal segments = 1:1.1-1.6:2.1-2.6; mandible with third preapical tooth
larger than first; thoracic proleg sclerite with about 45 teeth; anal ring with 96-102 rows
with 20-25 hooks each; larval cuticle without trichomes.
Distribution: Argentina: Corrientes; Brazil: Rio de Janeiro, Rio Grande do Sul, São Paulo.
Bionomics: Lutzsimulium pernigrum is found on vegetation in small creeks with clear water.

Lutzsimulium flavopubescens (LUTZ)


Simulim flavopubescens LUTZ, 1910: 248-249, 263, 265; PINTO, 1931: 702.
Eusimulium flavopubescens LANE & PORTO, 1939: 169; LANE & VULCANO, 1943: 431, 434-436.
Lutzsimulim flavopubescens PY-DANIEL, 1982: 308; COSCARÓN 1991: 82.
Female (based on lectotype): Frontal sulcus very evident and frons wider than in
Lutzsimulium hirticosta; fronto-ocular triangle about as long as wide; claws, calcipala and genitalia
similar to those of Lutzsimulium hirticosta; sensory organ of palp about half of basal segment length.
Male: Unknown.
Pupa: Cocoon with soft weave; length dorsally 2.9 mm, ventrally 3.6 mm; gill length
1.8-2.6 mm; gill arborescent with 12 branches arising from short primary branches and these
from very short trunk; head, thorax and abdomen with tubercles; head with 2+2 frontal and
1+1 facial single trichomes; thorax dorsally with 5+5 trichomes, 3 of them curved apically;
trichomes on distal portion of abdomen from gradually curved to strongly curved at apex.
Larva: Ratio of proximal, medial and distal antennal segments = 1:1.5-1.6:2.3-2.4;
postgenal cleft not deep, postgenal bridge two times longer than hypostomium length; cervical
sclerites wide, reaching postocciput; anal ring with 108-110 rows with 18-22 hooks each.
Distribution: Brazil: Rio de Janeiro.
Bionomics: Unknown.
Discussion: There are few specimens of Lutzsimulium flavopubescens; most of them
are mounted on slides, and it is difficult to see some characters of taxonomic value. The
bionomic information is dubious and very poor. The best information comes from pupae.
Available information, based especially on pupal and larval morphology, suggests that
Lutzsimulium flavopubescens is very similar to L. pernigrum.
110 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Kempfsimulium PY-DANIEL & NUNES DE MELLO (Figs. 26, 27, 136.)


Kempfsimulium PY-DANIEL, 1982: 294.
Type-species: Simulium simplicicolor LUTZ, 1910: 251.
Adults: Antennae each with 11 segments; frons of female as long as wide with
strong median sulcus bifurcated basally (Fig. 26 A); mandible with only internal teeth;
apical segment of maxillary palp about as long as subapical segment; mesepisternal sulcus
well developed; katepisternum wider than high; furcasternum with small lateral projec-
tions (Fig. 26 L); wings (Fig. 26 F) with Sc without hair-like setae; R1 with hair and setae,
Cu2 well curved distally; calcipala well developed and rounded apically; pedisulcus from
absent to faint in some males (Fig. 26 J); female claw with small subtriangular subbasal
tooth (Fig. 26 K); female eighth sternite with sclerotized area extending transversely from
one side to other; gonapophyses subtriangular with rounded apical border and internal
margin more sclerotized (Fig. 26 M); paraproct relatively short with small distal curvature
(Fig. 26 N); genital fork with median branch longer than lateral branches and well
sclerotized (Fig. 26 O); spermatheca oval, with sclerotized area reduced to neck of
spermathecal duct and without spiculae internally (Fig. 26 P-Q).
Male: Gonostylus about 1/3 length of gonocoxite, subconical and acuminate with two
apical teeth distally (Figs. 27 A-C); ventral plate subtrapezoidal, elongated and narrowed basally
(Figs. 27 A, D); endoparameres with basal plates and denticles inserted on membranous area
that cover also aedeagus and median sclerite (Fig. 27 A); median sclerite subovoidal.
Pupa: Cocoon with loose weave and evident threads; leaving gill and portion of
thorax free; gill with filamentous branches emerging close to base (Fig. 27 F); body well
sclerotized; frontoclypeus and thorax dorsally without tubercles; cephalothorax trichomes
single with 1+1 frontal and 2+2 facials small (Fig. 27 E); and 5-6 trichomes mostly curved
distally on each side of thorax disc (Fig. 27 F); abdominal chaetotaxy very similar to that
of Lutzsimulium, but showing two transverse carinae on tergites.
Larva: Cephalic apotome wider before posterior border, with median and lateral
punctiform spots scarcely evident (Fig. 27 H); cervical sclerites small and free from
postocciput; postgenal cleft deep with postgenal bridge shorter than hypostomium height;
antenna shorter than stem of cephalic fan length, with wide basal segments (Fig. 27 I);
combs of cephalic fan with similar setae of similar length; mandible with 3 preapical
teeth, median tooth reduced, 11 or 12 internal, and 2 marginal teeth (Fig. 27 K);
hypostomium teeth not arranged in groups (Fig. 27 J); lateral sclerite of thoracic proleg
subtriangular; larval cuticle without setae but with short cylindrical projections on cuticle
(Fig. 27 L); anal sclerite X-shaped, with abundant perianal scales; anal gill with 3 single
lobes; ninth abdominal segment without ventral tubercles.

CROSSKEY & HOWARD 1997, 2004 maintain Kempfsimulium in Lutzsimulium as a subgenus.


Differences between Lutzsimulium and Kempfsimulium females:
– General coloration brown-blackish, with coppery brown to blackish pilosity; Sc
with about 50 trichomes (Fig. 24 H); furcasternum branches with large projections
(Fig. 24 L); frons longer than wide with median sulcus relatively short and without
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 111

basal bifurcation (Fig. 24 A); spermatheca internally with spiculae; mesh-like


sclerotized area of spermathecal ducts also covering spermatheca neck and part
of spermathecal duct (Fig. 24 G); sclerotization of eighth sternite reduced to
medial area (Fig. 24 E). ................................................................................... Lutzsimulim
– General coloration brown-yellow greenish to reddish brown with light yellowish
brown pilosity; Sc without trichomes (Fig. 26 F); furcasternum branches with
small projections (Fig. 26 L); frons about as long as wide, with median sulcus long
with elongated basal bifurcate branches (Fig. 26 A); spermatheca internally
without spiculae; sclerotized area of spermathecal duct reduced to neck, without
mesh-like sclerotization at spermatheca union (Fig. 26 P); sclerotized area of
eighth sternite covering area from one side to other (Fig. 26 M). .... Kempfsimulium

Differences in the male, pupa and larva are given in the generic keys

Diferencias entre las hembras de Lutzsimulium y Kempfsimulium


– Coloración castaño negruzco, con pilosidad cobre-castaño; Sc con aproximada-
mente 50 tubérculos (Fig. 24 H); ramas del furcaesterno con grandes proyecciones
(Fig. 24 L); frente mas largo que ancho, con sulco mediano relativamente corto y
sin bifurcación basal (Fig. 24 A); espermateca con tubérculos internos; área
esclerotizada de los ductos de la spermatheca también cubriendo el cuello de la
espermateca y parte de los ductos, en aspecto de malla (Fig. 24 G); área
esclerotizada del octavo esternito reducido a las áreas medianas. (Fig. 24 E). .......
.............................................................................................................................. Lutzsimulim
– Coloración amarillo castaño verdoso a castaño rojizo con pilosidad castaño amarillen-
to; Sc sin tubérculos (Fig. 26 F); ramas del furcasterno con pequeñas proyecciones
(Fig. 26 L); frente tan largo como ancho, sulco mediano largo con alongadas ramas
bifurcadas basales (Fig. 26 A); espermateca sin tubérculos internos; área esclerotizada
de la espermateca reducido al cuello, sin aspecto de malla en la unión de la
espermateca (Fig. 26 P); área esclerotizada del octavo esternito cubriendo toda el área
transversalmente de un lado al otro (Fig. 26 M). ....................................... Kempfsimulium

Las diferencias en el macho, larva y pupa están el las claves a género.

Kempfsimulium simplicicolor (LUTZ) (Figs. 26 A-Q; 27 A-L.)


Simulium simplicicolor LUTZ, 1910: 251; 1922: 40; KNAB, 1911: 179; NEIVA & PENNA, 1916: 93; LUTZ,
1922: 90; PINTO, 1931: 712; FAIRCHILD, 1940: 706; VARGAS, 1945a: 197; VULCANO, 1967: 20;
LACEY & CHARLWOOD, 1980: 495, 497, 500-506; LACEY, 1981: 330.
Simulium hebeticolor LUTZ, 1910: 267; KNAB, 1915b: 179; SMART, 1945: 513; VULCANO, 1967: 20.
Eusimulium simplicicolor SURCOUF & GONZÁLEZ RINCONES, 1911: 286.
Lutzsimulium simplicicolor PY-DANIEL, 1981: 173; CROSSKEY & HOWARD, 1997: 16; 2007: 44.
Kempfsimulium simplicicolor PY-DANIEL, 1982: 300-308.
Lutzsimulium (Kempfsimulium) simplicicolor CROSSKEY & HOWARD, 1997: 16; 2004: 7.
112 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

F
B

C E A
R1

Rs

G
H
D

I
C
K
J

L
O
N
Q

P M
Figure 26. Kempfsimulium simplicicolor. A-Q: Female; A: frons; B: fronto-ocular triangle; C: third
palpomere with sensory vesicle; D: apex of mandible; E: basal portion of cibarium; F: wing, general
view; G: anterior portion of wings showing setae, hair and spiculae (C = costa, R1= radius 1, Rs=
secondary radius (radius 2) ; H-J: fore, mid and hind leg, respectively; K: claw; L: furcasternum; M:
eighth sternite and gonapophyses; N: cercus and paraproct; O: genital fork; P: spermatheca; Q:
portion of spermatheca with insertion base of spermathecal duct.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 113

A D

end. v.pl.
m.s.
ae.

G
E

I
K

J
L

Figure 27. Kempfsimulium simplicicolor. A-D: Male; A: genitalia, ventral view showing gonocoxite,
gonostylus, ventral plate, endoparameres base (end.) and aedeagal membrane (ae.); B: gonocoxite
and gonostylus, dorsal view; C: gonostylus, dorsal view; D: ventral plate (v.pl.) and median sclerite
(m.s.). E-G: Pupa; E: frontoclypeus; F: portion of thorax with gill, and trichomes; G: thoracic
trichomes magnified. H-L: Larva; H: head, dorsal view; I: antenna; J: hypostomium, anterior border;
K: apex of mandible with teeth; L: short cylindrical projections of body integument.
114 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Female: Wing length 2.3-2.7 mm; general color brownish; head brown, frons and
clypeus brown with grayish pollinosity; scutum from brown-yellow greenish to reddish
brown with decumbent hair orange to yellowish brown; scutellum and metanotum brown
reddish; pleurae grayish brown; wing veins light brown; legs light grayish brown, darkened
on tarsi; abdomen grayish brown with light brown hair; frons with deep frontal suture
(Fig. 26 A); fronto-ocular triangle about as high as wide (Fig. 26 B); sensory organ of
maxillary palp about 1/3 of basal segment (Fig. 26 C); cibarium smooth (Fig. 26 E); wing
chaetotaxy with thin hair brown and robust spine-like setae dark brown on C and R1 (Fig.
26 G); leg shape and coloration as in figure 26 H-J; claw subbasal tooth well developed
(Fig. 26 K); gonapophysis with abundant long hair and eighth sternite with about 21 setae
on each side (Fig. 26 M); genital fork with lateral branches relatively wide (Fig. 26 O);
discontinuous sclerotization of spermathecal neck reduced only to spermathecal duct
(Fig. 26 P-Q).
Male: Wing length 1.9-2.0 mm. general coloration as in female; gonocoxite and
gonostylus laterally flattened; gonostylus acuminate with 2 strong terminal teeth (Figs. 27
A-C); ventral plate basally with strong concavity and well-sclerotized margin (Figs. 27 A,
D); median sclerite and membrane of aedeagus with abundant microtrichia (Fig. 27 A).
Pupa: Cocoon with threads evident; gill length 0.9-1.3 mm; frontoclypeus as in
figure 27 E; gill light brown with 12 branches, arborescent, becoming thinner from base to
apex (Fig. 27 F); thoracic trichomes variable from straight to well curved (Fig. 27 G);
abdomen without tubercles but with robust spines and hooks.
Larva: Maximum length 5.5-7.0 mm; color grayish brown-green; cephalic apotome
yellowish, scarcely ornamented (Fig. 27 H), without hair; ratio of proximal, medial and
distal antennal segments = 1:2.0-2.5:3.0-3.6; distal segment more darkened (Fig. 27 I);
hypostomium with median teeth similar in height to intermediate and lateral teeth (Fig. 27
J); cephalic fan with 35-43 rays; anal ring with 110-130 rows with about 22 hooks in each.
Distribution: Brazil: Mato Grosso, Pará, Rondônia (Fig. 136).
Bionomics: The species breeds on Podostemaceae in rivers with strong current.
Females are haematophagous and were collected on humans, cows, horses and monkeys.

Simulium LATREILLE
Simulium LATREILLE, 1802: 428.
Type-species: Oestrus columbacensis SCOPOLI, 1780: 133 (monotype).
Adults: Color in general yellowish to black, scutum ornamentation homogeneous to
variable with filiform to flattened hair uniformly distributed or grouped and simulating
scales; wing without basal cell; basal sector of R with or without hair, shorter than
distance between Rs base and wing apex; radial sector unbranched; C and distal 2/3 of R1
with spiniform setae; hind basitarsus generally with calcipala; tarsomere I with deep
pedisulcus; legs with hair filiform or with petaloid shapes intermixed; female claws with or
without subbasal tooth; cibarium basally from light and smooth to well sclerotized and
with teeth of variable disposition; gonapophysis, paraproct, gonocoxite, gonostylus and
ventral plate of variable size and shape; spermatheca without sclerotized union with
spermathecal duct; endoparameres generally with abundant teeth.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 115

Pupa: Cocoon well developed, generally compact, with slipper to shoe shape;
frontoclypeus and thorax with trichomes single to multibranched, sometimes in elevated
number (nearly one hundred); abdominal segments with weak sclerotization; distal tergites
without special setae; terminal spines scarcely developed.
Larva: Cephalic apotome with variable ornamentation; cervical sclerites isolated;
mandible generally with 1 or 2 marginal teeth; distal antennal segment no longer than two
basal segments together; postgenal cleft varied from very deep to not evident; hypostomi-
al teeth very similar and median tooth about same level as lateral teeth; rectal papillae
single to having more than 10 diverticula on each lobe.
Distribution: This genus is distributed all over the world, and has the greatest
number of species, with great morphological variation (COSCARÓN, 1987; CROSSKEY &
HOWARD, 1997, 2004). For this reason, it is subdivided into a large number of subgenera;
some authors elevate these subgenera to the generic level (PY-DANIEL & MOREIRA
SAMPAIO, 1994, 1995).

KEYS TO SIMULIUM SUBGENERA

Females
1 General coloration black or blackish brown ................................................................ 2
– General coloration brown (reddish brown to yellowish) ......................................... 22
2(1) Body without special ornamentation; scutum hairs homogeneously distributed;
claws with basal tooth large, subovoid, elongated; basal portion of cibarium
smooth, without reinforcement at sides ............................... Eusimulium, Nevermannia
– Body ornamented with light vittae or spots; scutal hairs distributed homogeneously
or in groups; claws with or without subbasal tooth, but if present, it is
proportionally smaller and subtriangular (except Byssodon with basal tooth) ............ 3
3(2) With hair on basal sector of R ........................................................................................ 4
– Without hair on basal sector of R .................................................................................. 9
4(3) Grayish pollinose coloration; scutum with light area limited to 1 median and 1+1
sublateral stripes (Fig. 29 A); abdomen blackish or grayish with 1+1 silver
transverse bands (Fig. 28 A), or with bands on posterior border of tergites III-IV
(Fig. 51 A) ......................................................................................... Pternaspatha (in part)
– Blackish coloration; scutum with or without light submedian stripes or anterior
spots; abdomen blackish without silvery transverse bands on tergites III-IV ......... 5
5(4) Paraproct subtriangular, about twice as long as wide at base (Figs. 57 D, N); basal
portion of cibarium smooth ........................................................................... Chirostilbia
– Paraproct not subtriangular, curved distally, slightly longer than wide at base (Figs.
64 G, L, P); cibarium smooth or with teeth ................................................................. 6
6(5) Basal portion of cibarium with teeth arranged in elevated median subconical
group (Fig. 64 M) ..............................................................................................Inaequalium
– Cibarium generally smooth or with small teeth: if teeth present, these arranged in
median elevated group, scutum with 1+1 light anterior spots or 3 black
longitudinal stripes separated by nacreous stripes that change tone with different
light positions ...................................................................................................................... 7
116 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

7(6) Cibarium with very small teeth or without teeth (Figs. 102 C, F, G, L); paraprocts
relatively short not lobulate, scarcely acuminate apically and directed downwards
(Figs. 103 B, D, G, J) ............................................................................. Aspathia (in part)
– Cibarium with 1 row of teeth (Figs. 128 D, H) or without teeth but with
reinforced border (Figs. 132 B, L); paraproct relatively long, lobulate and scarcely
sclerotized, not acuminate (Figs. 128 C, E, J; 132 D, G, N) ..................................... 8
8(7) Scutum with hair grouped, simulating scales (Fig. 132 I), without anterior grayish
subquadrate spots; cibarium smooth (Figs. 132 B, L) .............. Thyrsopelma (in part)
– Scutum with hair homogeneously distributed, with anterior grayish subquadrate
light spots (Fig. 128 A); cibarium with teeth arranged in 1 row (Figs. 128 D, H)
............................................................................................................. Trichodagmia (in part)
9(3) Cibarium without teeth ................................................................................................... 10
– Cibarium with teeth ......................................................................................................... 16
10(9) Scutum with hair grouped like scales (Fig. 5 A); claw without subbasal tooth or
much reduced (Figs. 54 D, E); fronto-ocular triangle absent; paraprocts very thin
distally (Figs. 54 I, Q, X) ................................................................................... Notolepria
– Scutum with hair homogeneously distributed; claw with or without subbasal tooth,
fronto-ocular triangle well developed; paraprocts not thinner distally .................... 11
11(10) Species grayish pollinose; abdomen grayish with tergites III-V blackish or velvety
brown, generally with anteromedian or 1+1 silvery lateral spots; calcipala
frequently reduced or absent ......................................................................................... 12
– Species blackish; abdomen blackish without silver spots on tergites III-V; calcipala
well developed .................................................................................................................. 13
12(11) Claws with subbasal tooth (Fig. 51 F); scutum with blackish areas delimited by
grayish median and sublateral stripes; abdominal tergites III-V blackish with 1+1
sublateral posterior whitish spots (Fig. 51 A); paraprocts subrectangular, shorter
than wide (Fig. 51 H) ..................................................................................... Pternaspatha
– Claws without subbasal tooth (Fig. 99 C); scutum grayish uniform except 1+1
light median anterior spots; tergites III-V blackish or velvety brown, with or
without spots anteromedially (Figs. 98 A, P; 99 A); paraprocts subtriangular (Fig.
99 G), as long as wide .......................................... Psilopelmia (BLANCASI sp. group)
13(11) Paraproct subtriangular, with distal border arched (Figs. 118 A, B); scutum with
1+1 anterior light submedian spots continued posteriorly by thin stripe joining
with grayish posterior area ..................................... Hemicnetha (OVIEDOI sp. group)
– Paraproct not subtriangular, with distal border truncate or acuminate posteriorly;
scutum ornamentation variable ..................................................................................... 14
14(13) Paraprocts with distal edge truncate and slightly acute posteriorly (Figs. 103 B, K);
scutum with median and 1+1 submedian black stripes delimited by nacreous
vittae that invert coloration with change of illumination position (Figs. 101 A, B,
F, G) ......................................................................................................................... Aspathia
– Paraprocts with distal edge truncate or rounded but not acuminate posteriorly;
scutum without contrasting silvery vittae (Figs. 113 A, D, G, H; 120 A, B, K, L)
.............................................................................................................................................. 15
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 117

15(14) Paraproct as long as wide at base, with external surface mostly nude and shiny,
with depressions and distal border truncated (Fig. 120 I); gonapophysis as long as
wide .............................................................................................................................Hearlea
– Paraproct longer than wide at base, external surface mostly covered by
microtrichia, without depressions and distal border curved (Figs. 113 C, F, L, M,
R, U), gonapophysis longer than wide (Figs. 113 C, K, Q, T, V) ..............................
..................................................... Hemicnetha (in part BRACHYCLADUM sp. group)
16(9) Scutum with abundant gray pollinosity, showing, with some illumination, 3-5
longitudinal light stripes; cibarium with weak concavity covered by small similar
acute teeth; paraprocts subtriangular, about as long as wide; claws without teeth
...................................................................................................................................... Psilozia
– Scutum with sparse pollinosity, with or without 1+1 silvery longitudinal stripes;
cibarium with or without teeth, generally with median depression; paraprocts not
subtriangular, with curved distal edge, generally longer than wide and sometimes
thinner distally; claws with or without teeth .............................................................. 17
17(16) Claws with large subovoidal basal tooth; posterior edge of tergites III-V
frequently with 1+1 transverse silvery bands .................................................. Byssodon
– Claws with small subtriangular subbasal tooth or without tooth; tergites III-V
without silvery transverse bands ................................................................................... 18
18(17) Cibarium generally without teeth, but if teeth present, not in median depression
and at sides (Figs. 102 F, G, K, O); scutum with 1+1 submedian longitudinal
silvery stripes generally reaching posterior grayish area (Figs. 101 A, F) or darker
with change of light direction (Figs. 101 B, G); fronto-ocular triangle well
developed ................................................................................................................ Aspathia
– Cibarium with median depression sclerotized or not and with small teeth at sides
(Figs. 71 P; 76 N; 80 D; 84 M; 91 Q); scutum with or without silvery longitudinal
stripes that can reach to posterior silvery area; fronto-ocular triangle generally
absent or reduced ............................................................................................................. 19
19(18) Cibarium with central area well sclerotized and with small rounded teeth (Figs. 80
D, P, T); infra-frontal sutures well developed (Figs. 80 A, V) ....................................
....................................................................Coscaroniellum (QUADRIFIDUM sp. group)
– Cibarium with central area not sclerotized, teeth acute (Figs. 71 P; 76 N, X); infra-
frontal sutures not developed ........................................................................................ 20
20(19) Scutum black, with or without silvery submedian stripe mostly frequent on
anterior area and wedge shaped (Figs. 71 L, O, Q, U); when reaching posterior
silvery area, without gray shadow spots on silvery stripe ................... Psaroniocompsa
– Scutum grayish to black with silvery stripe reaching posterior grayish area,
generally with gray shadow spots on silvery stripes (Figs. 76 A-F, H, I, L, M, Q, R)
.............................................................................................................................................. 21
21(20) Cibarium with small teeth about same size and absent on median depression (Figs.
76 N, X); fronto-ocular triangle very short (Figs. 76 G, W); abdomen without
silvery posterior transverse spots on tergites III-V or very short (Fig. 76 K) ........
............................................................................................................................ Cerqueirellum
118 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

– Cibarium with submedian teeth larger than other teeth and with small teeth on
median area (Fig. 80 Y); fronto-ocular triangle relatively deep; abdomen with
silvery posterior transverse spots on tergites III-V about 1/3 of tergite height
(Fig. 80 X); scutum with submedian silvery vittae narrow (Fig. 80 W) ....................
.......................................................... Coscaroniellum (QUADRIVITTATUM sp. group)
22(1) Species yellowish-reddish to greenish gray; if scutum black, margins are yellowish
(Figs. 84 B-D); paraprocts subtriangular (Figs. 85 D-P) longer than wide at base
and distally relatively thin (Figs. 92 A-N) ................................................................... 23
– Species reddish brown to dark grayish; scutum generally with lyre-shaped pattern
(Fig. 128 A); paraprocts subrectangular and distally not acuminate (Figs. 128 C, E, J)
.............................................................................................................................................. 28
23(22) Scutum gray-greenish to brown-yellowish; cibarium smooth not sclerotized (Fig.
60 D) .............................................................. Chirostilbia (SUBPALLIDUM sp. group)
– Scutum orange to reddish or black bordered by yellow and frequently with
submedian silvery stripes; cibarium with sclerotized margin or with small teeth ..
.............................................................................................................................................. 24
24(23) Scutum with 1+1 or more nacreous or silvery longitudinal stripes (Figs. 91 B-E);
abdomen generally with 3 longitudinal rows of subquadrate blackish spots (Figs.
91 G-I); cibarium with median depression well sclerotized, bordered with weak
lobules and at sides with teeth on weakly elevated areas (Figs. 91 J-Q); paraprocts
thinner distally (Figs. 92 A-N) ............................ Psilopelmia (ESCOMELI sp. group)
– Scutum generally without 1+1 or more nacreous or silvery longitudinal stripes
(Figs. 84 A-E); abdomen without subquadrate blackish spots; cibarium without
teeth or with teeth generally larger and with different disposition from above
(Figs. 84 I-W); paraprocts not thin distally (Figs. 85 D-K) ........... Ectemnaspis......25
25(24) Scutum yellow or black, in the latter case, totally with yellow margins (Figs. 84 A-
E); cibarium without or with teeth, in the latter case, with plates or small lobules at
median area (Figs. 84 I-P) .............................................................................................. 26
– Scutum yellow or reddish, cibarium with teeth but median area without plates
.............................................................................................................................................. 27
26(25) Cibarium without teeth but with well sclerotized edge (Figs. 84 I-J) .......................
............................................................................................... BICOLORATUM sp. group
– Cibarium with teeth and median area with plates or lobules (Figs. 84 K-P) ..........
.......................................................................................................... ROMANAI sp. group
27(25) Cibarium with well-developed teeth grouped on 1+1 weak elevations (Figs. 84 Q-S)
................................................................................................... PERFLAVUM sp. group
– Cibarium with very small teeth and with 1+1 submedian processes generally with
teeth on apex (Figs. 84 T-W) ...................................................... DINELLII sp. group
28(22) Gonapophysis subovoidal shorter than wide (Fig. 132 M); paraproct slightly
acuminate distally (Fig. 132 N); scutum with hairs grouped (Fig. 132 I) and
arranged in longitudinal rows ....................................................... Trichodagmia (in part)
– Gonapophysis subtriangular or subovoidal, longer than wide or about as long as
wide; paraprocts not acuminate distally; scutum with hairs homogenously
distributed or in groups, but not arranged in longitudinal rows ........................... 29
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 119

29(28) Cibarium scarcely sclerotized on median area and without teeth or spiculae (Fig.
113 B); paraprocts slightly longer than wide, subrectangular and with abundant
microtrichia (Fig. 113 C, F) ...................................... Hemicnetha (PAYNEI sp. group)
– Cibarium well sclerotized on median area, with thin teeth acuminate and scarcely
sclerotized (Figs. 113 J; 128 D, H); paraprocts about 2 times longer than wide, and
with sparse microtrichia (Figs. 113 L, M; 128 C, E, J) ............................................. 30
30(29) Gonapophysis more than 2 times longer than wide, acute apically (Fig. 113 K);
cibarium with small teeth disordered (Fig. 113 J) .........................................................
................................................................... Hemicnetha (BRACHYCLADUM sp. group)
– Gonapophysis less than 2 times longer than wide, and blunt apically (Figs. 128 B,
I); cibarium with conspicuous teeth arranged along border to base of cornuae
(Figs. 128 D, H) ............................................................................................... Trichodagmia

Males
1 General coloration black, sometimes abdomen with greenish tones; with or
without hair on basal sector of R ................................................................................... 2
– General coloration variable form dark brown to yellowish, generally with hair on
basal sector of R .............................................................................................................. 23
2(1) With hair on basal sector of R ........................................................................................ 3
– Without hair on basal sector of R ................................................................................ 11
3(2) Gonostylus longer than gonocoxite (Figs. 133 B, C, H, J); ventral plate relatively
short and with or without median process and carina (Figs. 133 D, I, K) ............ 4
– Gonostylus shorter than gonocoxite; ventral plate with variable shape ................ 5
4(3) Scutum with hairs homogeneously arranged and without silvery spots; ventral
plate distally with small, median but not lateral processes (Fig. 129 E) ...................
............................................................................................................................. Trichodagmia
– Scutum with hairs disposed in groups like scales and with 1+1 silvery anterolateral
spots sometimes continued submedially reaching to posterior grayish area (Figs.
133 A, G); ventral plate distally with large median and lateral processes (Figs. 133
D, I) ..................................................................................................................... Thyrsopelma
5(3) Gonostylus distal spur much reduced or absent ................................ Chirostilbia......6
– Gonostylus with distal spur ............................................................................................. 7
6(5) Gonostylus without lateral crest (Figs. 58 B, F, G, I, L, O) ........................................
......................................................................................................... PERTINAX sp. group
– Gonostylus with lateral crest (Figs. 61 D, F, H) ............................................................
................................................................................ SUBPALLIDUM sp. group (in part)
7(5) Scutum without ornamentation; gonostylus flattened and curved, with
subtriangular internal lobe ....................................................... Eusimulium, Nevermannia
– Scutum, with anterior lighting, generally ornamented with spots; gonostylus with
variable shape ...................................................................................................................... 8
8(7) Gonostylus (microscope slide mounted) subquadrate with distal concavity and
subapical spur (Figs. 28 N; 30 R; 41 I) ............................................... Pternaspahta......9
– Gonostylus (microscope slide mounted) subtriangular to subtrapezoidal or
subcylindrical and with apical spur .............................................................................. 10
120 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

9(8) Gonostylus with tubercles (Figs. 28 N; 30 R; 31 S) .....................................................


......................................................................................... NIGRISTRIGATUM sp. group
– Gonostylus without tubercles (Figs. 41 I; 47 K; 52 K) ... NEMORALE sp. group
10(8) Gonostylus subtriangular or subtrapezoidal, with 1 apical spur (Figs. 65 D, F-H, J-N,
P); ventral plate without lateral constriction (Figs. 65 C, E, I, O) ............. Inaequalium
– Gonostylus subcylindrical with weak distal curvature and 1 apical spur (Figs. 86
H-N) (except wolffhuegeli, in which it is relatively short and has several apical spurs);
ventral plate with weak lateral constriction (Figs. 87 A-H) .........................................
............................. Ectemnaspis (BICOLORATUM sp. group; ROMANAI sp. group)
11(2) Gonostylus much shorter than gonocoxite ................................................................ 12
– Gonostylus as long as or longer than gonocoxite ..................................................... 19
12(11) Gonostylus without apical spur .................................................................................... 13
– Gonostylus with apical spur .......................................................................................... 14
13(12) Scutum without ornamentation; gonostylus subovoidal with reinforced edge
(Figs. 54 N, S, T) ................................................................................................. Notolepria
– Scutum with 1+1 submedian anterior silvery spots (Figs. 61 A, B); gonostylus
subtrapezoidal with sublateral crest (Figs. 61 D, F, H, I) ............................................
......................................................................... Chirostilbia (SUBPALLIDUM sp. group)
14(12) Scutum with 1+1 submedian silvery vittae, reaching or not to grayish posterior
area; gonostylus subtriangular or subtrapezoidal ...................................................... 15
– Scutum without 1+1 submedian silvery vittae, but with anterior subquadrate or
subtriangular silvery spots (Figs. 86 A-D; 91 A-D); gonostylus subrectanglular or
subcylindrical ..................................................................................................................... 17
15(14) Gonostylus about 2 times longer than wide at base; and apical spur spatuliform
(Figs. 81 C, G, H) ......................................................................................... Coscaroniellum
– Gonostylus only slightly longer than wide at base; terminal spur subconical .... 16
16(15) Gonostylus subtriangular with apical or subapical spur (Figs. 77 J, O; 81 L) .........
................ Byssodon, Cerqueirellum, Coscaroniellum (QUADRIVITTATUM sp. group),
Psaraniocompsa (SIOLLII sp. group)
– Gonostylus subquadrate or subtrapezoidal with submedian spur (Figs. 72 F, H, I)
.................................................................. Psaraniocompsa (INCRUSTATUM sp. group)
17(14) Gonostylus subcylindrical, generally with weak dorsal curvature and 1 apical spur
(Figs. 86 H-N; 93 K; 118 F, H) ............... Ectemnaspis (BICOLORATUM sp. group;
ROMANAI sp. group); Psilopelmia and Hemicnetha (OVIEDOI sp. group)
– Gonostylus subtrapezoidal with 2-15 marginal spurs .............................................. 18
18(17) Ventral plate more than 2 times wider than long (Fig. 87 I); gonostylus with 7-15
marginal spurs (Fig. 86 O) ................................. Ectemnaspis ( ROMANAI sp. group)
– Ventral plate as wide as long; gonostylus with 2-4 subapical spurs .............. Psilozia
19(11) Gonostylus subcylindrical and thin basally, generally narrower than half of
gonocoxite, distal edge wide and generally with basal process .............................. 20
– Gonostylus basally wider than half of gonocoxite, distal edge wide and without
basal process ...................................................................................................................... 21
20(19) Ventral plate generally becoming narrower apicomedially (Figs. 106 M-P);
gonostylus with basal process present (Figs. 105 B, C, E-H, J-L) .............. Aspathia
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 121

– Ventral plate not becoming narrower apicomedially (Fig. 121 D, G ); gonostylus


without basal process but with small subbasal carina (Figs. 121 C, F) ......... Hearlea
21(19) Gonostylus flattened and sinuous, scarcely narrowed distally (Figs. 114 A, E, G, I,
M, N); endoparameres with large teeth and base well sclerotized (Fig. 114 K) .....
..................................................................................................................... Hemicnetha......22
– Gonostylus subtriangular, narrowed and well curved apically (Figs. 133 B, C, H, J);
endoparameres without teeth and base not well sclerotized (Fig. 133 F) ................
.............................................................................................................................. Thyrsopelma
22(21) Ventral plate wider than long, with large median process (Figs. 114 B, C, F, H, L)
.................................................... PAYNEI sp. group, BRACHYCLADUM sp. group
– Ventral plate about as long as wide, without median process but with median
carina (Figs. 114 J, O) ........................................................... MEXICANUM sp. group
23(1) Gonostylus almost 2 times longer than gonocoxite ................................................. 28
– Gonostylus shorter or slightly longer than gonocoxite ........................................... 24
24(23) Gonostylus with 3-6 spurs or more (Fig. 129 L); endoparameres without teeth
(Fig. 129 C) ....................................................................................... Trichodagmia (in part)
– Gonostylus with 1 or no spurs; endoparameres generally with large teeth (Figs. 61
I; 99 I) ................................................................................................................................. 25
25(24) Basal sector of R without hair; scutum dull grayish; gonostylus with internal
border straight (Figs. 98 V; 99 H) ...................... Psilopelmia (BLANCASI sp. group)
– Basal sector of R with hair (Fig. 60 C); scutum not grayish; gonostylus with
internal border curved .................................................................................................... 26
26(25) Gonostylus without apical spur, with sublateral crest and distally not curved (Figs.
61 D, F, H, I) ................................................ Chirostilbia (SUBPALLIDUM sp. group)
– Gonostylus with apical spur, without sublateral crest and distally slightly curved
.............................................................................................................................................. 27
27(26) Ventral plate generally longer than wide at base (Figs. 87 J-L); gonostylus without
distal curvature (Figs. 86 P, Q, R) ............... Ectemnaspis (PERFLAVUM sp. group)
– Ventral plate generally shorter than wide at base (Figs. 87 M-O; 94 A-L);
gonostylus generally with distal curvature (Figs. 86 S; 93 H-R) ................................
............................................................... Psilopelmia, Ectemnaspis (DINELLII sp. group)
28(23) Scutum dark brown with hairs grouped like scales; gonostylus distally relatively
thick (Fig. 133 J) ............................................................................................... Thyrsopelma
– Scutum brown-reddish with hairs not grouped like scales; gonostylus distally
relatively thin (Figs. 129 B, D) ...................................................................... Trichodagmia

Pupae
1 Gill arborescent with relatively thick branches, with acute apices well sclerotized
and spine shaped, with 12-50 branches (Figs. 134 B, E, G, K), cocoon compact
and with ventral side projected anteriorly .................................. Thyrsopelma (in part)
– Gills with variable shape but without acute, well-sclerotized, spine-shaped apices;
cocoon of variable shape ................................................................................................. 2
2(1) Gills thick, asymmetrical, generally with curvature .................................................... 3
– Gills generally filiform, symmetrical, without lateral curvature ............................... 6
122 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

3(2) Frontoclypeus and exposed portion of thorax with single trichomes (Fig. 121 H);
gills with striations (Figs. 122 A-K; 123 A-O; 124 A-K); cocoon without dorsal
projection ...................................................................................................................Hearlea
– Frontoclypeus and exposed portion of thorax with multibranched trichomes with
2-8 branches; gill without striations; cocoon generally with dorsal projection ..... 4
4(3) Cocoon generally without dorsal projection, anterior edge straight (Figs. 66 A, B,
D, L; 70 A, B, P, Q) ...........................................................................................Inaequalium
– Cocoon projected dorsally (Figs. 73 H-K; 88 C, D, F; 107 I) .................................. 5
5(4) Frontoclypeus and thorax with trichomes bifid (Figs. 73 O-Q) ....... Psaroniocompsa
– Frontoclypeus and thorax with trichomes of 3-8 branches .......................................
........................................................................................................... .Ectmemnaspis, Aspatha
6(2) Cocoon with compact weave and anterior portion elevated, generally protecting
gill base (Figs. 115 A-D; 59 L, O, S); gill filaments relatively thick ......................... 7
– Cocoon without compact weave and generally with anterior portion not elevated
or protecting gill; gill filaments relatively thin ........................................................... 13
7(6) Cocoon anterior aperture with festoons protecting gill base (Figs. 115 A-D) ..... 8
– Cocoon anterior aperture without festoons protecting gill base ............................. 9
8(7) Gill of 8 branches with apices blunt and resistant (Figs. 116 A, B); frontoclypeus
generally reinforced at base and enlarged at facial area, with female and male
having similar facial area (Figs. 115 O-T) ............. Hemicnetha (PAYNEI sp. group)
– Gill of10 branches frequently with membranous apices blunt, not resistant (Fig.
59 N); frontoclypeus not reinforced at base and not enlarged at facial area, with
male frons basally narrower than in female ................................ Chirostilbia (in part)
9(7) Gill with 6 filaments ........................................................................................................ 10
– Gill with more than 6 filaments .................................................................................... 11
10(9) Frontoclypeus and thorax with numerous tubercles, some of them acute like short
spines (Figs. 119 B, D, E); cocoon scarcely elevated anteriorly (Fig. 119 A) ..........
...................................................................................... Hemicnetha (OVIEDOI sp. group)
– Frontoclypeus and thorax without tubercles (Fig. 115 R); cocoon frequently very
elevated anteriorly (Fig. 115 E) .......... Hemicnetha (BRACHYCLADUM sp. group)
11(9) Gill with 18-20 branches arranged in bunch (Fig. 130 L) ...... Trichodagmia (in part)
– Gill with 8-16 branches ................................................................................................... 12
12(11) Gill with 12 branches ..........................................................................................................
.................................... Hemicnetha (MEXICANUM sp. group), Trichodagmia (in part)
– Gill with 10 branches ............. Hemicnetha (BRACHYCLADUM sp. group in part)
13(6) Gill with 4 filamentous branches .................................................................................. 14
– Gill with more than 4 filamentous branches .............................................................. 17
14(13) Frontoclypeus and thorax with tubercles acuminate (Fig. 82 D) ..............................
.......................................................................................................... Coscaroniellum (in part)
– Frontoclypeus and thorax with tubercles not acuminate ........................................ 15
15(14) Cocoon reduced ventrally (Figs. 98 X; 99 K); cephalic and thoracic trichomes
single (Fig. 98 a-b) ................................................. Psilopelmia (BLANCASI sp. group)
– Cocoon not reduced ventrally; cephalic and thoracic trichomes with 1-8 branches
.............................................................................................................................................. 16
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 123

16(15) Cephalic and thoracic trichomes generally bifid (Figs. 73 O, P, Q) ..........................


................................................................................................. Psaroniocompsa, Cerqueirellum
– Cephalic and thoracic trichomes with 1-8 branches .....................................................
.................................................................. Eusimulium, Byssodon, Nevermannia, Psilopelmia
17(13) Gill with 6 filamentous branches .................................................................................. 18
– Gill with more than 6 filamentous branches .............................................................. 20
18(17) Thoracic trichomes numbering more than 10 on each side and frequently very
modified (Figs. 40 U-V; 44 R; 53 S, T; 108 G, H; 109 A, G) .....................................
............................................................................................ Pternaspatha, Aspathia (in part)
– Thoracic trichomes numbering about 5 on each side .............................................. 19
19(18) Cephalic and thoracic trichomes bifid (rarely single or trifid) ...................................
................................................................ Psaroniocompsa, Cerqueirellum, Aspathia (in part)
– Cephalic and thoracic trichomes generally with more than 3 branches, not bifid
..................................................................... Byssodon, Inaequalium, Notolepria, Ectemnaspis
(BICOLORATUM and ROMANAI sp. group), Psilopelmia
20(17) Gill with 8-10 branches ................................................................................................... 21
– Gill with more than 10 branches ......................................................................................
...................... Psilozia, Chirostilbia, Aspathia, Ectemnaspis (PERFLAVUM sp. group)
21(20) Thoracic trichomes numbering more than 10 and frequently much modified (Figs.
29 N, P; 31 Y, Z); gill with 8 branches ....................................................... Pternaspatha
– Thoracic trichomes numbering 5 on each side; gill with 8-10 branches .............. 22
22(21) Cocoon weave spongy with elevated anterior margin and dorsal carina (Figs. 88 B,
H, I, J) ................. Ectemnaspis (BICOLORATUM and ROMANAI sp. group in part)
– Cocoon weave compact, without elevated anterior margin or dorsal carina (Figs.
62 C, D, G; 78 A, G, J) ............................. Chirostilbia (SUBPALLIDUM sp. group),
Notolepria, Cerqueirellum, Coscaroniellum, Psilozia, Byssodon, Aspathia, Ectemnaspis
(PERFLAVUM and DINELLII sp. groups), Psilopelmia

Larvae
1 Anal ring with more than 150 rows of hooks ............................................................. 2
– Anal ring with less than 120 rows of hooks ................................................................ 6
2(2) Body cuticle with lanceolate trichomes (Figs. 135 G, L, S); hypostomium with
median tooth and intermediate teeth not well differentiated (Figs. 135 D, I, N);
abdomen not dorsally flattened on distal portion (Fig. 135 A); cervical sclerites
enlarged, each more than 2 times wider than long (Fig. 135 Q) ........... Thyrsopelma
– Body cuticle without lanceolate trichomes; hypostomium with median tooth and
generally intermediate teeth well differentiated (Figs. 117 N, Q; 126 J; 131 G);
abdomen dorsally flattened distally (Figs. 131 A; 127 G); cervical sclerites not
enlarged, each less than 2 times wider than long (Fig. 131 I) .................................. 3
3(2) Anal sclerite frequently with sclerotized reinforcement, in some cases with ventral
branches enlarged and completing a ring (Figs. 127 B-J); anterior margin of
hypostomium generally arc shaped with lateral and intermediate teeth reduced
(Figs. 126 A, B, I, H, J) ...........................................................................................Hearlea
124 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

– Anal sclerite without sclerotized reinforcement nor with ventral branches


completing a ring; anterior margin of hypostomium with variable shape ............. 4
4(3) Cephalic apotome darkened on distal half in form of triangle (Fig. 63 A) ............
............................................................................................................................... Chirostilbia
– Cephalic apotome with different ornamentation ........................................................ 5
5(4) Hypostomium with lateral edges descending abruptly from lateral teeth, giving
plateau-like appearance (Figs. 117 P, Q; 131 G); mandible with preapical tooth
shorter than apical tooth (Figs. 117 L; 131 E, K) .........................................................
........................................... Hemicnetha (BRACHYCLADUM sp. group), Trichodagmia
– Hypostomium with lateral borders descending gradually from lateral teeth in
curved manner (Figs. 117 N, O, R); mandible with preapical tooth as long as or
longer than apical tooth (Figs. 117 K, M) ......................................................................
...................................... Hemicnetha (PAYNEI sp. group; MEXICANUM sp. group)
6(1) Anal ring with 100-120 rows of hooks; cephalic fan rays with setate of similar size
(Fig. 119 J); mandible with pair of large marginal teeth (Fig. 119 K) ......................
...................................................................................... Hemicnetha (OVIEDOI sp. group)
– Anal ring with less than 100 rows of hooks; cephalic fan rays alternating between
groups of small setae, and some larger ones; mandible with only 1 marginal tooth,
if there are 2 the second is very small (Figs. 29 S; 79 B; 83 C, O, R; 90 Q) ........ 7
7(6) Cephalic apotome without ornamentation or with small spot on median basal
area; with small numerous and short hairs on cephalic capsule (Fig. 79 C);
generally with dark band around first abdominal segment; antenna thin and
surpassing apex of cephalic fan stem (Fig. 79 H) ....................................................... 8
– Cephalic apotome with evident ornamentation; without or with very scarce hair
on cephalic capsule; antenna generally thick and surpassing, or not, length of
cephalic fan stem ................................................................................................................ 9
8(7) Mandibles with row of inferior supramarginal setae (Fig. 79 B); medial antennal
segment as long as or longer than proximal segment (Figs. 79 D, E, I) ..................
............................................................................................................................ Cerqueirellum
– Mandible without row of inferior supramarginal setae; medial antennal segment
about 1/2 to 4/5 of proximal segment (Figs. 83 B, N, Q) ................. Coscaroniellum
9(7) Postgenal cleft scarcely developed, with postgenal bridge longer than
hypostomium length (Figs. 112 A, B, D, G); rectal papillae without diverticula on
lobes .................................................................................................................................... 10
– Postgenal cleft well developed, with postgenal bridge as long as or shorter than
hypostomium length; rectal papillae generally with diverticula on lobes ............. 13
10(9) Antennae with thin transverse striations and surpassing apex of cephalic fan stem
(Figs. 111 A, B, H, L); medial antennal segment longer than proximal segment
(Figs. 111 B, H, L, M) ........................................................................................... Aspathia
– Antennae without striations and not surpassing apex of cephalic fan; medial
antennal segment shorter than proximal segment .................................................... 11
11(10) Ninth sternite without 1+1 tubercles ............... Psilopelmia (BLANCASI sp. group)
– Ninth sternite with 1+1 conspicuous tubercles ........................................................ 12
12(11) Antennae surpassing apex of cephalic fan stem ................ Eusimulium, Nevermannia
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 125

– Antennae about same length as apex of cephalic fan stem ............................Psilozia


13(9) Rectal papillae without diverticula on lobes ....... Psilopelmia (BLANCASI sp. group)
– Rectal papillae with diverticula on lobes ..................................................................... 14
14(13) Cephalic apotome with isolated spots: anteromedian, posteromedian, ante-
rolateral and posterolateral spots generally positive (Figs. 30 Y; 55 C; 56 K; 75 L)
.............................................................................................................................................. 15
– Cephalic apotome with variable ornamentation, different from above ............... 19
15(14) Anal sclerite without scales (Fig. 63 G) ....................................................................... 16
– Anal sclerite with scales (Fig. 63 P) .............................................................................. 17
16(15) Antenna surpassing apex of cephalic fan stem (Figs. 75 D, L); medial antennal segment
generally shorter than distal segment (Figs. 75 A, E, M) ........................... Psaroniocompsa
– Antenna not surpassing apex of cephalic fan stem; medial antennal segment
generally longer than distal segment (Figs. 90 M, N) ...................................................
...................... Ectemnaspis (BICOLORATUM sp. group; PERFLAVUM sp. group)
17(15) Body with maximum length less than 4.5 mm ............................................. Notolepria
– Body with maximum length over 5.0 mm .................................................................. 18
18(17) Medial antennal segment shorter than apical segment (Fig. 63 N) ...........................
......................................................................... Chirostilbia (SUBPALLIDUM sp. group)
– Medial antennal segment longer than apical segment (Fig. 29 S) ..............................
............................................................................................. Byssodon, Pternaspatha (in part)
19(14) Cephalic apotome darkened on basal half, leaving light window on median
subbasal portion (Figs. 90 E, F; 97 A-G); anal sclerite without scales .....................
............................. Ectemnaspis (BICOLORATUM sp. group; ROMANAI sp. group;
DINELLII sp. group), Psilopelmia
– Cephalic apotome with variable ornamentation, different from above; anal sclerite
with or without scales ..................................................................................................... 20
20(19) Anal sclerite with scales; medial antennal segment longer than distal segment
(Figs. 29 S; 31 b; 35 X) .................................................................................. Pternaspatha
– Anal sclerite without scales (Fig. 68 C); medial antennal segment shorter than
distal segment (Figs. 68 D, L) ........................................................................................ 21
21(20) Cephalic apotome with dark median basal diffuse spot that becomes thinner
anteriorly, delimiting 2 light elongated areas (Figs. 68 B, M; 70 M) .......Inaequalium
– Cephalic apotome with strong basal pigmentation generally shaped like triangle
(Fig. 63 A) .............................................................. Chirostilbia (PERTINAX sp. group)

CLAVES DE SUBGÉNEROS DE SIMULIUM


Hembras
1 Coloración general negro o castaño negruzco ............................................................ 2
– Coloración general castaño (castaño rojizo hasta amarillento) .............................. 22
2(1) Cuerpo sin especial ornamentación; pelos del escudo homogéneamente
distribuidos; uñas con diente basal grande de forma subovoidal alargada; porción
basal del cibario liso, sin refuerzos a los costados ............. Eusimulium, Nevermannia
126 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

– Cuerpo ornamentado con franjas o manchas claras; escudo con pelos distribuidos
homogéneamente o en grupos; uñas con o sin diente subbasal, si existe es
proporcionalmente más pequeño y de forma subtriangular (excepto Byssodon con
diente basal) ......................................................................................................................... 3
3(2) Con pelos en sector basal de R ....................................................................................... 4
– Sin pelos en sector basal de R ......................................................................................... 9
4(3) Coloración gris polinosa; área clara del escudo delimitada a 1 banda mediana y
1+1 banda sublateral (Fig. 29 A); abdomen negruzco o grisáceo con 1+1 bandas
plateadas transversales (Fig. 28 A ), o sobre el borde posterior de los tergitos III-
IV (Fig. 51 A) ................................................................................ Pternaspatha (en parte)
– Coloración negruzca; escudo con o sin bandas submedianas o manchas en área
anterior claras; abdomen negruzco sin bandas plateadas transversales sobre borde
posterior de tergitos III-IV .............................................................................................. 5
5(4) Paraprocto de forma subtriangular, alrededor de 2 veces más largo que ancho en
la base (Figs. 57 D, N); porción basal del cibario liso ............................... Chirostilbia
– Paraprocto de forma no subtriangular, curvado distalmente, un poco más largo
que ancho en la base (Figs. 64 G, L, P); cibario liso o con dientes ......................... 6
6(5) Porción basal del cibario con dientes dispuestos en un grupo mediano elevado
subcónico (Fig. 64 M) ......................................................................................Inaequalium
– Porción basal del cibario generalmente liso o con dentículos pequeños: si
presentes, están dispuestos en una elevación mediana, el escudo tiene 1+1
manchas anteriores claras o 3 bandas longitudinales separadas por bandas
nacaradas que cambian de tonalidad con diferente posición de la luz ................... 7
7(6) Porción basal del cibario con dientes muy pequeños o sin ellos (Figs. 102 C, F, G,
L); paraproctos relativamente cortos no lobulados, escasamente aguzados
apicalmente y dirigidos hacia abajo (Figs. 103 B, D, G, J) ......... Aspathia (en parte)
– Porción basal del cibario con una hilera de dientes (Figs. 128 D, H), o sin dientes
pero con bordes reforzados (Figs. 132 B, L); paraprocto relativamente largo,
lobulado y escasamente esclerotizado, no aguzado (Figs. 128 C, E, J; 132 D, G, N)
................................................................................................................................................ 8
8(7) Pelos del escudo agrupados simulando escamas (Fig. 132 I), sin manchas anteriores
grisáceas subcuadradas; cibario liso (Figs. 132 B, L) ................ Thyrsopelma (en parte)
– Pelos del escudo homogéneamente distribuidos; con manchas anteriores grisáceas
claras subcuadradas (Fig. 128 A); cibario con dientes dispuestos en una hilera
(Figs. 128 D, H) ............................................................................ Trichodagmia (en parte)
9(3) Porción basal del cibario sin dientes ............................................................................ 10
– Porción basal del cibario con dientes .......................................................................... 16
10(9) Pelos del escudo agrupados simulando escamas (Fig. 5 A); uña sin diente subbasal
o muy reducido (Figs. 54 D, E); triángulo fronto-ocular ausente; paraproctos muy
finos distalmente (Figs. 54 I, Q, X) ................................................................. Notolepria
– Pelos del escudo homogéneamente distribuidos; uña con o sin diente subbasal
triángulo fronto-ocular bien desarrollado; paraproctos no finos distalmente .... 11
11(10) Especies gris polinoso; abdomen grisáceo con tergitos III-V negruzcos o
castaño aterciopelado, generalmente con una mancha plateada mediana anterior
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 127

o con 1+1 manchas plateadas laterales; calcipala frecuentemente reducida o


ausente ................................................................................................................................ 12
– Especies negruzcas, abdomen negruzco sin manchas plateadas en tergitos III-V;
calcipala bien desarrollada .............................................................................................. 13
12(11) Uñas con diente subbasal (Fig. 51 F); escudo con áreas negruzcas limitadas a
bandas grisáceas mediana y sublaterales; tergitos abdominales III-V negruzcos
con 1+1 manchas blanquecinas sublaterales posteriores (Fig. 51 A); paraproctos
de forma subrectangular, más cortos que anchos (Fig. 51 H) ............... Pternaspatha
– Uñas sin diente subbasal (Fig. 99 C); escudo grisáceo uniforme, excepto 1+1
manchas anteriores medianas; tergitos III-V negruzcos o castaño aterciopelado,
con o sin manchas en área mediana anterior (Figs. 98 A, P; 99 A); paraproctos de
forma subtriangular (Fig. 99 G) tan largos como anchos ...........................................
................................................................................... Psilopelmia (BLANCASI sp. grupo)
13(11) Paraprocto subtriangular con borde distal arqueado (Figs. 118 A, B, ); escudo con 1+1
manchas anteriores submedianas continuadas posteriormente por una fina banda,
uniéndose con área posterior grisácea .................... Hemicnetha (OVIEDOI sp. grupo)
– Paraproctos de forma no subtriangular, con borde distal truncado o aguzado
posteriormente; escudo de ornamentación variable ................................................. 14
14(13) Paraprocto con borde distal truncado y suavemente aguzado posteriormente
(Figs. 103 B, K); escudo con una banda negra mediana y 1+1 submediana
longitudinales limitadas por bandas nacaradas que invierten la coloración con el
cambio de posición de la luz (Figs. 101 A, B, F, G) ...................................... Aspathia
– Paraprocto con borde distal truncado o redondeado pero no aguzado
posteriormente; escudo sin bandas plateadas contrastantes (Figs. 113 A, D, G, H;
120 A, B, K, L) ................................................................................................................. 15
15(14) Paraprocto tan largo como ancho en la base, con superficie externa mayormente
desnuda, brillante y con depresiones, y borde distal truncado (Fig. 120 I);
gonapófisis tan largas como anchas .....................................................................Hearlea
– Paraprocto más largo que ancho en la base, superficie externa mayormente
cubierta por microtriquias, sin depresiones y con borde distal curvado (Figs. 113
C, F, L, M, R, U); gonapófisis más larga que ancha (Figs. 113 C, K, Q, T, V) .......
.................................................. Hemicnetha (en parte BRACHYCLADUM sp. grupo)
16(9) Escudo con abundante polinosidad gris, mostrando, con cierta iluminación, 3-5
bandas claras; cibario con suave concavidad cubierta por dientes pequeños agudos
similares; paraprocto subtriangular con borde distal curvado, generalmente más
largo que ancho y a veces más fino distalmente; uñas sin dientes ................ Psilozia
– Escudo con escasa polinosidad, con o sin 1+1 banda plateada longitudinal; cibario
con o sin dentículos, generalmente con una depresión mediana; paraprocto no
subtriangular, generalmente más largo que ancho y a veces más fino distalmente;
uñas con o sin diente ....................................................................................................... 17
17(16) Uñas con gran diente basal subovoidal; borde posterior de los tergitos III-V
frecuentemente con 1+1 banda plateada transversa ...................................... Byssodon
– Uña con pequeño diente subtriangular subbasal o ausente; tergitos III-V sin banda
plateada transversal .......................................................................................................... 18
128 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

18(17) Cibario generalmente sin dentículos, si los hay no están presentes en la depresión
mediana, y con dentículos a los costados (Figs. 102 F, G, K, O ); escudo con 1+1
bandas plateadas submedianas, longitudinales, alcanzando el área grisácea
posterior (Figs. 101 A, F) u obscuras con cambio de dirección de luz (Figs. 101 B,
G); triángulo frontoocular bien desarrollado .................................................. Aspathia
– Cibario con depresión mediana esclerotizada o no, con dientes pequeños a los
costados (Figs. 71 P; 76 N; 80 D; 84 M; 91 Q); escudo con o sin banda plateada
longitudinal que pueden alcanzar al área posterior plateada; triángulo frontoocular
generalmente ausente o reducido ................................................................................. 19
19(18) Cibario con área central bien esclerotizada y con pequeños dientes redondeados
(Figs. 80 D, P, T); suturas infrafrontales bien desarrolladas (Figs. 80 A, V) ...........
................................................................... Coscaroniellum (QUADRIFIDUM sp. grupo)
– Cibario con área central no esclerotizada y con agudos dientes (Figs. 71 P; 76 N,
X); suturas infrafrontales no desarrolladas ................................................................. 20
20(19) Escudo negro con o sin bandas submedianas, más frecuente en borde anterior y
márgenes laterales (Figs. 71 L, O, Q, U), cuando llegan al borde posterior no
presentan manchas sombreadas sobre las bandas plateadas .............. Psaroniocompsa
– Escudo grisáceo a negro, con banda plateada alcanzando al área grisácea
posterior, presentando manchas sombreadas sobre las bandas plateadas (Figs. 76
A-F, H, I, L, M, Q, R) ..................................................................................................... 21
21(20) Cibario con dientes de aproximadamente del mismo tamaño y ausentes de la
depresión mediana (Figs. 76 N, X); triángulo frontoocular muy corto (Figs. 76 G,
W); abdomen sin manchas plateadas transversas sobre tergitos III-V, o muy cortas
(Fig. 76 K) ........................................................................................................ Cerqueirellum
– Cibario con dientes submedianos notoriamente más grandes que los restantes, y
con pequeños dientes sobre área mediana (Fig. 80 Y); triángulo frontoocular
relativamente profundo; abdomen con manchas plateadas transversas sobre
tergitos III-V de aproximadamente 1/3 del largo del tergito (Fig. 80 X); escudo
con bandas plateadas submedianas angostas (Fig. 80 W) ............................................
.......................................................... Coscaroniellum (QUADRIVITTATUM sp. grupo)
22(1) Especies amarillo rojizo a gris verdoso; si el escudo es negro, los márgenes son
amarillentos (Figs. 84 B-D); paraprocto subtriangular (Figs. 85 D-P), más largo
que ancho en la base y distalmente relativamente fino (Figs. 92 A-N) ................ 23
– Especies castaño rojizo a grisáceo oscuro; escudo generalmente ornamentado en
forma de lira (Fig. 128 A); paraprocto subrectangullar y no aguzado distalmente
(Figs. 128 C, E, J) ............................................................................................................. 28
23(22) Escudo gris verdoso a castaño amarillento; cibario liso, no esclerotizado (Fig. 60
D) .................................................................... Chirostilbia (SUBPALLIDUM sp. grupo)
– Escudo anaranjado o rojizo, o negro bordeado de amarillo y frecuentemente con
bandas plateadas submedianas; cibario con bordes esclerotizados o con pequeños
dientes ................................................................................................................................. 24
24(23) Escudo con 1+1 o más bandas longitudinales nacaradas o plateadas (Figs. 91 B-
E); abdomen generalmente con 3 hileras longitudinales de manchas negruzcas
subcuadradas (Figs. 91 G-I ); cibario con depresión mediana bien esclerotizada
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 129

bordeada de suaves lobulaciones y a los costados con dientes sobre áreas


suavemente elevadas (Figs. 91 J-Q); paraproctos más finos distalmente (Figs. 92
A-N) .........................................................................Psilopelmia (ESCOMELI sp. grupo)
– Escudo generalmente sin 1+1 o más bandas longitudinales nacaradas o plateadas
(Figs. 84 A-E); abdomen sin hileras de manchas negruscas subcuadradas; cibario
sin o con dientes generalmente más grandes y con diferente disposición que en el
dilema opuesto (Figs. 84 I-W); paraproctos no finos distalmente (Figs. 85 D-K) .
...................................................................................................................Ectemnaspis ......25
25(24) Escudo amarillo a negro, en el último caso totalmente bordeado de amarillo (Figs.
84 A-E); cibario sin o con dientes, en el último caso con placas o pequeñas
lobulaciones en el área mediana (Figs. 84 I-P) .......................................................... 26
– Escudo amarillo o rojizo; cibario con dientes pero área mediana sin placas ......... 27
26(25) Cibario sin dientes pero con borde frontal bien esclerotizado (Figs. 84 I-J) ..........
............................................................................................... BICOLORATUM sp. grupo
– Cibario con dientes y área mediana con placas o lobulaciones (Figs. 84 K-P) ......
.......................................................................................................... ROMANAI sp. grupo
27(25) Cibario con dientes bien desarrollados y agrupados mostrando 1+1 elevaciones
suaves sobre el área mediana (Figs. 80 Q-S) ................... PERFLAVUM sp. grupo
– Cibario con dientes muy pequeños y con 1+1 elevaciones grandes, generalmente
con dientes pequeños en el ápice (Figs. 84 T-W) ................... DINELLII sp. grupo
28(22) Gonapófisis subovoidales más cortas que anchas (Fig. 132 M ); paraprocto
suavemente aguzado distalmente (Fig. 132 N); pelos del escudo agrupados (Fig.
132 I) y ordenados en hileras longitudinales .......................... Trichodagmia (en parte)
– Gonapófisis subtriangulares o subovoidales, más largas que anchas, o
aproximadamente tan largas como anchas; paraproctos no aguzados distalmente;
pelos del escudo homogéneamente distribuidos o en grupos, pero no dispuestos
en hileras longitudinales .................................................................................................. 29
29(28) Cibario escasamente esclerotizado sobre el área mediana y sin dientes ni espículas
(Fig. 113 B); paraproctos un poco más largos que anchos, subrectangulares y con
abundantes microtriquias (Figs. 113 C, F) ............ Hemicnetha (PAYNEI sp. grupo)
– Cibario bien esclerotizado sobre el área mediana, con finos dientes aguzados y
escasamente esclerotizados (Figs. 113 J; 128 D, H); paraprocto aproximadamente 2
veces más largo que ancho y con escasas microtriquias (Figs. 113 L, M; 128 C, E, J)
.............................................................................................................................................. 30
30(29) Gonapófisis más de 2 veces más larga que ancha, aguzadas apicalmente (Fig. 113
K); cibario con pequeños dientes y dispuestos desordenadamente (Fig. 113 J) .....
...................................................................Hemicnetha (BRACHYCLADUM sp. grupo)
– Gonapófisis menos de 2 veces más larga que ancha, y con ápice romo (Figs. 128 B,
I); cibario con dientes bien evidentes, dispuestos a lo largo del borde frontal
extendidos en la base de los brazos laterales (Figs. 128 D, H) .............. Trichodagmia

Machos
1 Coloración general negra, a veces el abdomen con tonalidad verdosa; con o sin
pelos en sector basal de R ................................................................................................ 2
130 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

– Coloración general variable, desde castaño oscuro a amarillento; generalmente


con pelos en sector basal de R ...................................................................................... 23
2(1) Con pelos en sector basal de R ....................................................................................... 3
– Sin pelos en sector basal de R ....................................................................................... 11
3(2) Gonostilo más largo que el gonocoxito (Figs. 133 B, C, H, J); placa ventral
relativamente corta y con o sin una prolongación mediana y carena (Figs. 133 D, I, K)
................................................................................................................................................ 4
– Gonostilo más corto que el gonocoxito; placa ventral de forma variable ............. 5
4(3) Escudo sin pelos homogéneamente dispuestos y sin manchas plateadas; placa
ventral distalmente con pequeña proyección mediana y sin proyecciones laterales
(Fig. 129 E) ....................................................................................................... Trichodagmia
– Escudo con pelos dispuestos en grupos aparentando escamas, y con 1+1 mancha
anterolateral o continuando submedianamente hasta el área grisácea posterior
(Figs. 133 A, G); placa ventral distalmente con prolongaciones mediana y laterales
(Figs. 133 D, I) .................................................................................................. Thyrsopelma
5(3) Gonostilo sin o con muy reducido espolón distal ............................. Chirostilbia......6
– Gonostilo con espolón distal ........................................................................................... 7
6(5) Gonostilo sin cresta lateral (Figs. 58 B, F, G, I, L, O) ........ PERTINAX sp. grupo
– Gonostilo con cresta lateral (Figs. 61 D, F, H) ..............................................................
............................................................................. SUBPALLIDUM sp. grupo (en parte)
7(5) Escudo sin ornamentación; gonostilo achatado y curvo, con lóbulo interno
subtriangular ............................................................................... Eusimulium, Nevermannia
– Escudo generalmente ornamentado con manchas anteriores claras; gonostilo de
variada forma ...................................................................................................................... 8
8(7) Gonostilo (visto en preparación microscópica) subcuadradado con concavidad
distal y espolón subapical (Figs. 28 N; 30 R; 41 I) .......................... Pternaspatha......9
– Gonostilo (visto en preparación microscópica) de forma subtriangular a
subtrapezoidal o cilíndrica y con espolón apical ....................................................... 10
9(8) Gonostilo con tubérculos (Figs. 28 N; 30 R; 31 S) ......................................................
.........................................................................................NIGRISTRIGATUM sp. grupo
– Gonostilo sin tubérculos (Figs. 41 I; 47 K; 52 K) ........... NEMORALE sp. grupo
10(8) Gonostilo subtriangular o subtrapezoidal, con un espolón apical (Figs. 65 D, F-
H, J-N, P); placa ventral sin constricciones laterales (Figs. 65 C, E, I, O) ...........
...............................................................................................................................Inaequalium
– Gonostilo subcilíndrico con suave curvatura distal y un espolón apical (Figs. 86
H-N) (excepto wolffhuegeli, donde es relativamente corto y con varios espolones
apicales; placa ventral con suave constricción lateral (Figs. 87 A-H) .......................
............................................ Ectemnaspis (BICOLORATUM y ROMANAI sp. grupo)
11(2) Gonostilo notoriamente más corto que el gonocoxito ............................................ 12
– Gonostilo similar o más largo que el gonocoxito ..................................................... 19
12(11) Gonostilo sin espolón apical ......................................................................................... 13
– Gonostilo con espolón apical ........................................................................................ 14
13(12) Escudo sin ornamentación; gonostilo subovoidal, con borde reforzado (Figs. 54
N, S, T) .................................................................................................................. Notolepria
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 131

– Escudo con 1+1 manchas plateadas submedianas (Figs. 61 A, B, ); gonostilo


subtrapezoidal con cresta sublateral (Figs. 61 D, F, H, I) ............................................
......................................................................... Chirostilbia (SUBPALLIDUM sp. grupo)
14(12) Escudo con 1+1 banda plateada submediana alcanzando o no área posterior
grisácea; gonostilo subtriangular o subtrapezoidal ................................................... 15
– Escudo sin 1+1 banda plateada submediana, pero presentando manchas plateadas
anteriores subcuadradas o subtriangulares (Figs. 86 A-D; 91 A-D); gonostilo
subrectangular o subcilíndrico ....................................................................................... 17
15(14) Gonostilo aproximadamente 2 veces más largo que ancho en la base; espolón
apical espatulado (Figs. 81 C, G, H) ......................................................... Coscaroniellum
– Gonostilo más corto y solamente más largo que ancho en la base; espolón
terminal subcónico ........................................................................................................... 16
16(15) Gonostilo subtriangular con espolón apical o subapical (Figs. 77 J, O; 81 L) ........
.................................................................................. Byssodon, Cerqueirellum, Coscaroniellum
(QUADRIVITTATUM sp. grupo; Psaroniocompsa (SIOLII sp. grupo)
– Gonostilo subcuadrado o subtrapezoidal con espolón submediano (Figs. 72 F, H, I)
.................................................................. Psaroniocompsa (INCRUSTATUM sp. grupo)
17(14) Gonostilo generalmente subcilíndrico, con suave curvatura dorsal y un espolón
apical (Figs. 86 H-N; 93 K; 118 F, H) .............................................................................
............................................ Ectemnaspis (BICOLORATUM y ROMANAI sp. grupo),
Psilopelmia y Hemicnetha (OVIEDOI sp. grupo)
– Gonostilo subtrapezoidal con 2-15 espolones marginales ...................................... 18
18(17) Placa ventral más de 2 veces ancha que larga (Fig. 87 I); gonostilo con 7-15
espolones en el margen distal (Fig. 86 O) ........ Ectemnaspis (ROMANAI sp. grupo)
– Placa ventral tan ancha como larga; gonostilo con 2-4 espolones apicales .............
...................................................................................................................................... Psilozia
19(11) Gonostilo subcilíndrico y fino basalmente, generalmente más angosto que la
mitad del gonocoxito, borde distal ancho y generalmente con una protuberancia
basal ..................................................................................................................................... 20
– Gonostilo basalmente más ancho que la mitad del gonocoxito, borde distal ancho
y sin protuberancia basal ................................................................................................ 21
20(19) Placa ventral generalmente haciéndose más angosta medio distalmente (Figs. 106
M-P); gonostilo con protuberancia basal (Figs. 105 B, C, E-H, J-L) ......... Aspathia
– Placa ventral no haciéndose más angosta medio distalmente (Fig. 121 D, G );
gonostilo sin protuberancia basal pero con pequeña carena basal (Figs. 121 C, F)
...................................................................................................................................... Hearlea
21(19) Gonostilo achatado, flexuoso y escasamente afinado distalmente (Figs. 114 A,
E, G, I, M, N); endoparámeros con largos dientes y base bien esclerotizada (Fig.
114 K) ........................................................................................................ Hemicnetha......22
– Gonostilo subtriangular afinándose distalmente y notoriamente curvo (Figs. 133
B, C, H, J); endoparámeros sin dientes y base no bien esclerotizada (Fig. 133 F) .
.............................................................................................................................. Thyrsopelma
22(21) Placa ventral notoriamente más ancha que larga, con una protuberancia mediana
ancha (Figs. 114 B, C, F, H, L) .............. PAYNEI y BRACHYCLADUM sp. grupo
132 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

– Placa ventral aproximadamente tan larga como ancha, sin protuberancia mediana,
pero con carena mediana (Figs. 114 J, O) ....................... MEXICANUM sp. grupo
23(1) Gonostilo cerca de 2 veces más largo que el largo del gonocoxito ...................... 24
– Gonostilo más corto o escasamente más largo que el largo del gonocoxito ...... 28
24(23) Gonostilo con 3-6 espolones o más (Fig. 129 L); endoparámeros sin dientes (Fig.
129 C) ............................................................................................. Trichodagmia (en parte)
– Gonostilo con 1 espolón o ausente; endoparámeros generalmente con dientes
grandes (Figs. 61 I; 99 I) ................................................................................................. 25
25(24) Sector basal de R sin pelos; escudo gris opaco; gonostilo con borde interno recto
(Figs. 98 V; 99 H) .................................................. Psilopelmia (BLANCASI sp. grupo)
– Sector basal de R con pelos (Fig. 60 C); escudo no gris opaco; gonostilo con borde
interno curvo .................................................................................................................... 26
26(25) Gonostilo sin espolón apical, con cresta sublateral no curvada distalmente (Figs.
61 D, F, H, I) ................................................ Chirostilbia (SUBPALLIDUM sp. grupo)
– Gonostilo con espolón apical, sin cresta sublateral y suavemente curvada
distalmente ......................................................................................................................... 27
27(26) Placa ventral generalmente más larga que ancha en la base (Figs. 87 J-L); gonostilo sin
curvatura distal (Figs. 86 P, Q, R) .................. Ectemnaspis (PERFLAVUM sp. grupo)
– Placa ventral generalmente más corta que ancha en la base (Figs. 87 M-O; 94 A-
L); gonostilo generalmente con curvatura distal (Figs. 86 S; 93 H-R) .....................
............................................................... Psilopelmia, Ectemnaspis (DINELLII sp. grupo)
28(23) Escudo castaño oscuro con pelos agrupados simulando escamas, gonostilo
relativamente grueso distalmente (Fig. 133 J) ............................................ Thyrsopelma
– Escudo castaño rojizo con pelos no agrupados; gonostilo relativamente fino
distalmente (Figs. 129 B, D) .......................................................................... Trichodagmia

Pupas
1 Branquias con ramas relativamente gruesas con ápice agudo y bien esclerotizado
en forma de espina, con 12-50 ramas (Figs. 134 B, E, G, K); capullo resistente y
proyectado ventralmente hacia delante .................................... Thyrsopelma (en parte)
– Branquias de forma variable pero sin ápice agudo bien esclerotizado en forma de
espina; capullo de forma variable ................................................................................... 2
2(2) Branquia notoriamente gruesas, asimétricas, generalmente con curvaturas .......... 3
– Branquias con ramas generalmente filiformes, simétricas y sin curvaturas laterales
................................................................................................................................................ 6
3(2) Frontoclípeo y porción expuesta del tórax con tricomas simples (Fig. 121 H);
branquias con estriaciones (Figs. 122 A-K; 123 A-O; 124 A-K); capullo sin
prolongación dorsal .................................................................................................Hearlea
– Frontoclípeo y porción expuesta del tórax con tricomas multirramosos con 2-8
ramas; branquias sin estriaciones; capullo generalmente con una prolongación
dorsal .................................................................................................................................... 4
4(3) Capullo generalmente sin prolongación dorsal, borde anterior recto (Figs. 66 A, B,
D, L; 70 A, B, P, Q) ...........................................................................................Inaequalium
– Capullo prolongado dorsalmente (Figs. 73 H-K; 88 C, D, F; 107 I) ...................... 5
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 133

5(4) Tricomas del frontoclípeo y tórax bífidos (Figs. 73 O-Q) ................. Psaroniocompsa
– Tricomas del frontoclípeo y tórax con 3-8 ramas .................... Ectemnaspis, Aspathia
6(2) Capullo con tejido resistente y porción anterior elevada, generalmente
protegiendo la base de la branquia (Figs. 59 L, O, S; 115 A, D); filamentos de las
branquias relativamente gruesos ..................................................................................... 7
– Capullo con tejido no resistente y generalmente con la porción anterior no elevada
protegiendo la branquia; filamentos de las branquias relativamente finos .......... 13
7(6) Abertura anterior del capullo con festones protegiendo la base de la branquia
(Figs. 115 A-D) ................................................................................................................... 8
– Abertura anterior del capullo sin festones .................................................................... 9
8(7) Branquia con 8 ramas, frecuentemente con el ápice romo y resistente (Figs. 116 A,
B); frontoclípeo generalmente reforzado en la base y agrandado en el área facial,
mostrando las hembras y machos similar ancho del área facial (Figs. 115 O-T) ...
........................................................................................ Hemicnetha (PAYNEI sp. grupo)
– Branquias con 10 ramas frecuentemente con el ápice romo no resistente (Fig. 59 N);
frontoclípeo no reforzado en la base y no agrandado en área facial, mostrando el
macho la base de la frente más angosta que la hembra ........... Chirostilbia (en parte)
9(7) Branquia con 6 filamentos ............................................................................................. 10
– Branquia con más de 6 filamentos ............................................................................... 11
10(9) Frontoclípeo y tórax con numerosos tubérculos, algunos de ellos aguzados como
cortas espinas (Figs. 119 B, D, E); capullo escasamente elevado anteriormente
(Fig. 119 A) ................................................................Hemicnetha (OVIEDOI sp. grupo)
– Frontoclípeo y tórax sin tubérculos (Fig. 115 R); capullo frecuentemente muy
elevado anteriormente (Fig. 115 E) ...............................................................................
............................................................... Hemicnetha (BRACHYCLADUM sp. grupo)
11(10) Branquia con 18-20 ramas ordenadas como un ramo (Fig. 130 L) ...........................
.........................................................................................................Trichodagmia ( en parte)
– Branquias con 8-16 ramas .............................................................................................. 12
12(11) Branquia con 12 ramas .......................................................................................................
................................. Hemicnetha (MEXICANUM sp. grupo), Trichodagmia (en parte)
– Branquias con 10 ramas ......................................................................................................
................................................ Hemicnethha (BRACHYCLADUM sp. grupo en parte)
13(6) Branquia con 4 ramas filamentosas .............................................................................. 14
– Branquia con más de 4 ramas filamentosas ............................................................... 17
14(13) Tubérculos de frontoclípeo y tórax aguzados (Fig. 82 D) .................. Coscaroniellum
– Tubérculos de frontoclípeo y tórax no aguzados ...................................................... 15
15(14) Capullo reducido ventralmente (Figs. 98 X; 99 K); tricomas cefálicos y torácicos
simples (Fig. 98 a-b) .............................................. Psilopelmia (BLANCASI sp. grupo)
– Capullo no reducido ventralmente; tricomas cefálicos y torácicos con 1-8 ramas
.............................................................................................................................................. 16
16(15) Tricomas cefálicos y torácicos generalmente bífidos (Figs. 73 O, P, Q) ..................
................................................................................................. Psaroniocompsa, Cerqueirellum
– Tricomas cefálicos y torácicos con 1-8 ramas ...............................................................
................................................................... Eusimulium, Byssodon, Nevermannia Psilopelmia
134 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

17(13) Branquia con 6 ramas filamentosas .............................................................................. 18


– Branquia con más de 6 ramas filamentosas ............................................................... 20
18(17) Tricomas torácicos en número sobrepasando 10 de cada lado y frecuentemente
muy modificado (Figs. 40 U-V; 44 R; 53 S, T; 108 G-H; 109 A, G) ........................
......................................................................................... Pternaspatha, Aspathia (en parte)
– Tricomas torácicos en número de alrededor de 5 de cada lado ............................ 19
19(18) Tricomas cefálicos y torácicos bífidos (raramente simples o trífidos) ......................
............................................................. Psaroniocompsa, Cerqueirellum, Aspathia (en parte)
– Tricomas cefálicos y torácicos no bífidos generalmente con más de 3 ramas .......
................................................. Byssodon, Inaequalium, Ectemnaspis (BICOLORATUM y
ROMANAI sp. grupo ), Notolepria, Psilopelmia
20(17) Branquias con 8-10 ramas .............................................................................................. 21
– Branquias con más de 10 ramas .......................................................................................
....................... Psilozia, Chirostilbia, Aspathia Ectemnaspis (PERFLAVUM sp. grupo)
21(20) Tricomas torácicos sobrepasando en número de 10 de cada lado y frecuentemente
muy modificados (Figs. 29 N, P; 31 Y, Z); branquias con 8 ramas ......... Pternaspatha
– Tricomas torácicos en número de 5 de cada lado; branquias con 8-10 ramas .......... 22
22(21) Tejido del capullo esponjoso, con margen anterior elevado y carena dorsal
(Figs. 88 B, H, I, J) .............................................................................................................
....................... Ectemnaspis (BICOLORATUM y ROMANAI sp. grupo, en parte)
– Tejido del capullo compacto, sin margen anterior elevado y sin carena dorsal (Figs.
62 C, D, G; 78 A, G, J) .............................. Chirostilbia (SUBPALLIDUM sp. grupo),
Notolepria, Cerqueirellum, Coscaroniellum, Psilozia, Byssodon, Aspathia, Ectemnaspis
(PERFLAVUM y DINELLII sp. grupo), Psilopelmia

Larvas
1 Anillo anal con más de 150 hileras de ganchos ........................................................... 2
– Anillo anal con menos de 120 hileras de ganchos ...................................................... 6
2(1) Cutícula del cuerpo con tricomas lanceolados (Figs. 135 G, L, S); hipostomio con
diente mediano y dientes intermediarios no bien diferenciados (Figs. 135 D, I, N);
abdomen no achatado dorsalmente en su porción distal (Fig. 135 A); escleritos
cervicales agrandados, más de 2 veces más ancho que largo (Fig. 135 Q) .............
.............................................................................................................................. Thyrsopelma
– Cutícula del cuerpo sin tricomas lanceolados; hipostomio con diente mediano y
generalmente los dientes intermediarios bien diferenciados (Figs. 117 N, Q; 126 J;
131 G); abdomen dorsalmente achatado distalmente (Figs. 127 G; 131 A);
escleritos cervicales no agrandados menos de 2 veces más ancho que largo (Fig.
131 I) ..................................................................................................................................... 3
3(2) Esclerito anal frecuentemente con refuerzo esclerotizado mostrando en algunos
casos ramas ventrales agrandadas completando un anillo (Figs. 127 B-J); margen
anterior del hipostomio generalmente en forma de arco, con dientes laterales e
intermediarios reducidos (Figs. 126 A, B, I, H, J) .............................................Hearlea
– Esclerito anal sin refuerzo esclerotizado ni alargadas las ramas ventrales completando
un anillo; margen anterior del hipostomio de forma variable ....................................... 4
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 135

4(3) Apotoma cefálico oscurecido en la mitad basal mostrando como un triángulo (Fig.
63 A) .................................................................................................................... Chirostilbia
– Apotoma cefálico con diferente ornamentación ......................................................... 5
5(4) Bordes laterales del hipostomio descendiendo abruptamente desde los dientes
laterales con aspecto de meseta (Figs. 117 P, Q; 131 G); mandíbula con diente
preapical más corto que el apical (Figs. 117 L; 131 E, K) ..........................................
........................................... Hemicnetha (BRACHYCLADUM sp. grupo), Trichodagmia
– Bordes laterales del hipostomio descendiendo suavemente desde los dientes
laterales con aspecto de curva (Figs. 117 N, O, R); mandíbula con diente preapical
tan largo o más que el apical (Figs. 117 K, M) ..............................................................
........................................................ Hemicnetha (PAYNEI; MEXICANUM sp. grupo)
6(1) Anillo anal con 100-120 hileras de ganchos; rayos del abanico cefálico con dientes
de similar tamaño (Fig. 119 J); mandíbulas con un par de dientes marginales muy
grandes (Fig. 119 K) ................................................ Hemicnetha (OVIEDOI sp. grupo)
– Anillo anal con menos de 100 hileras de ganchos; rayos del abanico cefálico
alternando un grupo de pequeños dientes con alguno más grande; mandíbula con
solo un diente marginal, y si existen 2 el segundo es muy pequeño (Figs. 29 S; 79
B; 83 C, O, R; 90 Q) .......................................................................................................... 7
7(6) Apotoma cefálico sin ornamentación o reducida a una pequeña mancha en el área
medio basal; con numerosos y cortos pelos sobre la cápsula cefálica (Fig. 79 C);
generalmente con banda oscura alrededor del primer segmento abdominal; antena
fina y sobrepasando el ápice del tallo del abanico cefálico (Fig. 79H) ................... 8
– Apotoma con evidente ornamentación; sin pelos o muy escasos en cápsula
cefálica; antena generalmente gruesa y sobrepasando o no el ápice del tallo del
abanico cefálico .................................................................................................................. 9
8(7) Mandíbulas con una hilera de cerdas supramarginal inferior (Fig. 79 B); artejo
antenal mediano similar o más largo que el proximal (Figs. 79 D, E, I) ..................
............................................................................................................................ Cerqueirellum
– Mandíbulas sin una hilera de cerdas supramarginal inferior; artejo antenal
mediano aproximadamente 1/2 a 4/5 del largo del proximal (Figs. 83 B, N, Q) ..
.......................................................................................................................... Coscaroniellum
9(7) Hendidura postgenal escasamente desarrollada, con puente postgenal más largo
que el hipostomio (Figs. 112, A, B, D, G); papilas rectales sin divertículos en los
lóbulos ................................................................................................................................ 10
– Hendidura postgenal bien desarrollada con puente postgenal tan largo o más
corto que el largo del hipostomio; papilas rectales generalmente con divertículos
en los lóbulos .................................................................................................................... 13
10(9) Antena con estriación transversal y sobrepasando el ápice del tallo del abanico
cefálico (Figs. 111 A, B, H, L); artejo antenal mediano más largo que el proximal
(Figs. 111 B, H, L, M) ........................................................................................... Aspathia
– Antena sin estriación transversal y no sobrepasando el ápice del tallo del abanico
cefálico; artejo mediano más corto que el artejo proximal ..................................... 11
11(10) Noveno esternito sin 1+1 tubérculos ............... Psilopelmia (BLANCASI sp. grupo)
– Noveno esternito con 1+1 tubérculos bien evidentes ............................................. 12
136 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

12(11) Antena sobrepasando el ápice del tallo del abanico cefálico ......................................
....................................................................................................... Eusimulium, Nevermannia
– Antena llegando aproximadamente al ápice del tallo del abanico cefálico ..............
...................................................................................................................................... Psilozia
13(9) Papilas rectales sin divertículos en los lóbulos ..............................................................
................................................................................... Psilopelmia (BLANCASI sp. grupo)
– Papilas rectales con divertículos en los lóbulos ......................................................... 14
14(13) Apotoma cefálico con manchas aisladas postero medianas, antero-laterales y
postero-laterales, generalmente positivas (Figs. 30 Y; 55 C; 56 K; 75 L) ............. 15
– Apotoma cefálico con ornamentación variable diferente a la señalada en el dilema
opuesto ............................................................................................................................... 19
15(14) Esclerito anal sin escamas (Fig. 63 G) ......................................................................... 16
– Esclerito anal con escamas (Fig. 63 P) ........................................................................ 17
16(15) Antena sobrepasando el ápice del tallo del abanico cefálico (Figs. 75 D, L); artejo
antenal mediano generalmente más corto que el distal (Figs. 75 A, E, M) .............
......................................................................................................................... Psaroniocompsa
– Antena no sobrepasando el ápice del tallo del abanico cefálico; artejo antenal
mediano generalmente más largo que el distal (Figs. 90 M, N) .................................
....................... Ectemnaspis (BICOLORATUM sp grupo; PERFLAVUM sp. grupo)
17(15) Largo máximo del cuerpo por debajo de 4,5 mm ....................................... Notolepria
– Largo máximo del cuerpo por arriba de 5,0 mm ...................................................... 18
18(17) Artejo antenal mediano más corto que el apical (Fig. 63 N) ......................................
......................................................................... Chirostilbia (SUBPALLIDUM sp. grupo)
– Artejo antenal mediano más largo que el apical (Fig. 29 S) ........................................
.......................................................................................... Byssodon, Pternaspatha (en parte)
19(14) Apotoma cefálico oscurecido en la mitad basal, dejando una ventana clara sobre la
porción medio basal (Figs. 90 E, F; 97 A-G); esclerito anales sin escamas ............
........................................................................ Ectemnaspis (BICOLORATUM sp. grupo;
ROMANAI sp. grupo; DINELLII sp. grupo), Psilopelmia
– Apotoma cefálico con ornamentación variable, diferente al del dilema opuesto;
esclerito anal con o sin escamas ................................................................................... 20
20(19) Esclerito anal con escamas; artejo antenal mediano más largo que el distal (Figs.
29 S; 31 b; 35 X) ............................................................................................. Pternaspatha
– Esclerito anal sin escamas (Fig. 68 C); artejo antenal mediano más corto que el
distal (Figs. 68 D, L) ........................................................................................................ 21
21(20) Apotoma cefálico con una mancha oscura difusa mediana basal que se vuelve
más fina hacia adelante, delimitando 2 áreas claras alargadas (Figs. 68 B, M; 70 M)
...............................................................................................................................Inaequalium
– Apotoma cefálico con fuerte pigmentación basal, generalmente con forma
triangular (Fig.63 A) ............................................. Chirostilbia (PERTINAX sp. grupo)
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 137

Simulium (Eusimulium) ROUBAUD


Eusimulium ROUBAUD, 1906a: 521.
Type-species: Simulium aureum FRIES, 1824: 16.
Only a short diagnosis is given.
Female: Coloration brown to black, without scutum ornamentation; calcipala well
developed; claws with wide basal tooth lanceolate; cibarium without teeth; paraprocts
short; spermatheca sclerotization reaching spermathecal duct.
Male: Gonocoxite larger than gonostylus; gonostylus with strong curvature, subtri-
angular and 1 terminal spur; ventral plate laterally compressed.
Pupa: Cocoon slipper shaped; generally with 4 gill filaments; thoracic trichomes
generally single.
Larva: Hypostomium with well-developed teeth, with median tooth at same level as
lateral teeth. Postgenal bridge nearly 2 times hypostomium length; ninth segment with
1+1 ventral tubercles, rectal papillae without diverticula.
Distribution: This taxon is primarily Holarctic, with only one species in the
Neotropics.

Simulium (Eusimulium) donovani VARGAS, 1943b: 359-360; VARGAS & DÍAZ NÁJERA,
1957b: 164-166; DALMAT, 1955: 117; ADLER et al., 2004: 328-329.
Simulium (Eusimulium) aureum (FRIES), 1824: 16; VARGAS et al., 1946: 166; VARGAS & DÍAZ NÁJERA,
1957b: 153, 164-166, 231; DALMAT, 1955: 116-121, 153, 164- 166, 231 (misidentification).
Simulium diazi DE LEÓN, 1945: 70.
The species was treated by ADLER et al. (2004).
Distribution: USA: Arizona, California, Colorado, Nevada, New Mexico, Oregon,
Texas; Mexico: Chiapas, Durango, Guanajuato, Michoacan, Morelos, Veracruz; Guatemala.

Simulium (Nevermannia) ENDERLEIN


Nevermannia ENDERLEIN, 1921a: 199.
Type-species: Simulium annulipes BECKER, 1908: 72-73 (= Simulium ruficorne MACQUART, 1838: 88).
Only a short diagnosis is given.
Female: Coloration brown to black, without scutum ornamentation; calcipala well
developed; claws with well-developed subbasal tooth; cibarium without teeth.
Male: Gonostylus distally with medially directed, subtriangular flange. Paramere
with 1 or 2 large, distal spines. Dorsal plate well developed.
Pupa: Cocoon slipper shaped, with or without anterodorsal projection. Gill of 4-8
filaments.
Larva: Hypostomium with well-developed teeth; lateral teeth moderately enlarged.
Ninth segment with 1+1 ventral tubercles, rectal papillae with or without diverticula.
Distribution: This taxon is primarily Holarctic and Oriental, with only one species
in the Neotropics.
138 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Simulium (Nevermannia) carbunculum ADLER, CURRIE & WOOD, 2004


The species was treated by ADLER et al. (2004).
Distribution: USA: Arizona, California, Colorado, Idaho, Nevada, New Mexico,
Utah, Wyoming; Canada: British Columbia, Saskatchewan; Mexico: Durango.
Discussion: Simulium pugetense was recorded from Durango, Mexico, by VARGAS &
DÍAZ NÁJERA (1957b: 153, 164-166, 231), but is actually Simulium carbunculum ADLER,
CURRIE & WOOD, 2004, as reported by ADLER et al. (2004).

Simulium (Byssodon) ENDERLEIN


Byssodon ENDERLEIN, 1925: 209.
Type-species: Simulium forbesi MALLOCH, 1914: 63-65 (= Simulium meridionale RILEY, 1887: 513-514).
Female: Small species; scutum blackish with submedian silvery stripes in shape of
lyre; claw generally with elongate basal tooth, as in Eusimulium; cibarium generally without
teeth; radius without hair; paraproct short weakly acuminate ventrally.
Male: Gonostylus shorter than gonocoxite; relatively thin and curved with one apical
spur; ventral plate flat, wider than long; endoparameres with large teeth.
Pupa: Gill with 2-6 or 22-26 branches; cocoon slipper or shoe shaped.
Larva: Cephalic apotome without clear ornamentation, with spots positive and
negative; postgenal bridge about half of hypostomium length; hypostomial teeth with
median tooth beneath level of lateral teeth; ninth segment with or without ventral
tubercles; rectal papillae without diverticula on lobes.
Distribution: This taxon is present in the Holarctic, Oriental and Afrotropical Regions.

Simulium (Byssodon) meridionale RILEY, 1887: 513-514.


Simulium tamaulipense TOWNSEND, 1897: 171-172 described from Mexico, is considered a synonym.
Distribution: Very well represented in Canada and USA; Mexico: Tamaulipas.
Discussion: Simulium tamaulipense was described from Mexico but it is considered a
synonym of Simulium meridionale.

Simulium (Psilozia) ENDERLEIN


Psilozia ENDERLEIN, 1936: 113-114.
Type-species: Psilozia groenlandica ENDERLEIN, 1936: 114 (= Simulium vittata ZETTERSTEDT, 1838: 803.
Simulium (Neosimulium) VARGAS et al., 1946: 103, 108, 160. Type-species Simulium vittata ZETTERSTEDT, 1838.
Female: Grayish pollinose; scutum black with 5 longitudinal silvery stripes; cibarium
with small regular teeth; basal sector of R without hair; calcipala small; claws without basal
tooth; paraprocts subtriangular, slightly longer than wide.
Male: Scutum black with silvery spots anteriorly; gonostylus about half gonocoxite
length; subtrapezoidal, with 2-3 apical spurs; ventral plate subtriangular with short basal
branches.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 139

Pupa: Cocoon slipper shaped, relatively short; gills with 10-24 filaments: cephalic
and thoracic trichomes simple.
Larva: Cephalic apotome with positive spots; postgenal bridge nearly 1.5 times
longer than hypostomium; ninth segment without ventral tubercles; rectal papillae
generally without diverticula.
Discussion: This subgenus has five species in the USA; three of them are also
present in Mexico.

Simulium (Psilozia) argus WILLISTON, 1893: 253-254; VARGAS & DÍAZ NÁJERA, 1957b:
263; PETERSON & KONDRATIEF, 1995: 36-37.
Simulium kamloopsi HEARLE: VARGAS et al., 1946: 171.
Distribution: Mexico: Distrito Federal, Nueva León, Oaxaca, Puebla, San Luis
Potosi, Sonora, Veracruz; Morelos by VARGAS & DÍAZ NÁJERA, 1948b, 1949, 1957b.
Discussion: For more information see ADLER et al., 2004: 354-355.

Simulium (Psilozia) encisoi VARGAS & DÍAZ NÁJERA, 1949: 292-295; 1957b: 180-182,
233, 263, 336.
Material not revised.
Distribution: Cited in Mexico: Hidalgo, Michoacan.
Discussion: For more information see ADLER et al., 2004: 355.

Simulium (Psilozia) vittatum ZETTERSTEDT, 1838: 803; VARGAS, 1942a: 243; VARGAS
& DÍAZ NÁJERA, 1948b: 324; PETERSON & KONDRATIEFF, 1995: 37; ADLER et al., 2004:
356-361, 363-364.
Material not revised.
Distribution: Mexico: Distrito Federal, Tabasco.
Discussion: For more information see ADLER et al., 2004: 356-364.

Simulium (Pternaspatha) ENDERLEIN (Figs. 28-53; 137, 138.)


Pternaspatha ENDERLEIN, 1930: 84, 88; EDWARDS, 1931: 151; ENDERLEIN, 1934b: 274; 1936: 115;
SMART, 1945: 465, 488; VARGAS, 1945a: 102; VARGAS & DÍAZ, 1951: 143; STONE, 1962: 206, 208;
1963: 17.
Type-species: Pternaspatha nigristrigata ENDERLEIN.
Acropogon ENDERLEIN, 1934b: 276; 1936: 118; SMART, 1945: 489; VARGAS, 1945a: 85; STONE, 1962:
206; 1963: 1. Type: Acropogon barbatipes ENDERLEIN.
Dasypelmoza ENDERLEIN, 1934b: 275; SMART, 1945: 488; VARGAS, 1945a: 90; VARGAS & DÍAZ, 1951:
146; STONE, 1962: 206; 1963: 5. Type-species: Simulium varipes PHILIPPI, 1865: 634.
Adults: Antennae 11 segmented; eyes of female with fronto-ocular triangle; row of
hairs between eyes of male; hind border of cibarium of female unarmed; mandibles and
lacinia of female toothed on both sides; mesonotum of female gray, with 1+1 sublateral
140 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

and one submedian, black, longitudinal stripes, median stripe divided longitudinally by
fine grayish line; scutum of male black, bordered with gray; anterior border with 1+1
small, white, submedian spots; delicate, white, central line often visible on anterior half of
disc of mesonotum; pleural tuft present; setae lacking from pleural membrane and
sternopleurum; costa with both hair and spinelike setae; radial sector not branched; basal
portion of R bare, very rarely with hairs dorsally; R1 with both hair and spiniform setae;
numerous long hairs on fore coxa, femur, and tibia of male; fore tarsi narrow; hind
basitarsus of female narrow; of male, from narrow to conspicuously widened, spindle-
shaped; calcipala small to absent; pedisulcus well developed; claws of female with small
subbasal tooth; basal fringe of abdomen of male exceptionally long; last tergites of female
dull; gonostylus subquadrate, shorter than gonocoxite, with single apical spine; posterior
surface glabrous or granulose; ventral plate transverse, in many instances with 1+1
translucent areas; basal arms short; median sclerite elongate, widened and deeply incised
on distal half; spermatheca subglobular; outer surface not sculptured; inner surface with
minute spicules arranged in groups of one to five; genital fork slender; stem strongly
pigmented; arms not pigmented; apical expansions of arms partly translucent, partly
pigmented; paraprocts short, rather rounded or subtriangular.
Pupa: Cocoon from slipper shaped to shoe shaped, with distinct anteroventral
bridge; close-woven, threads not conspicuous; rim of aperture slightly reinforced, not
festooned; gills shorter than pupa, with six or eight slender filaments; their surface with
granules forming spiral pattern; apex of filaments rounded; exposed portion of thorax
with numerous trichomes, from approximately 10+10 to more than 100+100; cephalic
and thoracic trichomes of greatly varied structure: hair like, spatulate, ribbon-shaped,
simple or branched; all trichomes either of single type or various types intermixed;
abdominal tergite II with 3+3, rarely with 4+4, simple spines; tergites III-IV with 4+4
simple spines; tergites VI-IX on anterior half with transverse rows of small, spinelike or
scale-like tubercles; sternites V-VII with 2+2 strong, simple or bifid spines; sternites III-
VIII on anterior portion of disc with 1+1 fields of minute scales; apical hook of
abdomen not developed.
Larva: Thorax slightly swollen; abdomen gradually expanded, slightly clubbed
posteriorly; no ventral tubercles at posterior of abdomen; body surface glabrous, or with
only very sparse, simple setae; setae more numerous at base of anal sclerite; hypostomial
setae arranged in two irregular series on each side, very rarely only in one series; postgenal
cleft subtriangular, narrowed apically; postgenal bridge from well developed to extremely
short, but postgenal cleft in no case completely attaining hypostomial groove; anal ring
with 80-90 rows composed of 13 to 20 hooks each (Figs. 137, 138).
Distribution: Cool and cold temperate western and extreme southern South
America, north as far as central Ecuador.

KEYS TO SIMULIUM (PTERNASPATHA) SPECIES

Females
The female of Simulium (Pt.) bordai is not known, and we do not have sufficient
information to include Simulium (Pt.) schoenemanni in the key.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 141

1 Basal portion of R bare (Fig. 39 I) ................................................................................. 2


– Basal portion of R setose (Fig. 51 D) ............................................ Simulium annulatum
2(1) Abdomen (Fig. 36 C) dull silver-gray; each of terga II-V with relatively small,
central, velvety black spot, and with lateral dark spots absent; calcipala very small
(Fig. 36 G) ...........................................................................................Simulium strigidorsum
– Abdominal terga II-V with more extensive dark markings; calcipala absent or
present .................................................................................................................................. 3
3(2) Calcipala absent (Figs. 32 A; 50 J) .................................................................................. 4
– Calcipala present, although very small in some species ............................................. 9
4(3) Terga IV and V without 1+1 silver-white spots, entirely black, although in some
specimens narrowly margined with white along hind borders (Fig. 50 A) ..............
....................................................................................................... Simulium prodexargenteum
– Terga IV and V with 1+1 large silver-white spots (Fig. 28 A) ................................. 5
5(4) Claws without subbasal tooth (Fig. 32 B) ..................................... Simulium horcochuspi
– Claws with subbasal tooth (Fig. 28 J) ............................................................................ 6
6(5) Abdominal tergites III-V with 1+1 well-developed lateral blackish spots meeting
anteriorly with median spot (Figs. 28 A; 29 B); wing length 2.3-3.5 mm .............. 7
– Abdominal tergites III-V with 1+1 small isolated lateral blackish spots (Figs. 35 B;
37 B); wing length 3.3-3.7 mm ........................................................................................ 8
7(6) Abdominal tergites III-V with light grayish coloration extending to lateral area of
tergites (Fig. 29 B); fore and mid femora light brown, not conspicuously darkened
apically; wing length 2.8-3.5 mm ............................................................. Simulium limay
– Abdominal tergites III-V with light grayish coloration extending or not to lateral
area of tergites (Fig. 28 A); fore and mid femora conspicuously darkened at apex;
wing length 2.3-2.7 mm ................................................................ Simulium nigristrigatum
8(6) Abdominal tergite VI with median black spot (Fig. 37 B); 1+1 lateral black spots
on tergites III-V reaching to posterior border; length/width ratio of hind
basitarsus = 7.2-7.6 ............................................................................. Simulium deagostinii
– Abdominal tergite VI without median blackish spot (Fig. 35 B); 1+1 lateral black
spots on tergites III-V not reaching posterior border; length/width ratio of hind
basitarsus = 6.6 ................................................................................Simulium diamantinum
9(3) Tergum III without 1+1 light-colored spots, thus terga III-V entirely dark except
in some species with narrow white line along lateral portions of hind margin (Fig.
51 A); mid-longitudinal light-colored line of scutum invisible or only faintly
perceptible ......................................................................................................................... 10
– Tergum III with 1+1 gray or whitish spots (Figs. 47 A; 48 A; 49 A; 50 A), in some
specimens small and not easily seen but in all cases perceptible; central light-
colored line of scutum invariably clearly visible ....................................................... 11
10(11) Bases of tibiae conspicuously pigmented, but apex of hind basitarsus lacking
conspicuous pigment; terga III-V entirely dark (Fig. 52 A); paraprocts roundly
salient; cerci relatively high (Fig. 52 H) .................................................. Simulium caprii
– Bases of tibiae lacking distinct pigment, with hind basitarsus conspicuously
pigmented on apical third (Fig. 53 H); posterior border of terga III-V narrowly
142 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

bordered with gray (Fig. 53 A); paraprocts very short and truncate, cerci relatively
low (Fig. 53 L) ....................................................................................... Simulium stelliferum
11(9) Terga IV and V lacking 1+1 gray or white spots, only in some cases their hind
margins narrowly bordered with white laterally (Figs. 47 A; 50 A); tibiae with
distinct dark basal or subbasal annulus ....................................................................... 12
– Terga IV and V, in some cases only tergum V, with 1+1 light-colored spots of
varied sizes (Figs. 33 A; 38 C; 40 A; 41 A); tibiae with or without dark subbasal or
basal annulus ..................................................................................................................... 16
12(11) Calcipala not well developed (Fig. 50 J) ................. (some)......Simulium prodexargenteum
– Calcipala well developed, distinct (Figs. 39 K; 48 C) ............................................... 13
13(12) Dark area separating white spots on abdominal tergum II very narrow, acuminate
anteriorly (Fig. 48 A); tergum VI entirely silver-white ............... Simulium albicinctum
– Dark area separating white spots on abdominal tergum II wide (Figs. 39 B; 47 A;
49 A), widened anteriorly or as wide anteriorly as posteriorly; tergum VI dark at
center and laterally ........................................................................................................... 14
14(13) Setae and spines on R1 arranged in several irregular rows (Fig. 39 I) ......................
.................................................................................................................... Simulium nemorale
– Setae and spines on R1 arranged in single row (similar to Fig. 42 P) ................... 15
15(14) Calcipala very small, narrowly triangular (Fig. 49 E) ...........................Simulium luchoi
– Calcipala larger, rounded (Fig. 47 F) ............................................Simulium hectorvargasi
16(11) Frons very wide (Fig. 38 D); paraproct with ventral posterior angle elongated (Fig.
38 N); scutum black, with variable ornamentation in accordance with light
incidence (Figs. 38 A, B) ....................................................................... Simulium cotopaxi
– Frons not wide (Fig. 40 B); paraproct with ventral posterior angle rounded (Fig. 40
L); scutum grayish, with median and 1+1 sublateral stripes blackish .................. 17
17(16) Tibiae not conspicuously darkened at base (Figs. 40 F, G, H) ............................... 18
– Tibiae conspicuously darkened at base ....................................................................... 23
18(17) 1+1 light-colored spots on terga III-V very small, in some cases difficult to
perceive (Fig. 40 A; 41 A); wing length 2.9-3.9 mm ................................................. 19
– 1+1 light-colored spots on terga III-V larger, all distinct (Fig. 33 A); wing length
2.5-3.6 mm ......................................................................................................................... 20
19(18) Light submedian spots on tergites III-V crossing from anterior to posterior
border of segment and with parallel sides (Fig. 40 A); claws with small subbasal
tooth (Fig. 40 J) ................................................................................... Simulium barbatipes
– Light submedian spots on tergites III-V not reaching anterior border, and with
enlarged base (Fig. 41 A); claws without subbasal tooth (Fig. 41 F) ........................
.................................................................................................................... Simulium philippii
20(18) Calcipala well developed (Fig. 33 E) ............................................................................ 21
– Calcipala minute (Fig. 31 I) ............................................................................................ 22
21(20) Wing length 3.2-3.5 mm; hind tibia with pigment often only at extreme apex ......
......................................................................................................................... Simulium simile
– Wing length 2.6-2.8 mm; pigment of hind tibiae always more widely extended, on
apical two-fifths of segment (Fig. 34 D) .......................................... Simulium pulchrum
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 143

22(20) Fronto-ocular triangle longer than wide; wing length greater than 3.0 mm; Sc with
approximately 10 hairs .............................................................................. Simulium dureti
– Fronto-ocular triangle not longer than wide; wing length 3.0 mm or less; Sc
without or with at most 1 or 2 hairs ........................................................ Simulium pichi
23(17) Setae and spines on R1 and setae on Rs arranged in several irregular rows (Fig. 39
I); white spots on terga IV and V faint, narrowly elongate (Fig. 39 C) ...................
.................................................................................................................... Simulium nemorale
– Setae and spines on R1 and setae on Rs arranged in single row, as in male (Fig. 42
P); white spots on terga IV and/or V generally distinct, rounded or rectangular
(Fig. 42 A) .......................................................................................................................... 24
24(23) Paired white spots on tergum V as large as those on tergum III (Fig. 44 A) .........
.............................................................................................................. Simulium albilineatum
– Paired white spots on tergum V distinctly smaller than those on tergum III (Figs.
42 A; 46 A) ........................................................................................................................ 25
25(24) Tooth of claws exceptionally small (Fig. 43 F); central area of eighth sternum lighter
than lateral portions; paraprocts pointed (Fig. 43 G) .................... Simulium yacuchuspi
– Tooth of claws normally developed (Fig. 46 A); central area of eighth sternum
darker than lateral portions; paraprocts broadly rounded ....................................... 26
26(25) Abdomen (Fig. 42 A) with 1+1 silver-white spots of tergum III only slightly
surpassing posterior half of tergum anteriorly; tergum IV with 1+1 small whitish
spots; central whitish area of tergum VI not bisected longitudinally by black stripe
....................................................................................................................... Simulium herreri
– Abdomen (Fig. 46 A) with paired whitish spots of tergum III coming close to
anterior border of tergum; tergum IV lacking spots; light-colored whitish area of
tergum VI bisected longitudinally by dark stripe ...................... Simulium quechuanum

Males
The males of Simulium (Pt.) bordai, Simulium (Pt.) schoenemanni and Simulium (Pt.) strigidorsum
are not known and have, therefore, not been included in the keys. The male of Simulium
(Pt.) horcochuspi is only known from a poorly preserved pharate specimen and also has not
been included; if better known, it might key out as Simulium (Pt.) deagostinii.

1 Basal portion of R bare (Fig. 39 I) ................................................................................. 2


– Basal portion of R setose (Fig. 51 D) ............................................ Simulium annulatum
2(1) Scape, pedicel and basal palpomeres with very long hair (Figs. 38 Q, R); scape and
pedicel combined about as long as flagellum .................................. Simulium cotopaxi
– Scape, pedicel and basal palpomeres with short hair; scape and pedicel combined
about 1/3 as long as flagellum ........................................................................................ 3
3(2) Calcipala absent (Figs. 50 P-R) ........................................................................................ 4
– Calcipala present, although in some species very small ............................................. 8
4(3) Hind tibiae conspicuously darkened at base (Figs. 37 L; 50 P-Q) .......................... 5
– Hind tibiae not conspicuously darkened at base ......................................................... 6
5(4) Abdomen with paired silvery spots on terga II-VII (Fig. 50 L); gonostylus with
apical surface smooth (similar to Fig. 42 S) ........................ Simulium prodexargenteum
144 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

– Abdomen with paired silvery spots only on segments II, VI, and VII; apical
surface of gonostylus granulose (similar to Fig. 37 M) .............. Simulium deagostinii
6(4) Hind basitarsus 5.0 times as long as wide ............................................. Simulium limay
– Hind basitarsus 3.3-3.8 times as long as wide .............................................................. 7
7(6) 1+1 silvery whitish spots not reaching anterior rim on tergites III-V; wing length 2.1-
2.5 mm; length/width ratio of hind basitarsus = 3.3-3.5 .......... Simulium nigristrigatum
– 1+1 silvery whitish spots reaching anterior rim on tergites III-V; wing length 2.8-
3.0 mm; length/width ratio of hind basitarsus = 3.0 ............. Simulium diamantinum
8(3) Apical surface of gonostylus smooth (Fig. 47 K) ....................................................... 9
– Apical surface of gonostylus tuberculate (Fig. 28 N) .............................................. 20
9(8) Base of hind tibia conspicuously darkened (Figs. 47 J; 49 L; 50 P, Q; 53 N) .... 10
– Base of hind tibia not conspicuously darkened (Fig. 40 O) ...... Simulium barbatipes
10(9) Hair and spines on R1 and Rs arranged in 1 or 2 irregular rows (as in female, Fig.
39 I); hind basitarsus narrower, about 4.0 times as long as wide .......................... 11
– Hair and spines on R1 and Rs arranged in single row (Fig. 42 P); hind basitarsus
wider, not more than 3.5 times as long as wide ........................................................ 14
11(10) Wing length > 3.5 mm .......................................................................... Simulium nemorale
– Wing length < 3.5 mm .................................................................................................... 12
12(11) Wing length 2.7-2.9 mm; hind basitarsus narrower (4.3-5.5 times as long as wide)
(Fig. 53 N) .............................................................................................. Simulium stelliferum
– Wing length 3.1-3.3 mm; hind basitarsus wider (3.5-4.0 times as long as wide)
.............................................................................................................................................. 13
13(12) Hind basitarsus 3.5 as long as wide .................................................... Simulium philippii
– Hind basitarsus 3.8-4.0 as long as wide .................................................. Simulium caprii
14(10) Abdominal terga III-V entirely black, or III with 1+1 minute light-colored spots
(Fig. 47 I) ........................................................................................................................... 15
– Abdominal tergum III or V with distinct 1+1 silvery white spots (Figs. 42 M; 43
H; 50 L) .............................................................................................................................. 18
15(14) Basal 3/5 of posterior femur whitish, sharply contrasting with dark apex (Fig. 49
L); posterior basitarsus <3.0 times as long as wide (Fig. 49 L) .............................. 16
– Basal three-fourths of hind femur light brown (Fig. 47 J); posterior basitarsus at
least 3.0 times as long as wide (Fig.47 J) ..................................................................... 17
16(15) Calcipala very small, subtriangular (Fig. 49 M) .....................................Simulium luchoi
– Calcipala somewhat larger, distinctly rounded (Fig. 48 G) .......... Simulium albicinctum
17(15) Calcipala very small (Fig. 46 C) ..................................................... Simulium quechuanum
– Calcipala larger (Fig. 47 F) .............................................................Simulium hectorvargasi
18(14) Hair of abdomen silvery; tergum III uniformly dark, but V with 1+1 distinct
white spots (Fig. 42 M); dark spot at bases of fore and mid femora short ............
....................................................................................................................... Simulium herreri
– Hair of abdomen light brass colored; tergum III with 1+1 small but distinct
silvery white spots at hind border, about half as high as height of segment; spot at
bases of fore and mid femora narrowly elongate ..................................................... 19
19(18) Posterolateral margins of abdominal terga IV and V narrowly bordered with silver
or white (Fig. 50 L) ................................................................... Simulium prodexargenteum
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 145

– Posterolateral margin of abdominal terga IV and V without light colored areas


except rarely on V (Figs. 43 H; 44 I) ........ Simulium albilineatum, Simulium yacuchuspi
20(8) Calcipala very small (similar to Fig. 30 P) ................................................................... 21
– Calcipala well developed (similar to Fig. 33 E) .......................................................... 22
21(20) Wing length 3.0 mm; hind basitarsus wide, 3.2-3.4 times as long as wide; under
certain angles of illumination, at least anterior 1/3 of scutum grayish ...................
........................................................................................................................ Simulium dureti
– Wing length 2.3-2.6 mm; hind basitarsus narrower, 3.8 times as long as wide; only
about 1/5 of anterior portion of scutum gray ..................................... Simulium pichi
22(20) Length of wing 2.8-3.0 mm; abdomen with small but distinct, paired, light-
colored spots on tergum III (Figs. 33 J, K) ........................................... Simulium simile
– Length of wing 2.6-2.7 mm; tergum III lacking any trace of white spots ..............
................................................................................................................... Simulium pulchrum

Pupae
Simulium (Pt.) albicinctum, Simulium (Pt.) schoenemanni, and Simulium (Pt.) strigidorswn are not
included in the key below because their pupae are not known.

1 Gill with 8 filaments .......................................................................................................... 2


– Gill with 6 filaments ........................................................................................................ 16
2(1) Lower or both filaments of ventral primary branch of gill sharply diverging from
remaining filaments, closely adhering to body of pupa (Figs. 42 X, U; 46 F; 49 O;
50 U) ..................................................................................................................................... 3
– All filaments of gill approximately parallel, at least on apical half ......................... 5
3(2) Many trichomes of head and thorax shortly lanceolate, generally bifid (Figs. 49
Q-S) ................................................................................................................Simulium luchoi
– Trichomes of head and thorax not as above ............................................................... 4
4(3) Trichomes of thorax elongate-lanceolate, numerous, about 250+250 (Figs. 42
X, Y) .......................................................................................... (some)......Simulium herreri
– Trichomes of thorax fewer, about 12+12, simple or bifid (Fig. 50 W) ...................
....................................................................................................... Simulium prodexargenteum
5(2) Trichomes of head and thorax very short, a few branched but most simple, from
hair-like to spine-like, those of thorax concentrated in transverse row at base of
exposed portion (Fig. 39 X) ................................................................. Simulium nemorale
– Trichomes of head and thorax much longer, their arrangement not as above .... 6
6(5) Trichomes of head and thorax very numerous, flattened, their branches ribbon-
like (Figs. 33 R-T; 34 I) ..................................................................................................... 7
– Trichomes of head and thorax not as above ............................................................... 8
7(6) Length of cocoon along base 5.2-6.8 mm; frontoclypeus (Fig. 33 R) with about
400 tubercles, with cephalic tubercles not surpassing area of facial trichomes, this
region reinforced by sclerotized edge; thoracic trichomes not numbering more
than 250+250, average 140+140 ............................................................. Simulium simile
– Length of cocoon along base 3.8-4.5 mm; frontoclypeus (Fig. 34 I) with about 800
tubercles; cephalic tubercles surpassing area of facial trichomes, this region
146 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

smooth, not reinforced at edges; thoracic trichomes numbering not more than
130+130, average 70+70 ..................................................................... Simulium pulchrum
8(6) Disc of exposed portion of thorax without tubercles, or tubercles only along area
adjacent to rim of aperture of cocoon (Figs. 39 X; 47 Q) ....................................... 9
– Disc of exposed portion of thorax with numerous tubercles (Fig. 30 V) .......... 11
9(8) Frontal trichomes reduced to minute cones (Fig. 47 O); thorax with about
150+150 trichomes (47 Q) .............................................................Simulium hectorvargasi
– Frontal trichomes normally developed, hair-like; thorax with fewer than 100+100
trichomes ............................................................................................................................ 10
10(9) Thorax with about 70+70 trichomes (Figs. 52 Q-R), these trichomes either simple
or, frequently, with 2 or 3 branches ........................................................ Simulium caprii
– Thorax with not more than 20+20 trichomes, these generally with 4 or more
branches (Fig. 28 T) ....................................................................... Simulium nigristrigatum
11(8) Trichomes of thorax lacking from anterior portion of sclerite ............................. 12
– Trichomes of thorax practically covering all of exposed surface of sclerite
(similar to figure 45 D) ................................................................................................... 13
12(11) Thorax with about 20+20 trichomes (Fig. 29 N); tubercles of frontoclypeus
sparse, their surface minutely spinose; male pupa with tubercles only at base of
frontoclypeus (Fig. 29 Q) .......................................................................... Simulium limay
– Thorax with about 60+60 trichomes (Fig. 30 V); tubercles of frontoclypeus more
numerous, their surface smooth; male pupa with frontoclypeus entirely covered
by tubercles (Fig. 30 U) ............................................................................... Simulium pichi
13(11) Cocoon with leaden sheen; surface of tubercles of frontoclypeus smooth; frontal
trichomes (Figs. 32 F, H) arranged in groups of 3 or 4, distinctly separated from
facial trichomes which are single; frontal trichomes (Fig. 32 H) single or in groups
of 2; trichomes of thorax most frequently with 1 to 3 branches (Fig. 32 I) ..........
................................................................................................................ Simulium horcochuspi
– Cocoon without leaden sheen; tubercles of frontoclypeus more or less distinctly
tuberculate; frontal trichomes (Fig. 31 X) more numerous, contiguous with facial
trichomes which number more than 1; ocular trichomes (Figs. 31 X; 37 P)
numbering more than 2; trichomes of thorax most frequently with more than 3
branches (Fig. 31 Z) ......................................................................................................... 14
14(13) Number of thoracic trichomes on each side 35-80 (Fig. 37 Q); cocoon with large
anteroventral projection thick, translucent (Fig. 37 O) ............... Simulium deagostinii
– Number of thoracic trichomes on each side 150-220; cocoon with large
anteroventral projection thick, not translucent .......................................................... 15
15(14) Number of thoracic trichomes on each side 150 (Fig. 31 Y); trichome length 0.10-
0.19 mm; cocoon maximum length at base 4.3 mm .......................... Simulium dureti
– Number of thoracic trichomes on each side 160-220; trichome length 0.24-0.35
mm; cocoon maximum length at base 5.2-6.0 mm ................. Simulium diamantinum
16(1) One or both filaments of ventral primary branch of gill sharply diverging from
remaining filaments, downward and backwardly directed (Figs. 42 U, X; 43 L, M;
46 F) .................................................................................................................................... 17
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 147

– Filaments of gill approximately parallel, or, if one or both filaments of ventral


branch somewhat diverging, then not backwardly directed (Figs. 53 Q) ............. 19
17(16) All thoracic trichomes long, hair-like (similar to Figs. 43 O, P) or elongate-
lanceolate (Figs. 42 X, Y) ............................................................................................... 18
– Thoracic trichomes either hair-like, simple or branched, and, most frequently,
spatulate (Fig. 46 H) ........................................................................ Simulium quechuanum
18(17) Both filaments of ventral primary branch of gill sharply diverging from
remaining filaments (Fig. 42 U); 1+1 areas of tubercles at base of frontoclypeus
(Fig. 42 V); trichomes of thorax elongate-lanceolate, subequal (Figs. 42 X, Y)
....................................................................................................................... Simulium herreri
– Generally only lower filament of primary ventral branch of gill diverging from
remaining filaments (Fig. 43 M); no tubercles in area mentioned (Fig. 43 N);
trichomes of thorax slender, hair-like, long and short (Figs. 43 O, P) .....................
................................................................................................................. Simulium yacuchuspi
19(16) Most thoracic trichomes roughly stellate (Figs. 53 S, T) ............. Simulium stelliferum
– Thoracic trichomes not stellate ..................................................................................... 20
20(19) Thoracic trichomes both long, hair-like, and shortly lanceolate (Figs. 40 U, V) ....
................................................................................................................. Simulium barbatipes
– All thoracic trichomes hair-like, simple or branched (Figs. 44 R; 45 D, F; 51 N-O)
.............................................................................................................................................. 21
21(20) Practically all thoracic trichomes simple (Fig. 51 N) ................... Simulium annulatum
– Many thoracic trichomes branched (Figs. 44 R; 45 D-F) ........................................ 22
22(21) Thoracic trichomes (Fig. 44 R) simple or bifid, only rarely with more than 2
branches; their number 250-300 on each side ........................... Simulium albilineatum
– Thoracic trichomes (Figs. 45 D-F) practically never simple, generally with 2 or
more branches, and their number 120-170 on each side ........................................ 23
23(22) Thoracic trichomes 5-6 on each side, very short and with 8 branches (Fig. 38 b);
tubercles of frontoclypeus and thorax rugose ................................ Simulium cotopaxi
– Thoracic trichomes 5-170 on each side, relatively elongated with 1-6 branches;
tubercles of frontoclypeus and thorax smooth ......................................................... 24
24(23) Thoracic trichomes of 60-80 on each side; trichomes mostly bifid with short
branches (Figs. 41 M, N); frontoclypeus with about 1200 tubercles ........................
.................................................................................................................... Simulium philippii
– Thoracic trichomes of 120-170 on each side; trichomes mostly with 3 to 6
branches (Figs. 45 D-F); frontoclypeus without tubercles ................ Simulium bordai

Larvae
Simulium (Pt.) albicinctum, Simulium (Pt.) bordai, Simulium (Pt.) luchoi, Simulium (Pt.) schoenemanni
and Simulium (Pt.) strigidorsum are not included in this key because their larvae are not known.

1 Postgenal bridge extremely reduced, its length about 1/5 or less of depth of
postgenal cleft (Figs. 29 V; 50 c) ..................................................................................... 2
– Postgenal bridge much larger, about 1/2 of depth of postgenal cleft or larger
(Figs. 42 e; 46 J; 51 T ) ...................................................................................................... 9
148 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

2(1) Rectal papillae with 3 simple lobes, at most with 1 pair of secondary lobules .... 3
– Rectal papillae with about 3 pairs of diverticula on each lobe ................................ 6
3(2) Rectal papillae with only 3 simple lobes (Figs. 30 b; 41 U) ...................................... 4
– Rectal papillae with 1 secondary diverticulum close to base of lateral lobes, or
with 2 pairs on middle lobe ............................................................................................. 5
4(3) Cephalic apotome very light colored, only slightly darkened at posterior margin
(Fig. 34 J) .............................................................................. (some)......Simulium pulchrum
– Cephalic apotome with overall light or medium brown pigment, also perceptibly
darkened at hind border (Fig. 50 Y) ..................................... Simulium prodexargenteum
5(3) Rectal papillae with diverticula close to base of each of lateral lobe (Fig. 28 Y);
first 2 antennal segments extremely light colored; hypostomial setae arranged in 1,
rarely 2, irregular series, with 6-8 setae on each side (Fig. 28 V) ..............................
............................................................................................................ Simulium nigristrigatum
– Rectal papillae with 2 diverticula on middle lobe; first 2 antennal segments
distinctly pigmented (Fig. 33 W); hypostomial setae arranged in 1 to 3 rows, with
about 14-18 setae on each side (Fig. 33 Y) ...................................... Simulium pulchrum
6(2) Maximum length about 6.7 mm; maximum width of cephalic capsule about
0.6 mm .......................................................................................................... Simulium limay
– Maximum length greater than 7.5 mm, maximum width of cephalic capsule more
than 0.7 mm ........................................................................................................................ 7
7(6) Maximum length 8.0 mm; maximum width of cephalic capsule 0.75 mm;
postgenal cleft approximately as wide at base as deep .............................................. 8
– Maximum length 9.0 mm; maximum width of cephalic capsule 0.80 mm;
postgenal cleft wider at base than deep ................................................. Simulium simile
8(7) Cephalic apotome light brown (Fig. 35 W); ratio of proximal, medial and distal
antennal segments = 1:1.8-1.9:1.2-1.3 ........................................ Simulium diamantinum
– Cephalic apotome dark brown (Fig. 37 T); ratio of proximal, medial and distal
antennal segments = 1:1.3-1.5:0.8-1.0 ............................................ Simulium deagostinii
9(1) Lobes of rectal papillae each with 16-20 very long and slender diverticula (Fig.
43 X) ...................................................................................................... Simulium yacuchuspi
– Lobes of rectal papillae with fewer or no diverticula .............................................. 10
10(9) Postgenal bridge about as long as postgenal cleft depth (Fig. 42 e) ..................... 11
– Postgenal bridge much shorter, about half as long as postgenal cleft depth (Figs.
46 J; 47 V; 53 a) ................................................................................................................ 12
11(10) Rectal papillae with 1+1 diverticula (Fig. 42 f); ratio of proximal, medial and distal
antennal segments = 1:1.10-1.25:1.00-1.08 .......................................... Simulium herreri
– Rectal papillae without diverticula (Fig. 41 U); ratio of proximal, medial and distal
antennal segments = 1:1.0-1.2:.0.8 ..................................................... Simulium philippii
12(10) Lobes of rectal papillae simple, without diverticula ................................................. 13
– Lobes of rectal papillae with diverticula ..................................................................... 19
13(12) Base of anal sclerite with scale-like cuticular structures, from simple to trifid
.................................................................................................................... Simulium nemorale
– Base of anal sclerite generally without cuticular structures, or only with simple
spicules ............................................................................................................................... 14
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 149

14(13) Sclerotized area between anterior and posterior arms of anal sclerite extensive
(Fig. 53 c); base of anal sclerite with simple spicules .............................................. 15
– Sclerotized area between anterior and posterior arms of anal sclerite
comparatively narrow; base of anal sclerite lacking spicules .................................. 17
15(14) Anterior half of cephalic apotome conspicuously lighter than basal portion of
sclerite (Figs. 53 V, W) ..................................................... (some)......Simulium stelliferum
– Pigmentation of cephalic apotome not with strong contrasts ............................... 16
16(15) First marginal tooth of mandible about 4 times as long as second (Fig.46 I)
.............................................................................................................. Simulium quechuanum
– First marginal tooth of mandible about 2 times as long as second (Fig. 47 U)
..............................................................................................................Simulium hectorvargasi
17(14) Pattern of cephalic apotome positive; maximum size of mature larva 7.0 mm
.......................................................................................................................... Simulium pichi
– Pattern of cephalic apotome negative (Fig. 31 a); size of mature larva 8.0 mm or
more .................................................................................................................................... 18
18(17) Distal antennal segment shorter than proximal (Fig. 31 b) .............. Simulium dureti
– Distal antennal segment about as long as proximal (Fig. 32 J) ..................................
................................................................................................................ Simulium horcochuspi
19(12) Base of anal sclerite lacking spicules or scale-like cuticular structures ................ 20
– Base of anal sclerite with scale-like cuticular structures ......................................... 22
20(19) Hypostomium with anterior border depressed and median tooth below lateral
teeth (Fig. 38 i); rectal papillae with 11-14 diverticula on each lobe (Fig. 38 n);
medial antennal segment relatively elongated (Fig. 38 f), ratio of antennal segment
lengths = 1:1.4-1.7:0.8 ........................................................................... Simulium cotopaxi
– Hypostomium with anterior border not depressed and median tooth almost as
prominent as lateral teeth; rectal papillae with 4-6 diverticula on each lobe; medial
antennal segment not overly elongated, ratio of antennal segment lengths = 1:1.2-
1.3:0.8-0.9 ........................................................................................................................... 21
21(20) Sclerotized area between anterior and posterior arms of anal sclerite compa-
ratively small (similar to Fig. 32 K) ................................................. Simulium annulatum
– Sclerotized area between anterior and posterior arms of anal sclerite extensive
............................................................................................... (some)......Simulium stelliferum
22(19) Anal ring with not more than 70 rows of hooks; distal antennal segment as long
as proximal; pattern of cephalic apotome as shown in figure 44 S .......................
.............................................................................................................. Simulium albilineatum
– Anal ring with 80 rows of hooks or more; distal antennal segment slightly but
distinctly shorter than first; pattern of cephalic apotome different ..................... 23
23(22) Spicules at base of anal sclerite not numerous ........... (some)......Simulium stelliferum
– Spicules at base of anal sclerite much more numerous (Fig. 40 b) ....................... 24
24(23) Pattern of cephalic apotome as shown in figure 40 W .............. Simulium barbatipes
– Pattern of cephalic apotome difficult to perceive, as shown in figure 52 S ...........
......................................................................................................................... Simulium caprii
150 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

CLAVES DE ESPECIES DE SIMULIUM (PTERNASPATHA)


Hembras
Simulium (Pt.) bordai no se conoce la hembra y Simulium (Pt.) schoenemanni no tiene
suficientes caracteres para incluirlo en la clave.

1 Porción basal de R desnuda (Fig. 39 I) ......................................................................... 2


– Porción basal de R con pelos (Fig. 51 D) ...................................... Simulium annulatum
2(1) Abdomen (Fig. 36 C) gris plateado opaco; tergitos II-V cada uno con una mancha
relativamente pequeña, central, negro aterciopelada y con manchas laterales
oscuras ausentes; calcipala muy pequeña (Fig. 36 G) ................ Simulium strigidorsum
– Tergitos abdominales II-V con manchas oscuras más extensas; calcipala ausente o
presente ................................................................................................................................ 3
3(2) Calcipala ausente (Figs. 32 A; 50 J) ................................................................................ 4
– Calcipala presente, aunque en algunas especies muy pequeña ................................. 9
4(3) Tergitos IV y V sin 1+1 manchas blanco plateadas de coloración enteramente
negra, aunque en algunos especimenes con angosta banda blanquecina a lo largo
del borde posterior (Fig. 50 A) ............................................... Simulium prodexargenteum
– Tergitos IV y V con 1+1 manchas blancas plateadas (Fig. 28 A) ........................... 5
5(4) Uñas sin diente subbasal (Fig. 32 B) .............................................. Simulium horcochuspi
– Uñas con diente subbasal (Fig. 28 J) .............................................................................. 6
6(5) Tergitos abdominales III-V con 1+1 manchas laterales negruzcas uniéndose
anteriormente con una mancha mediana (Figs.28 A; 29 B); largo del ala 2,3-
3,5 mm ................................................................................................................................. 7
– Tergitos abdominales III-V con 1+1 manchas laterales negruzcas pequeñas (Figs.
35 B; 37 B); largo del ala 3,3–3,7 mm ........................................................................... 8
7(6) Tergitos abdominales III-V con coloración gris claro extendiéndose al área
lateral de los tergitos (Fig. 29 B); fémur anterior y mediana castaño claro, no
conspicuamente oscurecidos apicalmente; longitud ala 2,8-3,5 mm ...................
......................................................................................................................... Simulium limay
– Tergitos abdominales III-V con coloración gris clara extendiéndose o no al área
lateral de los tergitos (Fig. 28 A); fémures anterior y medio conspicuamente
oscurecidos en el ápice; longitud del ala 2,3-2,7 mm ............. Simulium nigristrigatum
8(6) Tergito abdominal VI con una mancha mediana negra (Fig. 37 B); tergitos III-V
con 1+1 mancha negra lateral alcanzando al borde posterior; relación largo/
ancho del basitarso posterior = 7,2-7,6 .......................................... Simulium deagostinii
– Tergito abdominal VI sin una mancha mediana negra (Fig. 35 B); tergitos III-V
con 1+1 manchas negras laterales no alcanzando el borde posterior; relación
largo/ancho del basitarso posterior = 6,6 ................................. Simulium diamantinum
9(3) Tergito III sin 1+1 manchas blanquecinas, con tergitos III-V enteramente
oscuros, excepto en algunas especies con angosta línea a lo largo de bordes
laterales del margen posterior (Fig. 51 A); línea medio longitudinal clara del
escudo invisible o débilmente perceptible .................................................................. 10
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 151

– Tergito III con 1+1 mancha gris o blanquecina (Figs. 47 A; 48 A; 49 A; 50 A);


difícil de observar en algunos especímenes pequeños pero en todos los casos
perceptible; línea media longitudinal del escudo claramente visible ..................... 11
10(9) Bases de las tibias conspicuamente pigmentadas pero el ápice del basitarso
posterior faltando pigmento; tergitos III-V enteramente oscuros (Fig. 52 A);
paraproctos redondeados, salientes; cercos relativamente altos (Fig. 52 H) ...........
......................................................................................................................... Simulium caprii
– Bases de las tibias faltando evidente pigmento, con basitarso posterior bien
pigmentado en el tercio apical (Fig. 53 H); borde posterior del tergito III-V
angostamente pigmentado de gris (Fig. 53 A); paraproctos muy cortos y
truncados; cercos relativamente bajos (Fig. 53 L) ......................... Simulium stelliferum
11(9) Tergitos IV-V faltando las 1+1 manchas grises o blancas, solamente en algunos
casos con margen posterior angostamente pigmentado con blanco lateralmente
(Figs. 47 A; 50 A); tibia con nítido anillo oscuro basal o subbasal ....................... 12
– Tergitos IV y V, en algunos casos solo tergito V con 1+1 manchas claras de
variado tamaño (Figs. 33 A; 38 C; 40 A; 41 A); tibias con o sin anillo oscuro basal
o subbasal .......................................................................................................................... 16
12(11) Calcipala no desarrollada (Fig. 50 J) .............. (algunos)......Simulium prodexargenteum
– Calcipala bien desarrollada (Figs. 39 K; 48 C) ........................................................... 13
13(12) Área oscura separando las manchas blancas del tergito abdominal II muy angosta
(Fig. 48 A); tergito VI enteramente blanco plateado .................. Simulium albicinctum
– Área oscura separando las manchas blancas del tergito abdominal II ancha (Figs.
39 B; 47 A; 49 A), ensanchado anteriormente o tan ancho anterior como
posteriormente; tergito VI oscuro en el centro y lateralmente .............................. 14
14(13) Cerdas y espinas sobre R1 dispuestas en varias hileras irregulares (Fig. 39 I) ........
.................................................................................................................... Simulium nemorale
– Cerdas y espinas sobre R1 dispuestas en una hilera simple (similar a figura 42 P)
.............................................................................................................................................. 15
15(14) Calcipala muy pequeña, angostamente triangular (Fig. 49 E) ............Simulium luchoi
– Calcipala más grande, redondeada (Fig. 47 F) ...........................Simulium hectorvargasi
16(11) Frente muy ancha (Fig. 38 D); paraprocto con ángulo ventral posterior alargado
(Fig. 38 N); escudo negro, con variable ornamentación de acuerdo a la incidencia
de la luz (Figs. 38 A-B) ......................................................................... Simulium cotopaxi
– Frente no ancha (Fig. 40 B); paraprocto con el ángulo ventral posterior
redondeado (Fig. 40 L); escudo grisáceo, mostrando una banda mediana y 1+1
sublaterales negruzcas ..................................................................................................... 17
17(16) Tibia no conspicuamente oscurecida en la base (Figs. 40 F, G, H) ...................... 18
– Tibia conspicuamente oscurecida en la base .............................................................. 23
18(17) Tergitos III-V con 1+1 manchas claras muy pequeñas, en algunos casos difíciles
de percibir (Figs. 40 A; 41 A); longitud ala 2,9-3,9 mm .......................................... 19
– Tergitos III-V con 1+1 manchas claras más grandes, bien distinguibles (Fig. 33 A);
longitud ala 2,5-3,6 mm .................................................................................................. 20
152 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

19(18) Manchas claras submedianas sobre tergitos III-V cruzando desde el borde
anterior al posterior del segmento y de lados paralelos (Fig. 40 A); uñas con
pequeño diente subbasal (Fig. 40 J) ................................................ Simulium barbatipes
– Manchas claras submedianas sobre tergitos III-V no arribando al borde anterior y
con base agrandada (Fig. 41 A); uñas sin diente subbasal (Fig. 41 F) ......................
.................................................................................................................... Simulium philippii
20(18) Calcipala bien desarrollada (Fig. 33 E) ........................................................................ 21
– Calcipala pequeña (Fig. 31 I) ......................................................................................... 22
21(20) Longitud ala 3,2-3,5 mm; tibia posterior con pigmento frecuentemente solo en el
ápice ............................................................................................................... Simulium simile
– Longitud ala 2,6-2,8 mm; tibia posterior con pigmento siempre más extendido,
sobre los 2/5 del ápice del segmento (Fig. 34 D) .......................... Simulium pulchrum
22(20) Triángulo frontoocular más largo que ancho; longitud ala sobrepasando 3,0 mm;
Sc con aproximadamente 10 pelos ......................................................... Simulium dureti
– Triángulo frontoocular no más largo que ancho; longitud ala 3,0 mm o menos; Sc
sin o con 1 ó 2 pelos cuando mas ............................................................ Simulium pichi
23(17) Cerdas y espinas sobre R1 y cerdas sobre Rs dispuestas en varias hileras irregulares
(Fig. 39 I); manchas blancas sobre tergitos IV y V débiles angostamente alargadas
(Fig. 39 C) ................................................................................................ Simulium nemorale
– Cerdas y espinas sobre R1 y cerdas sobre Rs dispuestas en una hilera como en el
macho (Fig. 42 P); manchas blancas sobre tergitos IV y V notorias redondeadas o
rectangulares (Fig. 42 A) ................................................................................................. 24
24(23) Tergito V con 1+1 manchas blancas tan grandes como las del tergito III (Fig. 44
A) ......................................................................................................... Simulium albilineatum
– Tergito V con 1+1 manchas blancas notoriamente más pequeñas que aquellas del
tergito III (Figs. 42 A; 46 A) .......................................................................................... 25
25(24) Diente de las uñas excepcionalmente pequeño (Fig. 43 F); área central del octavo
esternito más clara que las porciones laterales; paraprocto relativamente aguzado
(Fig. 43 G) ............................................................................................. Simulium yacuchuspi
– Diente de las uñas normalmente desarrollado (Fig. 46 A); área central del octavo
esternito más oscura que las porciones laterales; paraprocto anchamente
redondeado ........................................................................................................................ 26
26(25) Tergito III del abdomen (Fig. 42 A) con 1+1 manchas blanco plateadas solo
sobrepasando ligeramente la mitad anterior del tergito; tergito IV con 1+1
manchas blanquecinas pequeñas; área central blanquecina del tergito VI no
dividida longitudinalmente por una banda negra ............................... Simulium herreri
– Tergito III del abdomen (Fig. 46 A) con 1+1 manchas blanquecinas, llegando muy
cerca del borde anterior del tergito; tergito IV sin manchas; área blanquecina del
tergito VI dividido longitudinalmente por una banda oscura Simulium quechuanum

Machos
Los machos de Simulium (Pt.) bordai, Simulium (Pt.)schoenemanni y Simulium (Pt.) strigidorsum no
se conocen y Simulium horcochuspi no posee caracteres suficientes para incluirlo en la clave. Si
se conociera mejor Simulium horcochuspi saldría en la clave como Simulium (Pt.) deagostinii.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 153

1 Porción basal de R sin pelos (Fig. 39 I) ........................................................................ 2


– Porción basal de R con pelos (Fig. 51 D) ...................................... Simulium annulatum
2(1) Escapo, pedicelo y palpómeros basales con pelos muy largos (Figs. 38 Q, R);
escapo y pedicelo combinados son tan largos como el flagelo .........................
.................................................................................................................... Simulium cotopaxi
– Escapo, pedicelo y palpómeros basales con pelos cortos; escapo y pedicelo
combinados son 1/3 del largo del flagelo .................................................................... 3
3(2) Calcipala ausente (Figs. 50 P-R) ...................................................................................... 4
– Calcipala presente, aunque en algunas especies muy pequeña ................................. 8
4(3) Tibia posterior conspicuamente oscurecida en la base (Figs. 37 L; 50 P-Q) ........ 5
– Tibia posterior no conspicuamente oscurecida en la base ........................................ 6
5(4) Abdomen con 1+1 manchas plateadas en tergitos II-VII (Fig. 50 L); gonostilo con
superficie apical lisa (similar a la figura 42 S) ..................... Simulium prodexargenteum
– Abdomen con 1+1 manchas plateadas solamente en tergitos II, VI y VII;
superficie distal granulosa (similar a figura 37 M) ....................... Simulium deagostinii
6(4) Basitarso posterior 5,0 veces tan largo como ancho ........................... Simulium limay
– Basitarso posterior 3,3-3,8 veces tan largo como ancho ........................................... 7
7(6) Tergitos abdominales III-V con 1+1 manchas blanco plateadas no llegando al
borde anterior; largo ala 2,1-2,5; relación largo /ancho de basitarso posterior =
3,3-3,5 ............................................................................................... Simulium nigristrigatum
– Tergitos abdominales III-V con 1+1 manchas blanco plateadas llegando al borde
anterior; largo ala 2,8-3,0 mm; relación largo/ancho del basitarso posterior = 3,0
.............................................................................................................Simulium diamantinum
8(3) Superficie apical del gonostilo liso (Fig. 47 K) ............................................................ 9
– Superficie apical del gonostilo con tubérculos (Fig. 28 N) ..................................... 20
9(8) Base de tibia posterior notoriamente oscurecida (Figs. 47 J; 49 L; 50 P-Q; 53 N)
.............................................................................................................................................. 10
– Base de tibia posterior no notoriamente oscurecida (Fig. 40 O) .........................
................................................................................................................. Simulium barbatipes
10(9) Pelos y espinas de R1 y Rs ordenados en 1 o 2 hileras irregulares (como en la
hembra, Fig. 39 I); basitarso posterior angosto, alrededor de 4,0 veces tan largo
como ancho ....................................................................................................................... 11
– Pelos y espinas sobre R1 y Rs dispuestos en 1 hilera simple (Fig. 42 P ); basitarso
posterior más ancho, no mayor de 3,5 veces más largo que ancho ...................... 14
11(10) Largo del ala > 3,5 mm ........................................................................ Simulium nemorale
– Largo del ala < 3,5 mm .................................................................................................. 12
12(11) Largo del ala 2,7-2,9 mm; basitarso posterior angosto (4,3-5,5 veces tan largo
como ancho) (Fig. 53 N) .................................................................... Simulium stelliferum
– Largo del ala 3,1-3,3 mm; basitarso posterior más ancho (3,5-4,0 veces tan largo
como ancho ) .................................................................................................................... 13
13(12) Basitarso posterior 3,5 veces tan largo como ancho ...................... Simulium philippii
– Basitarso posterior 3,8-4,0 veces tan largo como ancho .................... Simulium caprii
14(10) Tergitos abdominales III-V enteramente negro, o III con 1+1 manchas pequeñas
claras (Fig. 47 I) ................................................................................................................ 15
154 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

– Tergitos abdominales III-V con 1+1 manchas blancas plateadas (Figs. 42 M; 43 H;


50 L) .................................................................................................................................... 18
15(14) Fémur posterior con 3/5 basales blanquecinos, contrastando con el ápice oscuro
(Fig. 49 L); basitarso posterior < 3,0 veces largo como ancho (Fig. 49 L) ......... 16
– Fémur posterior con 3/4 basales castaño claro (Fig. 47 J); basitarso posterior por
lo menos 3.0 veces tan largo como ancho (Fig. 47 J) .............................................. 17
16(15) Calcipala muy pequeña, subtriangular (Fig. 49 M) ...............................Simulium luchoi
– Calcipala algo más grande, redondeada (Fig. 48 G) ................... Simulium albicinctum
17(15) Calcipala muy pequeña (Fig. 46 C) ............................................... Simulium quechuanum
– Calcipala más grande (Fig. 47 F) ..................................................Simulium hectorvargasi
18(14) Pelos del abdomen plateados; tergito III uniformemente oscuro, pero el V con
1+1 mancha blanca (Fig. 42 M); mancha oscura del fémur anterior y mediano
corto ............................................................................................................. Simulium herreri
– Pelos del abdomen bronceado claro; tergito III con 1+1 mancha blanca plateada
pequeña en borde posterior, aproximadamente la mitad de alto del segmento;
manchas en la base del fémur medio y anterior angostamente alargada ............. 19
19(18) Margen posterolateral de tergitos abdominales IV y V angostamente bordeado de
plateado o blanco (Fig. 50 L) .................................................. Simulium prodexargenteum
– Margen posterolateral de tergitos abdominales IV y V sin áreas plateadas o
blanquecinas (Figs. 43 H; 44 I), excepto raramente sobre el V .................................
........................................................................... Simulium albilineatum, Simulium yacuchuspi
20(8) Calcipala pequeña (similar figura 30 P) ....................................................................... 21
– Calcipala bien desarrollada (similar figura 33 E) ....................................................... 22
21(20) Longitud ala 3,0 mm; basitarso posterior ancho 3,2-3,4 veces tan largo como
ancho; bajo ciertos ángulos de iluminación, por lo menos el 1/3 anterior del
escudo grisáceo ........................................................................................... Simulium dureti
– Longitud del ala 2,3-2,6 mm; basitarso posterior más angosto, 3,8 veces tan largo
como ancho; solo alrededor de 1/5 de la porción anterior gris ........ Simulium pichi
22(20) Largo del ala 2,8-3,0 mm; abdomen con 1+1 manchas claras pequeñas pero
diferenciables sobre tergito III (Fig. 33 J K) ......................................... Simulium simile
– Largo del ala 2,6-2,7 mm; tergito III sin ninguna mancha blanquecina ..................
................................................................................................................... Simulium pulchrum

Pupas
En Simulium (Pt.) albicinctum, Simulium (Pt.) schoenemanni y Simulium (Pt.) strigidorsum no se
conoce la pupa.

1 Branquias con 8 filamentos .............................................................................................. 2


– Branquias con 6 filamentos ............................................................................................ 16
2(1) El más bajo o los 2 filamentos de la rama ventral primaria de la branquia
divergiendo de los restantes filamentos, disponiéndose muy cerca al cuerpo de la
pupa (Figs. 42 X, U; 46 F; 49 O; 50 U) ........................................................................ 3
– Todos los filamentos de la rama ventral primaria dispuestos aproximadamente
paralelos, por lo menos en la mitad apical .................................................................... 5
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 155

3(2) Mayoría de tricomas de cabeza y tórax ligeramente lanceolados, generalmente


bífidos (Figs. 49 Q-S) .................................................................................Simulium luchoi
– Tricomas de cabeza y tórax no como arriba ................................................................ 4
4(3) Tricomas de la cabeza lanceolados alargados, en número de alrededor de 250+250
(Figs. 42 X, Y) ..................................................................... (algunos)......Simulium herreri
– Tricomas del tórax simples o bífidos (Fig. 50 W), en número de alrededor de
12+12 ........................................................................................... Simulium prodexargenteum
5(2) Tricomas de la cabeza y tórax muy cortos, algunos ramificados pero la mayoría
simples, desde forma de pelo hasta de espina, presentando los del tórax
concentrados en una hilera transversa en la base de la porción expuesta (Fig. 39 X)
.................................................................................................................... Simulium nemorale
– Tricomas de la cabeza y tórax más largos con una disposición diferente a la de
arriba ..................................................................................................................................... 6
6(5) Tricomas de la cabeza y tórax numerosos, achatados, con ramas acintadas (Figs. 33
R-T; 34 I) ............................................................................................................................. 7
– Tricomas de la cabeza y tórax no como arriba ............................................................ 8
7(6) Largo del capullo en la base 5,2-6,8 mm; frontoclípeo (Fig. 33 R) con alrededor
de 400 tubérculos, con los tubérculos cefálicos no sobrepasando el área de los
tricomas faciales, presentando esta región reforzada por un borde esclerotizado;
tricomas torácicos no sobrepasando 250+250, con un promedio de 140+140
......................................................................................................................... Simulium simile
– Largo del capullo en la base 3,8-4,5 mm; frontoclípeo (Fig. 34 I) con alrededor de
800 tubérculos; tubérculos cefálicos sobrepasando el área de tricomas faciales,
presentando esta región lisa, no reforzada por bordes esclerotizados; tricomas
torácicos no sobrepasando 130+130, con un promedio de 70+70 ..........................
................................................................................................................... Simulium pulchrum
8(6) Disco de porción expuesta del tórax sin tubérculos o con tubérculos solo a lo largo
del área adyacente al borde de la abertura del capullo (Figs. 39 X; 47 Q) ............... 9
– Disco de porción expuesta del tórax con numerosos tubérculos (Fig. 30 V) ........ 11
9(8) Tricomas frontales reducidos a pequeños conos (Fig. 47 O); tórax con alrededor
de 150+150 tricomas (Fig. 47 Q) .................................................Simulium hectorvargasi
– Tricomas frontales normalmente desarrollados, en forma de pelo; tórax con
alrededor de 100+100 tricomas .................................................................................... 10
10(9) Tórax con alrededor de 70+70 tricomas (Figs. 52 Q-R), estos tricomas desde
simples a frecuentemente con 2 o 3 ramas ........................................... Simulium caprii
– Tórax no con más de 20+20 tricomas, estos generalmente con 4 o más ramas (Fig.
28 T) .................................................................................................. Simulium nigristrigatum
11(8) Tricomas del tórax faltando en porción anterior del esclerito ............................... 12
– Tricomas del tórax cubriendo prácticamente toda la superficie de la porción
expuesta del tórax (similar a figura 45 D) .................................................................. 13
12(11) Tórax con alrededor de 20+20 tricomas (Fig. 29 N); tubérculos del frontoclípeo
esparcidos, su superficie espinosa, tubérculos del frontoclípeo del macho
solamente presentes en la base (Fig. 29 Q) ........................................... Simulium limay
156 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

– Tórax con alrededor de 60+60 tricomas (Fig. 30 V); tubérculos del frontoclípeo
más numerosos, su superficie lisa; tubérculos del clípeo del macho cubriéndolo en
su totalidad (Fig. 30 U) ............................................................................... Simulium pichi
13(11) Capullo con brillo plomizo; superficie de los tubérculos lisa; tubérculos frontales
(Figs. 32 F, H) dispuestos en grupos de 3 o 4, bien separados de los tricomas
faciales los cuales son simples; tricomas frontales (Fig. 32 H) simples o en grupos
de 2; tricomas del tórax frecuentemente con 1 a 3 ramas (Fig. 32 I) .......................
................................................................................................................ Simulium horcochuspi
– Capullo sin brillo plomizo; tubérculos del frontoclípeo con superficie más o
menos tuberculada; tricomas frontales (Fig. 31 X) más numerosos, contiguos, con
los tricomas faciales en número mayor de 1; tricomas oculares (Figs. 31 X; 37 P)
en número superior a 2; tricomas del tórax muy frecuentemente con más de 3
ramas (Fig. 31 Z) .............................................................................................................. 14
14(13) Con 35-80 tricomas torácicos de cada lado (Fig. 37 Q); prolongación antero-
ventral del capullo gruesa y translúcida (Fig. 37 O) .................... Simulium deagostinii
– Con 150-220 tricomas torácicos de cada lado; prolongación antero-ventral del
capullo gruesa, no translúcida ....................................................................................... 15
15(14) Con alrededor de 150 tricomas torácicos de cada lado (Fig. 31 Y); longitud de tricomas
0,10-0,19 mm; largo máximo del capullo en la base 4,3 mm ................ Simulium dureti
– Con 180-220 tricomas torácicos de cada lado; longitud de tricomas 0,24-0,35 mm;
largo máximo del capullo en la base 5,2-6,0 mm ..................... Simulium diamantinum
16(1) Uno o ambos filamentos de la rama primaria ventral de la branquias divergiendo
fuertemente de los restantes filamentos, dirigidos hacia abajo y atrás (Figs. 42 U,
X, 43 L, M; 46 F) ............................................................................................................. 17
– Filamentos de branquias aproximadamente paralelos, o si 1o ambos filamentos de
la rama primaria ventral son algo divergentes, no están dirigidos hacia atrás (Fig.
53 Q) ................................................................................................................................... 19
17(16) Todos los tricomas torácicos largos, en forma de pelo (similar Fig. 43 O-P) ó
alargado y lanceolado (Figs. 42 X-Y) ........................................................................... 18
– Tricomas torácicos en forma de pelo, simple o ramificado o más frecuentemente
espatulados (Fig. 46 H) ................................................................... Simulium quechuanum
18(17) Ambos filamentos de la rama primaria ventral de la branquias fuertemente
divergentes de los restantes filamentos (Fig. 42 U); presencia de 1+1 áreas de
tubérculos en la base del frontoclípeo (Fig. 42 V); tricomas del tórax alargados,
lanceolados, subiguales (Figs. 42 X, Y) ................................................. Simulium herreri
– Generalmente solo el filamento más bajo de la rama primaria ventral divergiendo
de los restantes filamentos (Fig. 43 M); sin tubérculos en el área mencionada (Fig.
43 N); tricomas del tórax alargados, en forma de pelo, largos y cortos (Figs. 43 O, P)
................................................................................................................. Simulium yacuchuspi
19(16) Mayoría de tricomas torácicos de forma aproximadamente estrellada (Figs. 53 S, T)
.................................................................................................................. Simulium stelliferum
– Tricomas cefálicos no de forma estrellada ................................................................. 20
20(19) Triocomas torácicos desde alargados en forma de pelo o más cortos lanceolados
(Figs. 40 U, V) ...................................................................................... Simulium barbatipes
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 157

– Tricomas torácicos en forma de pelo simple o ramificado (Figs. 44 R; 45 D, F; 51


N-O) ................................................................................................................................... 21
21(20) Prácticamente todos los tricomas torácicos simples (Fig. 51 N) ...............................
.................................................................................................................Simulium annulatum
– Muchos tricomas torácicos ramificados (Figs. 44 R; 45 D-F) ................................ 22
22(21) Tricomas torácicos (Fig. 44 R) simples o bífidos, raramente con más de 2 ramas,
en número de 250-300 de cada lado ............................................ Simulium albilineatum
– Tricomas torácicos (Figs. 45 D-F) prácticamente nunca simples, generalmente con
2 o más ramas y en número de 120-170 de cada lado ............................................. 23
23(22) Tricomas torácicos en número de 5-6 de cada lado, muy cortos y con hasta 8
ramas (Fig. 38 b); tubérculos de frontoclípeo y tórax rugoso ...... Simulium cotopaxi
– Tricomas torácicos de 5-170 de cada lado, relativamente largos con 1-6 ramas;
tubérculos del frontoclípeo y tórax lisos ..................................................................... 24
24(23) Tricomas torácicos en número de 60-80 de cada lado; tricomas mayormente
bífidos, con ramas cortas (Figs. 41 M, N); frontoclípeo con alrededor de 1200
tubérculos ................................................................................................ Simulium philippii
– Tricomas torácicos en número de 120-170 de cada lado; tricomas mayormente
con 3 a 6 ramas (Figs. 45 D-F); frontoclípeo sin tubérculos ............ Simulium bordai

Larvas
En Simulium (Pt.) albicinctum, Simulium (Pt.) bordai, Simulium (Pt.) luchoi, Simulium (Pt.)
schoenemani y Simulium (Pt.) strigidorsum sus larvas son desconocidas.

1 Puente postgenal extremadamente reducido, su longitud aproximadamente 1/5 o


menos de la profundidad de la hendidura postgenal (Figs. 29 V; 50 c) ................. 2
– Puente postgenal más largo, aproximadamente la mitad de la profundidad de la
hendidura postgenal (Figs. 42 e; 46 J; 51 T) ................................................................. 9
2(1) Papilas rectales con 3 lóbulos simples, cuando mucho con 1 par de divertículos
................................................................................................................................................ 3
– Papilas rectales con 3 pares de divertículos en cada lóbulo ...................................... 6
3(2) Papilas rectales con solo 3 lóbulos simples (Figs. 30 b; 41 U) ................................. 4
– Papilas rectales con 1 divertículo próximo a la base de los lóbulos laterales o con
2 pares sobre el lóbulo mediano ..................................................................................... 5
4(3) Apotoma cefálico con coloración clara, solo ligeramente más oscuro sobre el
margen posterior (Fig. 34 J) ......................................... (algunos)......Simulium pulchrum
– Apotoma cefálico todo claro o castaño en el medio y también oscurecido en el
borde posterior (Fig., 50 Y) .................................................... Simulium prodexargenteum
5(3) Papilas rectales con un divertículo próximo a la base de los lóbulos laterales
(Fig. 28 Y); primeros 2 artejos antenales extremadamente claros; cerdas del
hipostomio ordenadas en 1, raramente 2 series irregulares, con 6-8 cerdas de
cada lado (Fig. 28 V) .................................................................... Simulium nigristrigatum
– Papilas rectales con 2 divertículos sobre el lóbulo mediano; primeros 2 artejos antenales
notoriamente pigmentados (Fig. 33 W); cerdas del hipostomio dispuestas desde 1 a 3
hileras, con alrededor de 14-18 cerdas de cada lado (Fig. 33 Y) ......... Simulium pulchrum
158 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

6(2) Longitud máxima alrededor de 6,7 mm; ancho máximo de la cápsula cefálica
alrededor de 0,6 mm .................................................................................. Simulium limay
– Longitud máxima superior a 7,5 mm; ancho máximo de la cápsula cefálica mayor
de 0,7 mm ............................................................................................................................ 7
7(6) Longitud máxima 8,0 mm; ancho máximo de la cápsula cefálica 0,75 mm;
hendidura postgenal aproximadamente tan profunda como ancha en la base ..... 8
– Longitud máxima 9,0 mm; ancho máximo de la cápsula cefálica 0,80 mm;
hendidura postgenal menos profunda que ancha en la base ............. Simulium simile
8(7) Apotoma cefálico castaño claro (Fig. 35 W); relación de los artejos antenales
proximal, mediano y distal = 1:1,8-1,9:1,2-1,3 .......................... Simulium diamantinum
– Apotoma cefálico castaño oscuro (Fig. 37 T); relación de los artejos antenales
proximal, mediano y distal = 1:1,3-1,5:0,8-1.0 ............................... Simulium deagostini
9(1) Lóbulos de las papilas rectales cada uno con 16-20 divertículos largos y delgados
(Fig. 43 X) ............................................................................................. Simulium yacuchuspi
– Lóbulos de las papilas rectales con pocos o sin divertículos .................................. 10
10(9) Puente postgenal aproximadamente tan largo como el de la hendidura postgenal
(Fig. 42 e) ........................................................................................................................... 11
– Puente postgenal más corto, aproximadamente la mitad del largo de la hendidura
postgenal (Figs. 46 J; 47 V; 53a ) ................................................................................... 12
11(10) Papilas rectales con 1+1 divertículo sobre cada lóbulo (Fig. 42 f); relación de los
artejos antenales proximal, mediano y distal = 1:1,10-1,25:1.00-1,08 ......................
....................................................................................................................... Simulium herreri
– Papilas rectales sin divertículos (Fig. 41 U); relación de los artejos antenales
proximal, mediano y distal = 1:1,0-1,2:0,8........................................ Simulium philippii
12(10) Lóbulos de papilas rectales simples, sin divertículos ................................................ 13
– Lóbulos de papilas rectales con divertículos .............................................................. 19
13(12) Base del esclerito anal con estructuras cuticulares como escamas desde simples a
trífidas ....................................................................................................... Simulium nemorale
– Base del esclerito anal generalmente sin estructuras cuticulares como escamas o
solamente como espículas simples ............................................................................... 14
14(13) Área esclerotizada entre ramas anteriores y posteriores del esclerito anal extensa
(Fig. 53 c); base del esclerito anal con espículas simples ......................................... 15
– Área esclerotizada entre ramas anteriores y posteriores del esclerito anal
comparativamente angosta; base del esclerito anal sin espículas ........................... 17
15(14) Mitad anterior del apotoma cefálico conspicuamente más claro que la porción
basal del esclerito (Figs. 53 V, W) .............................. (algunos)......Simulium stelliferum
– Pigmentación del apotoma cefálico sin fuertes contrastes ..................................... 16
16(15) Primer diente marginal de la mandíbula aproximadamente 4 veces más largo que
el segundo (Fig. 46 I) ....................................................................... Simulium quechuanum
– Primer diente marginal aproximadamente 2 veces más largo que el segundo (Fig.
47 U) ...................................................................................................Simulium hectorvargasi
17(14) Manchas del apotoma cefálico positivas; tamaño máximo de la larva 7,0 mm
.......................................................................................................................... Simulium pichi
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 159

– Manchas del apotoma cefálico negativas (Fig. 31 a); tamaño de la larva madura 8,0
mm o más .......................................................................................................................... 18
18(17) Artejo distal de antena más corto que el proximal (Fig. 31 b) ......... Simulium dureti
– Artejo distal de antena aproximadamente tan largo como el proximal (Fig. 32 J)
................................................................................................................ Simulium horcochuspi
19(12) Base del esclerito anal sin espículas o estructuras cuticulares como espinas ...... 20
– Base del esclerito con estructuras cuticulares como escamas ................................ 22
20(19) Hipostomio con borde anterior deprimido y con diente mediano por debajo de
los dientes laterales (Fig. 38 i); papilas rectales con 11-14 divertículos en cada
lóbulo (Fig. 38 n); artejo antenal mediano relativamente alargado (Fig. 38 f);
relación del largo de los artejos antenales proximal, mediano y distal = 1:1, 1.4-
1,7:0,8 ........................................................................................................ Simulium cotopaxi
– Hipostomio con el borde anterior no deprimido y diente mediano casi tan
prominentes como los dientes laterales; papilas rectales con 4-6 divertículos sobre
cada lóbulo; artejo mediano de la antena no muy alargado, relación de los artejos
antenales = 1:1,2-1,3:0,8-0,9 .......................................................................................... 21
21(20) Área esclerotizada entre las ramas anterior y posterior del esclerito anal
comparativamente pequeña (similar a figura 32 K) ..................... Simulium annulatum
– Área esclerotizada entre las ramas anterior y posterior del esclerito anal
comparativamente grande ........................................... (algunos)......Simulium stelliferum
22(19) Anillo anal con no más de 70 hileras de ganchos; artejo distal de la antena tan
largo como el proximal; ornamentación del apotoma cefálico como se muestra en
figura 44 S .......................................................................................... Simulium albilineatum
– Anillo anal con 80 hileras de ganchos o más; artejo distal de la antena ligeramente
más corto que el proximal; ornamentación del apotoma cefálico diferente ....... 23
23(22) Espículas en la base del esclerito anal no numerosos ..................................................
........................................................................................... (algunos)......Simulium stelliferum
– Espículas en la base del esclerito anal numerosos (Fig. 40 b) ................................ 24
24(23) Ornamentación del apotoma cefálico como se muestra en figura 40 W .................
................................................................................................................. Simulium barbatipes
– Ornamentación del apotoma cefálico poco intensa, como en figura 52 S ..............
......................................................................................................................... Simulium caprii

NIGRISTRIGATUM species group

Simulium (Pternaspatha) nigristrigatum (ENDERLEIN) (Fig. 28.)


Pternaspatha nigristrigata ENDERLEIN, 1930: 88;1934b: 275; STONE, 1962: 208.
Simulium nigristrigatum SMART, 1945: 509; VARGAS, 1945a: 170; STUARDO ORTIZ, 1946: 41; WYGODZIN-
SKY, 1949: 304; VARGAS & DIAZ, 1953: 142; WYGODZINSKY, 1958: 132.
Simulium (Simulium) nigristrigatum EDWARDS, 1931: 154; ORFILA, 1939: 1533.
[Simulium] (Pternaspatha) nigristrigata STONE, 1963: 17.
Simulium (Pternaspatha) nigristrigatum WYGODZINSKY & COSCARÓN, 1967: 69-70; COSCARÓN, 1991: 113.
Acropogon edwardsi ENDERLEIN, 1934b: 279; VARGAS, 1945: 134.
160 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Simulium edwardsi STUARDO ORTIZ, 1946: 41.


Simulium (Pternaspatha) bachmanni WYGODZINSKY & COSCARÓN, 1967: 58; COSCARÓN & WYGODZINSKY,
1972b: 209; COSCARÓN, 1991: 110; COSCARÓN & COSCARÓN ARIAS, 1996a: 703.
Female: Wing length 2.5-2.6 mm; head black; antennae, palpi, and labrum dark gray;
occiput, frons, and clypeus dark gray pollinose; pilosity of antennae very short, that of
palpi, clypeus, frons, and occiput somewhat longer, silver-gray; first and second antennal
segments much lighter than remaining segments, brownish; scutum dark gray, central dark
band divided by distinct, longitudinal, gray line; hair of thorax silver-gray; appressed setae
of scutum very dense, somewhat obliterating pattern; scutellum, metanotum, pleura, and
sterna dark gray; wings hyaline, veins light gray, hair and spines blackish, tufts at base of
wing silver-gray; stem of halteres tinged with brown; knob white; legs light yellow; hair
silvery gray to black; following regions darkened: all coxae, those of fore legs as intensely
as others; trochanters; femora, their apices very strongly so on all pairs; apical three-fifths
of mid and hind tibiae, fore tibiae almost entirely; entire fore tarsi; apical third of basal,
and apical half of second, segment of mid and hind tarsi, and their entire third to fifth
segments; exact proportions and relative intensity of pigmented areas as shown in figure
F-H; abdomen (Fig. A) matte silver-gray; first tergite darkened; tergites II-V each with one
large central and 1+1 lateral black spots connected by black line along anterior border of
tergites; these spots narrowed posteriorly; distance between central and lateral spots about
as large as width of central spot; tergite VI only very slightly darkened at middle
anteriorly; tergite VII with one very large, black, central spot; tergites VIII and IX gray,
VIII faintly darkened anteriorly at center; hair of abdomen from black to silver-gray;
antenna as in figure B; distal palpomere as long as two anterior united (Fig. C); sensory
vesicle of palp as in figure D; fronto-ocular triangle wider than high (Fig. E); wing: Sc with
two or three hairs on its basal and three or four hairs on its central portion; basal portion
of R glabrous; R1 with hair and setae arranged in one row, but occasional setae or spines
out of line; Rs with one somewhat irregular row of setae, in some specimens a few out of
line; shape and proportion of segments of legs as shown in figure F-H; calcipala not
developed (Fig. I); claws with well-developed tooth (Fig. J); eighth abdominal sternite as
shown in figure L; rather faintly pigmented, its central portion lighter than lateral areas;
posterior border of central portion slightly sclerotized; paraprocts short, their free
extremity truncate; genital fork as shown in figure K; median process distinctly
sclerotized, including its terminal knob; spermatheca oval, its inner surface with minute
spiculae, either single or arranged in groups of two.
Male: Length of wing, 2.5 mm; head black; eyes deep purple; antennae and palpi
piceous; clypeus gray pollinose; antennae with short pubescence only; setae of basal
segments not distinctly longer than remainder; occiput, clypeus, and basal segment of
maxillary palp with some long hair; isolated and rather short setae between eyes; scutum
with usual color pattern; basic color from dark matte gray to velvety black, depending on
the angle of view; appressed setae very light brass-colored; scutellum blackish, silvery gray
pollinose; metanotum blackish; pleura blackish, silvery gray pollinose; pleural tuft silvery
white; color of wings and halteres as in female; legs pale yellow to whitish, their hair from
silvery white to black; following regions dark: all coxae and trochanters; base and apex of
fore and mid femora, and apex of hind femur strongly pigmented, rest of femora slightly
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 161

B
D

J F C
A
G

L
E I
H K

N
O
P

R
T S

V U
W
X

Y
Figure 28. Simulium (Pternaspatha) nigristrigatum. A-L: Female; A: abdomen dorsal view in different
specimens; B: antenna; C: maxillary palp; D: sensory vesicle of maxillary palp; E: fronto-ocular
triangle; F-H: fore, mid and hind leg, respectively; I: distal portion of hind basitarsus and
pedisulcus; J: claw; K: genital fork; L: portion of eighth sternite and gonapophyses. M-P: Male; M:
hind basitarsus; N: gonostylus; O: ventral plate; P: endoparameres and median sclerite. Q-T: Pupa;
Q: general aspect; R: gill; S: frontoclypeus, with trichomes with high magnification at right; T:
portion of thorax with trichomes. U-Y : Larva; U: teeth of mandible; V: hypostomium; W:
hypostomium and postgenal cleft; X: anal sclerite; Y: rectal papillae.
162 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

pigmented; fore and mid tibiae with apical fourth intensely pigmented, and with dark
stripe running toward base, becoming evanescent in basal third; hind tibia intensely
pigmented on apical third, faintly more basad, very faintly near base; entire fore tarsus,
mid tarsus on apical half of basitarsus, and entire second through fifth segments; hind
tarsus on apical third of basitarsus, apical half of second, and entire third through fifth
segments; abdomen velvety black, its hair silvery white, including that of basal fringe;
tergites II, VI, and VII with 1+1 silvery gray spots, those of segments II and VI very large,
separated dorsally only by small black area on II, virtually confluent on VI; those of
segment VII much smaller; last segment of maxillary palp more than twice as long as
penultimate; diameter of sensory vesicle smaller than half diameter of third segment;
chaetotaxy of wings like that of female; femur of forelegs with large number of long
hairs; hind basitarsus 3.3-3.4 times as long as wide; calcipala not developed (Fig. M);
gonocoxite wider than long; gonostylus subquadrate, distal surface heavily granulose (Fig.
N); central portion of ventral plate somewhat salient; lateral projection and basal margin
distinctly sclerotized, disc uniformly pigmented (Fig. O); bifid portion of median sclerite
narrow and endoparameres with hard teeth (Fig. P).
Pupa: Cocoon (Fig. Q) slipper shaped, its anteroventral bridge very short; color
stramineous, surface smooth, closely woven, threads not distinct under medium magnifi-
cation (X 60); rim of aperture not perceptibly reinforced; length of cocoon at dorsum
along middle, 2.3-2.7 mm.; maximum length along base, 3.8 mm; length of pupal body
2.8-3.0 mm; length of gills, approximately 1.5 mm, viz., less than half as long as cocoon,
and about half as long as pupa; gills each with eight filaments, forming a compact group
(Fig. R); head and thorax of pupa light brown; head with 2+2 frontal and 1+1 facial
trichomes (Fig. S); trichomes simple or with two or three branches; thorax with exposed
portion without tubercles and not more strongly sclerotized than rest; region adjacent to
aperture of cocoon with tubercles only near dorsal midline; disc of thorax (Fig. T) with
about 10+10 long, simple or branched, hair like trichomes, latter with not more than four
branches; length of trichomes, 0.10-0.16 mm.
Larva: Maximum length, 5.2 mm; maximum width of head, 0.5 mm; general color
whitish, with transverse bands of hypodermal pigment on first three or four abdominal
segments dorsally; entire posterior portion of body pigmented; ventral surface pigmented
only on anterior portion of body; head ivory-colored; cephalic apotome without
perceptible pattern, only slightly darkened at posterior margin; body integument glabrous;
antennae very pale, only distal segment slightly pigmented; ratio of proximal, medial and
distal segments = 1:1.3:0.9; cephalic fan with 30-35 rays; toothing of mandibles as shown
in figure U; median tooth of hypostomium about as prominent as lateral ones; serrations
of lateral border of hypostomium exceptionally prominent (Fig. V); hypostomial setae
arranged in one, rarely in two, irregular series, numbering about six in each group, in
addition to two to four short, posterior setae; disc of hypostomium with one or two short,
single, bifid hairs; postgenal cleft narrow and exceptionally deep, its apex almost attaining
hypostomial groove (Fig. W); anal sclerite as shown in figure X, its base with numerous
minute scales with from one to four rays; anal ring with approximately 80 rows of 15-17
hooks each; rectal papillae consisting of three primary lobes; two lateral ones close to their
base, each with one secondary lobule (Fig. Y).
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 163

Distribution: Argentina: Buenos Aires, Chubut, Neuquen, Rio Negro.


Bionomics: Simulium nigristrigatum breeds in Patagonian rivers with fast flow over
trees branches and leaves, especially of Salix humboldtiana, Cortaderia sp., or over stones.
Females are slightly anthropophilic.
Discussion: COSCARÓN and COSCARÓN ARIAS (1996: 709) established that Simulium
bachmanni WYGODZINSKY & COSCARÓN is a synonym of Simulium nigristrigatum, based on
variation of the female abdominal coloration of Simulium bachmanni. Simulium bachmanni is very
well known in all stages, where the pupa provides the best characters to differentiate this
species. In the topotypical area of Simulium nigristrigatum (known only from adults), pupae and
larvae were found on several occasions and breeding males and females of Simulium bachmanni
were also obtained. These are similar to the Simulium nigristrigatum female syntype and male
lectotype from Zoologisches Museum of Humboldt University, Berlin. Simulium nigristrigatum is
closely related to Simulium limay. The male of Simulium nigristrigatum differs from that of
Simulium limay by its much wider basitarsus (at least four times as long as wide). The female of
Simulium nigristrigatum with the apically truncate paraprocts are similar in shape to those of
Simulium limay, but in the latter species tergites VIII and IX of the female are almost completely
black, and neither the fore nor the mid femur is totally darkened. The pupa of Simulium
nigristrigatum differs from that of Simulium limay by the absence of granulosity from the thoracic
disc, by the smaller number of thoracic trichomes and the very long ventral primary branch of
the gill. The very elongated postgenal cleft of the larva is remarkable in Simulium nigristrigatum.

Simulium (Pternaspatha) limay WYGODZINSKY (Fig. 29.)


Simulium limay WYGODZINSKY, 1958: 123; VULCANO, 1967: 15.
Simulium (Pternaspatha) limay STONE, 1963: 16; WYGODZINSKY & COSCARÓN, 1967: 69-70; COSCARÓN
& WYGODZINSKY, 1972b: 212; COSCARÓN, 1987: 18; 1991: 111.
Simulium (Pternaspatha) huemul WYGODZINSKY & COSCARÓN, 1967: 67; COSCARÓN, 1991: 111.
Simulium walterwittmeri WYGODZINSKY, 1958: 129.
Female: Wing length 2.8-3.5 mm; scutum gray pollinose with median and 1+1
sublateral bands blackish, covered by abundant gray-yellowish to silvery pilosity (Fig. A);
abdomen gray pollinose with grayish dark brown spot medially on tergite I; another
similarly median and 1+1 sublateral that don’t reach posterior border on tergites II-V,
small median anterior spot on tergite VI and another more expanded on tergites VII-IX
(Fig. B); legs grayish brown-yellow with apex of coxa, trochanter, apex of femur, tibia and
tarsi more darkened; calcipala very small (Fig. C); claws with subbasal tooth evident (Fig.
D); genitalia as shown in figures E-F; gonapophyses with abundant microtrichia and
paraprocts with soft curvature on distal angles.
Male: Wing length 2.5-3.0 mm; scutum black with 1+1 anterior, small subtriangular
spots light gray continued laterally and posteriorly by thin grayish band bordering the
scutum (Fig. G) and covered by grayish-tin pilosity; abdomen black with 1+1 silvery-gray
spots on tergites II and VI, and very reduced on tergite III, and lateral borders of tergites
VII; tergite IX grayish brown (Fig. H); hind leg with tibia and basitarsus scarcely stretched
(Figs. I, J); length/width ratio of hind basitarsus = 5.2-5.6; genitalia as shown in figure K;
gonostylus with tubercles on distal border (Fig. L).
164 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

J
D

B G
A
C H
E
O

I
K
M

R
T

Q V
S U
Figure 29. Simulium (Pternaspatha) limay. A-F: Female; A: scutum; B: abdomen, dorsal view; C:
portion distal of hind basitarsus with small calcipala and pedisulcus; D: claw E: eighth sternite,
gonapophyses, genital fork and spermatheca; F: paraproct and cercus. G-L: Male; G: scutum; H:
abdomen dorsal view; I: hind leg; J: hind basitarsus; K: ventral view of gonocoxite, gonostylus,
ventral plate, endoparameres and median sclerite; L: gonocoxite, gonostylus dorsal view. M-Q:
Pupa; M: general view; N: portion of thorax with trichomes and gill; O: gill; P: portion of thorax
with trichomes of different specimen than O; Q: frontoclypeus. R-V: Larva; R: cephalic apotome;
S: antenna; T-U: mandible teeth of different specimens; V: hypostomium and postgenal cleft.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 165

Pupa: Cocoon length dorsally 3.0-3.8 mm, on base 3.3-4.2 mm, maximum 4.8-5.5 mm;
cocoon light brown, with shoe-shaped strong basal projection, protecting gill base (Fig. M);
body pupa length 4.0-4.5, gill length 1.5-2.0 mm; gill with 8 branches disposed as shown in
figures N and O; frontoclypeus with 3 pairs of frontal trichomes, 1-2 pairs facials
multibranched; thorax with 20-55 trichomes specially positioned on free posterior border; with
4-8 thin branches emerging close to base (Figs. N, P); frontoclypeus with tubercles on entire
frons in female and reduced to basal area on male (Fig. Q); tubercles with smooth surface.
Larva: Length of mature larva, 6.7-7.5 mm; maximum width of head capsule, 0.7
mm; general body shape much as in Simulium annulatum; color of larva dark brownish
green, except unpigmented undersurface of posterior half of body; head dark brown,
pigment pattern of cephalic apotome as shown in figure R; body integument with isolated
short hair, more numerous in area of anal sclerite; antennae (Fig. S) with third segment
darkest, second segment with one or two small unpigmented areas; ratio of length of
segments proximal, medial and distal = 1:1.1:1.5:0.9; mouth brushes with 39-45 rays;
toothing of mandibles as shown in figures T, U; anterior border of hypostomium strongly
pigmented; hypostomial setae arranged in one or two irregular rows, with 14-16 setae in
each group; postgenal cleft (Fig. V) very deep, five times as deep as length of hypostomial
bridge; anal sclerite with some hair and without scales; anal ring with approximately 95
rows, each composed of 17 hooks; rectal papillae with three primary lobes, each with
about six diverticula; median primary lobe simple in some specimens.
Distribution: Argentina: Chubut, Mendoza, Neuquén, Río Negro, San Juan, Santa
Cruz on precordilleran area.
Bionomics: Simulium limay breeds in fast–flowing, clear rivers or creeks of large
volume, with tree branches and leaves. Females could be collected several kilometers from
breeding places on the Patagonian steppes.
Discussion: The closest species to Simulium limay is S. nigristrigatum with adult abdomen
ornamentation very similar, showing in the latter some variations. It can be separated from
Simulium nigristrigatum by the absence in the pupa of tubercles on the frontoclypeus and
dorsum of the thorax, scarce trichomes on the thorax (about 10), wider hind basitarsus in
the male (length/width ratio = 3.2-3.4) and in larvae the rectal papillae with one lobule on
the lateral lobes. Simulium walterwittmeri shows great similarity with S. nigristrigatum and S. limay
(COSCARÓN 1991: 113) and lives in the same river. Based on pupal characters that are closest
to Simulium limay, we consider S. walterwittmeri a synonym of S. limay.

Simulium (Pternaspatha) pichi WYGODZINSKY and COSCARÓN (Fig. 30.)


Simulium (Pternaspatha) pichi WYGODZINSKY & COSCARÓN, 1967: 70-74; COSCARÓN & WYGODZINSKY,
1972b: 213; COSCARÓN, 1991: 114-115.
Female: Wing length 2.4-2.9 mm; head blackish; color of appendages and hair like
that of Simulium nemorale; scutum dark gray, with three longitudinal, light gray lines: a
central very delicate one not attaining anterior border, and 1+1 wider submedian ones
that meet central line posteriorly, posterior fourth of scutum light gray; lateral borders of
scutum also light gray (Fig. A); scutum covered with very numerous appressed, silvery
hair; scutellum light brown, with long silvery or dark brown hair; metanotum brown,
166 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

tinged faintly with violaceous, delicately pollinose; pleural tuft silvery white; wing veins
whitish; setae at base of wings whitish; base of halteres brownish, knob yellowish white;
legs light brown; distribution of pigment on legs very similar to that of male; abdomen
dull silvery gray; distribution of pigmentation in accordance with figure B; frons like that
of Simulium nemorale; fronto-ocular triangle approximately as long as wide (Fig. C); shape
and proportions of antennal segments as shown in figure D; last segment of maxillary
palp (Fig. E) more than twice as long as penultimate, and approximately as long as two
preceding segments combined; diameter of sensory vesicle as large as half of diameter of
third segment; cibarium with well-sclerotized lateral projections (Fig. F); Sc in most cases
without, rarely with one or two, setae on its center; setae and spines on R1 arranged in one
row, in rare cases in irregular double row on basal portion of vein; setae on Rs arranged in
one series; calcipala very small, wider than long; claws with small tooth (Fig. G); posterior
portion of eighth sternite with 1+1 groups of approximately 35 long setae, their size
diminishing toward central region (Fig. H); central portion of sternite slightly less
pigmented than lateral areas; gonapophyses subtriangular, beset with microtrichia, their
internal borders more strongly sclerotized; paraprocts and cerci as shown in figure I; cerci
higher than wide, their distal border rounded; projecting portion of paraprocts subtrian-
gular, extreme apex rounded; genital fork (Fig. J) with median process well sclerotized, its
extreme apex widened; anterior projections narrow, with large lateral expansions at their
tips, their external border pigmented; spermatheca like that in Simulium nemorale.
Male: Wing length 2.3-2.6 mm; head, palpi, and labrum dark brown; eyes dark
purple; antennae blackish, first and second segments slightly lighter; frons, clypeus, and
labrum silvery pollinose; hair of palpi from silvery to black; antenna as in figure K;
scutum velvety black, lateral margins and anterior and posterior declivities silvery
pollinose, under certain angles with 1+1 submedian grayish spots adjacent to anterior gray
border (Fig. L); hair of scutum very dense, silvery, decumbent, giving scutum overall
grayish cast; metanotum dark brown, tinged with violaceous; pleura dark brown, grayish
pollinose; pleural tuft silvery white; legs from yellowish to brownish; distribution and
relative intensity of pigment of legs as shown in figures M, N, O; dorsal surface of
abdomen velvety black, with 1+1 silvery gray spots on tergites II, VI, and VII; ventral
surface of abdomen grayish brown; hair of abdomen, including that of basal fringe,
silvery white; last segment of maxillary palp longer than two preceding segments
combined; diameter of sensory vesicle about as large as half of diameter of third
segment, this segment notably darker than remainder; wings like those of female, but no
setae on Sc, and setae and spines on R1 invariably in single row, even at base of vein; hind
basitarsus narrow, 3.8 times as long as wide; calcipala very small, wider at base than long
(Fig. P); general structure of genitalia very similar to that of Simulium dureti; gonocoxite
and gonostylus as shown in figures Q, R; ventral plate with 1+1 distinct, translucent areas.
Pupa: Cocoon slipper shaped, with moderately extended anteroventral bridge (Fig. S);
color of cocoon light brown, surface smooth, closely woven, but threads perceptible under
moderate magnification; rim of aperture distinctly reinforced; gills covered by cocoon only
at their extreme base; length of cocoon along dorsal surface 2.7-3.4 mm; along ventral
surface 3.8-4.7 mm; length of body of pupa 2.8-3.0 mm; length of gill 1.2-1.6 mm; gills (Fig.
T) each with eight filaments, slightly diverging on basal, and approximately parallel on apical,
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 167

D
C

F A G
B

I
L J M
O
K
Q N

R S

V U

W a
b
Y
Figure 30. Simulium (Pternaspatha) pichi. A-J: Female; A: scutum; B: abdomen, dorsal view; C:
fronto-ocular triangle; D: antenna; E: maxillary palp; F: cibarium basal portion; G: claw; H: eighth
sternite; I: paraproct and cercus; J: genital fork. K-R: Male; K: antenna; L: scutum; M-O: fore, mid
and hind leg respectively ; P: distal portion of hind basitarsus; Q: gonocoxite and gonostylus; R:
gonostylus. S-W: Pupa; S: general aspect; T: gill; U: frontoclypeus; V: portion of thorax with
trichomes and gill base; W: thoracic trichome enlarged. X-Z: Larva; X: general aspect; Y: cephalic
apotome; Z: hypostomium; a: anal sclerite; c: rectal papillae.
168 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

half; frontoclypeus (Fig. U) with 3+3 branched frontal and 1+1 branched facial trichomes;
densely covered with small tubercles; exposed portion of thorax (Fig. V) covered with
numerous tubercles and branched trichomes, latter limited to posterior half of exposed
portion; trichomes slender, hair like, with two to five branches (Fig. W); average length of
trichomes 0.12 mm; total number of trichomes of thorax approximately 60+60; chaetotaxy
of abdomen like that in Simulium nemorale, but tubercles on tergites VI-VIII less numerous.
Larva: Maximum length, 6.7 mm; maximum width of head capsule, 0.6 mm; general
body shape as shown in figure X; color of larva light brown, with faint, dark, hypodermal
pigment on dorsal surface and on underside of anterior third of body; cephalic apotome
with faint brown pattern (Fig. Y); body integument smooth, hair present at base of anal
sclerite; antennae light brown, distal segment darkest; medial segment with two slight
constrictions, coinciding with unpigmented areas; ratio of lengths of proximal, medial
and distal segments = 1:1.1-1.15:0.9; cephalic fan with about 26 rays in large fan; anterior
border of hypostomium strongly pigmented (Fig. Z); median tooth almost as prominent
as lateral and outermost intermediate teeth, remaining intermediate teeth smaller;
sublateral tooth absent; lateral borders of hypostomium with seven or eight minute
serrations; number of hypostomial setae from 7 to 11 in each group, arranged in one,
rarely in two, irregular series; disc of hypostomium glabrous or with occasional simple
short seta; postgenal cleft deep, about twice as deep as postgenal bridge; anal sclerite as
shown in figure a; scales at base absent; anal ring with about 83 rows composed of about
18 hooks each; rectal papillae simple, composed of three simple lobes (Fig. b).
Distribution: Argentina: Chubut, Santa Cruz.
Bionomics: The larvae and pupae of Simulium pichi were collected in a crystal-clear
stream about 2 m wide and 1 m deep. The specimens were attached to stems and leaves of
Potamogeton sp. and Heleocharis sp., but mostly on reeds, generally on the downstream side of
the leaves.
Discussion: Simulium pichi resembles S. dureti, from which it differs by its smaller size, the
smaller extension of the anterior gray portion on the scutum of the male, the absence of a
median frontal blackish spot on tergite VI of the female, the absence of setae from Sc and the
more salient cerci in the same sex, fewer and less-branched cephalic and thoracic trichomes of
the pupa, and the different pigmentation of the larval head. Simulium limay also is similar, but
has fewer trichomes on the thorax (20-55) and the frontoclypeal tubercles have more rugosity,
and in the male pupa, the tubercles are only on the clypeal base. Simulium diamantinum also has
a similar pattern on the abdomen, but is larger (wing length 3.4-3.9 mm), and has the calcipala
absent, the claw subbasal teeth larger, the cocoon shoe shaped, more abundant frontal (5-15)
facial (3-6), and thoracic (160-120) trichomes, a relatively shorter proximal antennal segment in
larvae and more diverticula on the rectal papillae lobes (5-7).

Simulium (Pternaspatha) dureti WYGODZINSKY & COSCARÓN (Fig. 31.)


Simulium (Pternaspatha) dureti WYGODZINSKY & COSCARÓN, 1967: 62-67; COSCARÓN & WYGODZINSKY,
1972b: 209-210; COSCARÓN, 1991: 115-116.
Female: Wing length 3.6 mm; body with overall gray tinge; color of head like that of
Simulium nemorale, hair of clypeus silvery; color of scutum very similar to that of Simulium
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 169

nemorale, but grayish pattern more extended, occupying anteriorly one-third of surface of
scutum; appressed setae of scutum silvery, very dense, giving scutum overall grayish
aspect; scutellum grayish brown, its apical portion darker; pleural tuft silvery white; wing
veins stramineous; hair at base of wings silvery white; stem of halteres brown; their knob
yellowish white; legs light brown, their hair silvery to black; exact distribution and relative
intensity of pigment of legs as in figures F-H; overall color of abdomen (Fig. K) light
gray, dull; first segment darker; tergites II-V with well-developed, central, velvety black
spot, separated from 1+1 slightly less intense, lateral, dark spots by gray areas as wide as
central spots; tergite VI with very small, in some cases indistinct, black spot; lateral dark
areas present; tergites VII and VIII with large, dark gray spots at center, in some cases
entire tergites darkened; frons with obsolete median sulcus (Fig. A); fronto-ocular triangle
slightly longer than wide (Fig. B); shape and proportions of antennal segments as in figure
C; last segment of maxillary palp more than twice as long as penultimate and longer than
two preceding segments combined (Fig. D); diameter of sensory vesicle larger than half
diameter of third segment, its structure as shown in figure E; cibarium with well-
sclerotized, lateral projections; lacinia with about 25 teeth, mandible with approximately
40 teeth; Sc with not more than 10 setae on its central portion; setae and spines on R1
arranged in one, or in some cases two, irregular rows; setae on Rs arranged in one series;
shapes and proportions of segments of legs as shown in figures F, G, H; calcipala very
small but distinct, somewhat wider at base than long (Fig. I); claws with small tooth (Fig.
J); posterior portion of eighth sternite with 1+1 groups of long setae becoming
progressively shorter toward center; about 30 setae in each group (Fig. L); central portion
of eighth sternite less pigmented than lateral; gonapophyses subtriangular, covered with
microtrichia; their internal border sclerotized; paraprocts and cerci as shown in figure M;
cerci with apical border rounded, apex of paraprocts subrectangular; genital fork with
median process distinctly sclerotized, its extreme apex widened; anterior projections with
wide, lateral expansions; external border of distal portion pigmented; spermatheca like
that in Simulium nemorale.
Male: Wing length 3.0 mm; body with overall gray tinge; color of head and its
appendages like that in Simulium nemorale; scutum with color pattern of the group, but all
of anterior half gray under most angles except at middle, and posterior gray area also
more widely extended than usual (Fig. N); scutum with very numerous silvery, appressed
hair; pleura, halteres, and wings like those in female; legs light brown, their hair from
silvery to black; abdomen (Figs. O, P) with dorsal surface velvety black, undersurface
grayish; tergites II, VI, VII, and in some cases also VIII, with 1+1 light gray spots, smaller
and with bluish sheen on posterior segments; hair of abdomen including that of basal
fringe, silvery; last segment of maxillary palp approximately twice as long as penultimate;
diameter of sensory vesicle slightly less than half diameter of segment; wings like those in
female, but Sc devoid of hair, and spines and hair of R1 invariably in single row; shape,
proportion and coloration of segments of legs very close to those of female and as shown
on hind leg (Fig. Q); hind basitarsus 3.2-3.4 times as long as wide; calcipala very small but
distinct, approximately as long as wide at base; gonocoxite and gonostylus as in figure R;
gonostylus subquadrate, apical concavity beset with very numerous heavily pigmented
tubercles (Fig. S); ventral plate as in figure T, without distinct translucent areas.
170 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

A
G
C B F

D E I H

N P
K O M
L

S X
Q

V U

W Z
Y
d
b

a
e
f c
Figure 31. Simulium (Pternaspatha) dureti. A-M: Female; A: frons; B: fronto-ocular triangle; C:
antenna; D: maxillary palp; E: sensory vesicle of maxillary palp; F-H: fore, mid and hind leg,
respectively; I: apex of hind basitarsus and tarsomere I; J: claw; K: abdomen, dorsal view; L: eighth
sternite and gonapophyses; M: cercus and paraproct. N-T: Male; N: scutum; O-P: abdomen in
dorsal and lateral views; Q: hind leg; R: gonocoxite and gonostylus, ventral view; S: gonostylus,
dorsal view; T: ventral plate. U-Z: Pupa; U: lateral view; V-W: gill in different specimens; X:
frontoclypeus; Y: exposed portion of thorax with trichomes and tubercles; Z: trichomes with high
magnification. a-f: Larva; a: cephalic apotome; b: antenna; c: teeth of mandible; d: hypostomium; e:
cephalic capsule in ventral view; f: anal sclerite and trichomes.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 171

Pupa: Pupae single or forming compact groups; cocoon shoe shaped, with large,
anteroventral bridge; surface of all cocoons examined covered with foreign particles of
various sizes (Fig. U); cocoon closely woven, threads not individualized; rim of aperture
slightly reinforced; gills covered laterally on basal half by walls of cocoon, in some cases
only extreme apex visible; length of cocoon at dorsum along middle to border of
aperture, 3.3-4.0 mm; maximum length along base to anterior border of antero-ventral
bridge, 4.3 mm; length of body of pupa 3.5-4.5 mm; length of gill 2.0-2.5 mm, viz., less
than half as long as cocoon, and slightly more than half as long as pupa proper; gill with
eight filaments (Figs. V, W), head and thorax of pupa light brown; clypeus covered with
small tubercles; head with 1+1 groups composed each of approximately 17 branched
frontal and facial trichomes (Fig. X) and 1+1 groups of three branched, ocular trichomes
each; exposed portion of thorax more strongly sclerotized than remainder, covered with
tubercles and long, branched trichomes; tubercles are abundant and those situated toward
midline more or less pointed, spinelike; trichomes with as many as 8 branches (Figs. Y, Z);
their average length, 0.2 mm; total number of thoracic trichomes approximately 150+150;
chaetotaxy of abdomen like that in Simulium nemorale.
Larva: Maximum length, 8.5 mm; width of head capsule, 0.7 mm; color yellowish,
with dark greenish brown hypodermal pigment over whole body surface, especially
intense on posterior half dorsally; cephalic apotome dark with negative spots (Fig. a);
body integument smooth, with short setae at base of anal sclerite; antennae dark brown,
pigment most intense on subapical segment (Fig. b); ratio of proximal, medial and distal
segments = 1:1.3-1.45:0.7-0.8, viz., last segment distinctly shorter than first; cephalic fans
with approximately 46 rays in large fan; toothing of mandible as shown in figure c; two
external, one strong apical, three subapical, four internal, and two external teeth; anterior
border of hypostomium strongly pigmented (Fig. d); median tooth as prominent as lateral
ones, intermediate teeth gradually increasing in size from submedian to lateral ones;
sublateral tooth barely perceptible; lateral borders of hypostomium with five to seven very
small serrations; number of hypostomial setae about 14 in each group, arranged in two
irregular series; disc of hypostomium with some short, simple setae; postgenal cleft deep
(Fig. e), almost twice as deep as hypostomial bridge; anal sclerite as shown in figure f,
without scales at its base; anal ring with approximately 90 rows, each composed of about
17 hooks; rectal papillae with three simple lobes.
Distribution: Argentina: Jujuy, Salta; Bolivia: Potosí.
Bionomics: Larval stages live in torrential creeks at 2000 to 4300 m altitude, flowing
over stones and leaves.
Discussion: Simulium dureti is close to Simulium pichi and S. deagostinii. The differences for
S. pichi were treated under that species. Simulium deagostinii shows a more darkened abdominal
tergite VI in the female and no calcipala In the pupa, the frontal and facial trichomes are fewer
in number (5-6 on each side), and the larva has 6 diverticula on the median lobe and 4 on each
lateral lobe of the rectal papillae. Simulium diamantinum is another similar species that can be
differentiated by the absence of a blackish median anterior spot on female tergite VI, wider
blackish spot on tergite VII, shorter blackish 1+1 lateral spots on tergites III-V, calcipala
absent, thoracic trichomes more abundant (160-220), relatively shorter proximal segment of
the larval antenna, and more diverticula on the rectal papillae lobes (5-7).
172 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Simulium (Pternaspatha) horcochuspi COSCARÓN & WYGODZINSKY (Fig. 32.)


Simulium (Pternaspatha) horcochuspi COSCARÓN & WYGODZINSKY, 1972b: 210-212; COSCARÓN, 1991: 118.
Female: Wing length, 2.9-3.6 mm; color of head and thorax as in Simulium nemorale;
appressed setae of scutum silvery, not very dense; overall color of dark portions of
scutum distinctly blackish; scutellum and metanotum gray; pleural tuft silvery white; color
of wings, halteres, legs and abdomen much as in Simulium dureti; head and its appendages
and general shape of legs as in S. dureti; calcipala absent (Fig. A); claw without tooth, only
with occasional slight angular projection in place of tooth (Fig. B); genitalia as in S. dureti.
Male: Wing length about 2.8 mm; color of body and legs close to Simulium dureti;
general structure as in S. dureti, including genitalia; apex of gonostylus granulose; length/
width ratio of hind basitarsus = 3.5; calcipala absent.
Pupa: Pupae single or forming groups; cocoon with large anteroventral bridge (Fig.
C); cocoon very thin, translucent, with conspicuous leaden sheen; cocoon closely woven;
individual threads difficult to perceive; rim of aperture slightly reinforced; sides of
cocoon covering basal half or two thirds of gills; length of cocoon at dorsum along
middle to border of aperture 2.4-3.4 mm; maximum length along base to anterior border
of anteroventral bridge, 4.6-5.2 mm; length of body of pupa 2.9-3.2 mm; length of gills
1.5-1.8 mm., viz., about half as long as pupa proper; gills with eight filaments, arising from
three primary branches (Figs. D, E); frontoclypeus covered with small tubercles, more
numerous in male (Fig. F) than in female (Fig. G); surface of tubercles smooth; head with
3(-4)+3(-4) frontal, 1+1 facial and 1-2+1-2 ocular trichomes; these trichomes hair like,
with 1-4 branches (Fig. H); exposed portion of thorax covered with very numerous
tubercles forming irregular groups (Fig. I) and long hair like trichomes; number of
trichomes 75-85+75-85, each with from one to three, rarely four, branches, most
frequently two; average length of trichomes 0.25 mm; abdomen as in Simulium dureti.
Larva: Maximum length of mature larva, 8.0 mm; maximum width of head capsule,
0.75 mm; general body color whitish, but almost entire body with very intense dark brown
hypodermal pigment; pattern of cephalic apotome negative; general body shape as in
Simulium dureti; antennae as shown in figure J, similar to those of S. dureti, but distal segment
about same length as proximal; cephalic fans, maxillary palp, mandibles, hypostomium,
postgenal cleft, anal ring, and rectal papillae as in S. dureti; anal sclerite as figure K.
Distribution: Argentina: Jujuy.
Bionomics: Simulium horcochuspi was collected between 3900 and 4700 m of altitude
in creeks of reduced volume and crystalline water flowing over stones.
Discussion: Simulium horcochuspi is close to S. deagostinii and S. dureti but S. deagostinii
has a median black spot on female tergite VI. The female of S. horcochuspi differs from the
two related species mentioned, by the absence of a process on the claws; it differs further
from S. dureti, with which it might possibly be found together, by the light-colored area of
the anterior portion of the scutum, which is much less extensive than in S. dureti. The male
of S. horcochuspi, known only from a pharate specimen, cannot be distinguished from that
of S. deagostinii, but differs from that of S. dureti by the absence of a calcipala. The pupa
of S. horcochuspi is characterized by the leaden sheen of its cocoon, which distinguishes it
immediately from the two other species, and thoracic trichomes that in S. dureti number
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 173

B
H
A
C

I
E
D

F G K

Figure 32. Simulium (Pternaspatha) horcochuspi. A-B: Female; A: apex of hind basitarsus and
pedisulcus; B: claw. C-I: Pupa; C: pupa in lateral view; D-E: gill in different specimens; F-G: portion
of frontoclypeus, showing different number of tubercles (F: male, G: female); H: frontoclypeus; I:
trichomes of thorax. J-K: Larva; J: antenna; K: anal sclerite.

about 150. Further differential characters, mostly relating to the number and structure of
its cephalic and thoracic trichomes, serve to distinguish S. horcochuspi from S. dureti and S.
deagostinii, as shown in the key. In the larva, S. dureti and S. deagostinii have the distal
segment of the antenna shorter than the proximal.

Simulium (Pternaspatha) simile SILVA FIGUEROA (Fig. 33.)


Simulium simile SILVA FIGUEROA, 1917: 33; PINTO, 1931: 730; VARGAS, 1945a: 196; BEQUAERT, 1945:
114; STUARDO ORTIZ, 1946: 41; WYGODZINSKY, 1949: 304; 1958: 133; STONE, 1962: 207.
Simulium (Simulium) simile EDWARDS, 1931: 153; ORFILA, 1939: 1534.
Acropogon simile ENDERLEIN, 1934b: 280.
Simulium (Pternaspatha) simile WYGODZINSKY & COSCARÓN, 1967: 79; COSCARÓN & WYGODZINSKY,
1972b: 216-218; COSCARÓN, 1991: 119.
[Simulium (Pternaspatha)] simile STONE, 1963: 17.
Simulium figueroa SMART, 1944: 133; 1945: 505; VARGAS & DIAZ, 1953b: 141;VULCANO, 1967: 11.
Simulium illiesi WYGODZINSKY & COSCARÓN, 1967: 75.
Acropogon punctativentris ENDERLEIN, 1936: 118.
Simulium punctativentris SMART, 1945: 512; VARGAS, 1945a: 188; STUARDO ORTIZ, 1946: 41; VULCANO,
1967: 19.
174 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Simulium (Pternaspatha) punctativentris WYGODZINSKY & COSCARÓN, 1967: 115; COSCARÓN & WYGODZ-
INSKY, 1972b: 218.
Simulium barbatipes VARGAS & DIAZ, 1953b: 141 (part).
Female: Wing length 3.2-3.5 mm; head blackish; antennae, palpi, and labrum
piceous; scape and pedicel dark orange; frons and clypeus gray pollinose; pilosity of
antennae very short, that of palpi, clypeus, frons, and occiput somewhat longer, silvery
gray; scutum as in Simulium nemorale; hair of thorax silvery gray; appressed setae of scutum
very dense, somewhat obliterating pattern; scutellum, metanotum, pleura, and sterna dark
gray; pleura with silver pollinosity; legs similar to those of Simulium pulchrum; color of
abdomen blackish with tergite II with 1+1 large, tergites III-V with 1+1 small, and tergites
VI and VII again with 1+1 large, silvery white spots; tergites VIII and IX extensively
silvery white (Fig. A); fronto-ocular triangle about as long as wide (Fig. B); maxillary palp
as in figure C; diameter of sensory vesicle equal to half of width of third palpomere;
structure of vesicle as shown in figure D; lacinia with 28-29 teeth, mandible with
approximately 43 teeth; calcipala well developed (Fig. E); claws with relatively large tooth
(Fig. F); eighth sternite as illustrated in figure G, its central portion more strongly
sclerotized than lateral areas; gonapophyses triangular, beset with microchaetae, their
inner border faintly sclerotized; cerci with distal border rounded; paraprocts salient, their
free portion subtriangular (Fig. H); genital fork as illustrated (Fig. I).
Male: Wing length more than 3.5 mm; color of head and cephalic appendages like
those in Simulium nemorale; thorax black, with abundant, appressed, golden setae; abdomen
blackish with ornamentation as shown in figures J-K; hind basitarsus 4.5 times as long as
wide; calcipala well developed but shorter than wide at base; gonostylus with distal
portion widely concave, heavily tuberculate (Fig. L); ventral plate as in figure M.
Pupa: Cocoon shoe shaped (Figs. N-P), with large, anteroventral bridge; color of
cocoon light brown; surface smooth, closely woven, but threads individualized; rim of
aperture distinctly reinforced; gills protected laterally by wall of cocoon; length of cocoon
at dorsum 4.0-4.5 mm; total length to border of anteroventral bridge, 5.5-7.0 mm; length
of body of pupa 4.0-5.0 mm; length of gills 1.8-2.0 mm, slightly less than one-third of
length of cocoon, and not quite half as long as pupa proper; gills whitish, each with eight
filaments arranged in fan-like pattern (Figs. P, Q); head and thorax of pupa dark brown;
frontoclypeus (Fig. R) with approximately 40 frontal and facial, and apparently 4+4
ocular, trichomes, their shape as described below for thoracic trichomes; clypeus heavily
beset with tubercles; thorax (Fig. S) strongly sclerotized on exposed portion, latter heavily
beset with tubercles and with large number (approximately 140+140) of trichomes;
trichomes (Figs. T-V) narrow at extreme base, then flattened and band-shaped, apically
again narrowed and pointed; trichomes simple or divided near base into two to four
branches, latter closely parallel and difficult to individualize; tubercles of frontoclypeus at
base not distinctly surpassing area of frontal trichomes; tubercles in many cases with
sharply pointed protuberances (Fig. U); area of facial trichomes with conspicuously
marked depression (Fig. R); borders of depression heavily sclerotized.
Larva: Maximum length, 9.0 mm; maximum width of head capsule, 0.8 mm; body
dark greenish gray; head dark brown; antenna as in figure W; ratio of proximal, medial and
distal segments = 1:1.6:0.8; cephalic fans with 28-32 rays in large fan; toothing of
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 175

B E

C G
I

A L
D
H
K O

M N
J
P

S
T

U Q
V

W
Z
Y X
Figure 33. Simulium (Pternaspatha) simile. A-I: Female; A: abdomen, dorsal view; B: fronto-ocular
triangle; C: maxillary palp; D: sensory vesicle; E: hind basitarsus showing calcipala; F: claw; G:
portion of eighth sternite and gonapophyses; H: cercus and paraproct; I: genital fork. J-M: Male; J-
K: abdomen in dorsal and lateral views; L: gonocoxite and gonostylus in ventral view; M: ventral
plate. N-V: Pupa; N-P: lateral, dorsal and ventral view, respectively; Q: gill; R: frontoclypeus,
showing trichomes and tubercles; S: portion of exposed portion of thorax, with abundant
trichomes; T-V: different trichomes with high magnification. W-Z: Larva; W: antenna; Y: portion of
hypostomium; X: teeth of mandible; Z: cephalic capsule in ventral view.
176 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

mandibles as shown in figure X; pigment pattern of hypostomium as shown in figure Y;


median tooth more pronounced than lateral ones; lateral serrations distinct; hypostomial
setae arranged in two or three irregular rows 21-24 in each group; postgenal cleft pointed,
very deep, its apex very close to hypostomial groove (Fig. Z); anal sclerite at its base with
simple to trifid scales, few in number; anal ring with 95-118 rows, each composed of 18-
19 hooks; rectal papillae with three primary lobes, each bearing three pairs of long
secondary lobules; median pair in some cases with four pairs of secondary lobules.
Distribution: Argentina: Chubut, Neuquén, Rio Negro; Chile: from Tarapaca to
Bio-Bio.
Bionomics: Simulium simile is a medium- to high-altitude species which prefers
colder, very fast-flowing streams.
Discussion: Simulium simile is characterized by the unique chaetotaxy of the pupal
thorax. In the shape of its cocoon it resembles Simulium nemorale, but the latter has a very
different thoracic chaetotaxy. The male can be distinguished from that of similar species as
shown in the key. The female is not well enough known to be compared with females of
related species. Simulium simile is very close to S. pulchrum PHILIPPI redescribed below; the
differences between these two species are indicated in our keys, and summarized in a table
given under the heading of S. pulchrum. These species also differ in habitat preferences.

Simulium (Pternaspatha) pulchrum PHILIPPI (Fig. 34.)


Simulium pulchrum PHILIPPI, 1865: 633; KERTESZ, 1902a: 290; SILVA FIGUEROA, 1917: 30; PINTO, 1931:
729; VARGAS, 1945a: 187; SMART, 1945: 512; STUARDO ORTIZ, 1946: 42; VULCANO, 1967: 19.
Simulium (Pternaspatha) pulchrum WYGODZINSKY & COSCARÓN, 1967: 115; COSCARÓN & WYGODZINSKY,
1972b: 204, 213-216; COSCARÓN, 1991: 118-119.
Female: Wing length 2.6-2.8 mm; color of head, thorax, and legs as in S. simile; legs
brownish with apex of segments more darkened as shown in figures B-D; abdomen close
to Simulium limay but sublateral blackish spots on tergites III-V wider (Fig. A); fronto-
ocular triangle, slightly deeper than wide (Fig. E); shape and proportions of antennal
segments as shown in figure F; diameter of sensory vesicle of third palpomere equal to
half width of segment (Fig. G); lacinia with 25-26 teeth, mandible with 38-41 teeth;
calcipala, claws and wings as in Simulium simile; general structure of genitalia as in Simulium
simile, but paraproct less salient.
Male: Wing length 2.6-2.7 mm; color of head and its appendages, thorax wings, and
halteres as in Simulium simile; legs as in female, although somewhat darker; abdomen
velvety black dorsally, except dark gray on tergum IX; terga II, VI and VII with 1+1 large,
silvery white spots; terga III-V entirely black; antennae as shown in figure H; hind
basitarsus five times as long as wide; genitalia as in Simulium simile.
Pupa: Structure of cocoon as in Simulium simile; its length dorsally 2.2-2.7 mm,
maximum length along its base 3.8-4.5 mm; length of body of pupa 2.5-3.2 mm, of gills 1.1-
1.3 mm; head of pupa light brown; chaetotaxy of head (Fig. I) much as in S. simile, but
tubercles of frontoclypeus more numerous (about 800) than in S. simile, distinctly surpassing
level of frontal trichomes, attaining level of uppermost facial trichomes; protuberances of
tubercles not pointed; thorax of pupa much as in S. simile, but trichomes less numerous
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 177

B C
F
E G

H
A
D

I J

Figure 34. Simulium (Pternaspatha) pulchrum. A-G Female; A. abdomen, dorsal view; B-D: fore, mid
and hind leg, respectively ; E: fronto-ocular triangle; F: antenna; G: sensory vesicle of maxillary
palp. H: Male, antenna. I: Pupa, frontoclypeus with abundant tubercles and trichomes. J: Larva,
cephalic apotome of different specimens.

(normally 70-90, occasionally up to 130, on each side), and tubercles more numerous; length
of trichomes 0.15-0.28 mm, much shorter than in S. simile; abdomen as in S. simile.
Larva: Length of mature larva, 5.6-6.0 mm; maximum width of head capsule, 0.8 mm;
general body shape as in Simulium simile; color light greenish brown, head light brown; cephalic
apotome very light, dark pigment restricted to base (Fig. J); general aspect of antennae as in S.
simile; ratio of proximal, medial and distal segments = 1:1.5:1.1; cephalic fan with 29-33 rays;
maxillary palpi, hypostomium, postgenal cleft and anal sclerite as in S. simile; anal ring with 85-
100 rows of about 16 hooks each; rectal papillae in most specimens with three simple lobes,
with one pair of diverticula in specimens from the province of Aconcagua, and two pairs
diverticula on each lobe in specimens from southern part of range.
Distribution: Chile: Aconcagua, Colchagua, Curico, Nuble, O’Higgins, Santiago, Talca.
Bionomics: The larvae and pupae of this species were found attached to stems of
grasses (such as Scirpus sp.) trailing in the water, or to branches of trees like Salix trailing
in the stream, but only rarely to rocks. The species was collected mostly in large, fast-
flowing streams or irrigation ditches. It prefers clear streams, with only occasional
turbidity. This is a lowland species in Central Chile, collected generally at 400-800 m; the
highest elevation recorded was 1100 m.
Discussion: Simulium pulchrum is closely related to Simulium simile, from which it can
be distinguished as shown in table 1.
178 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Table 1. Differences between Simulium pulchrum and Simulium simile (All measurements are in
millimeters.)

Character Simulium pulchrum Simulium simile


ADULTS
Wing length, female 2.6-2.8 3.2-3.5
Wing length, male 2.6-2.7 3.2-3.5
Darkened portion of posterior tibia of Female Apical fifth Apical two-fifths
Tergum III of abdomen of male Black with 1 + 1 small
whitish spots
Terga IV and V of abdomen of male Black with 1 + 1 very small
whitish spots
PUPAE
Length of cocoon, dorsally 2.2-2.7 3.3-4.5
Length of cocoon, ventrally 3.8-4.5 5.2-6.8
Length of body of pupa 2.5-3.2 3.2-4.5
Length of gills 1.1-1.3 1.8-2.0
Number of tubercles of frontoclypeus +800 +400
Shape of projections of tubercles Not pointed Pointed
Area of facial trichomes almost smooth Distinct impression
limited by carinae
Length of thoracic trichomes 0.15-0.30 0.3-0.5
Maximum number of trichomes of thorax 130+130 250+250
Average number of trichomes of thorax 70+70 140 + 140
LARVAE
Color of head light brown dark brown
Color of body light greenish brown dark greenish brown
Maximum length 6.0 9.0
Maximum width of head capsule 0.6 0.8
Number of rows in anal ring 85-100 95-118
Number of hooks per row 14-17 18-19
Number of hypostomial setae 13-18 21-24
Number of diverticula to each rectalPapillae 1 or 2 3, rarely 4

Simulium (Pternaspatha) diamantinum COSCARÓN & COSCARÓN ARIAS (Fig. 35.)


Simulium (Pternaspatha) diamantinum COSCARÓN & COSCARÓN ARIAS, 1996a: 703.
Female: wing length 3.3-3.7 mm; general color gray pollinose; antennae, maxillary
palp and proboscis blackish; head brown, dark gray pollinose; frons, clypeus and occiput
blackish with gray pollinosity; occiput a little darker with hair grayish to black in
accordance with light; scutum grayish black with silvery-gray, submedian, lyre-shaped
pattern, lighter anteriorly and grayish laterally and posteriorly (Fig. A); with changing light
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 179

position lyre-shaped pattern becomes thicker, and in some lights its anterior border
becomes black, to silky bright light gray; scutum hair sparse and upright; scutellum
blackish-gray; scutum and scutellum hair light gray; metanotum velvety blackish; pleura
blackish gray; legs blackish with basal third of femur III, basal half of tibiae I-III, basal
half of basitarsus II and 3/4 of basitarsus III and base of tarsomeres II-III light
brownish; distribution of pigmentation of hind leg as in figure G; abdomen light grayish
pollinose, tergite I dark grayish; black subtriangular median spots on tergites II-V, lighter
and pollinose on VII, and 1+1 lateral, blackish, subrectangular spots not reaching
posterior border on tergites IV-V (Figs. B, C); hair of pleural tuft light gray; frons as in
figure D; fronto-ocular triangle about as long as wide (Fig. E); sensory vesicle of palp
about 1/3-1/4 of third palpomere, its structure as in figure F; mandible with 25+8-30+9
teeth, and lacinia 23-25 teeth; Sc with 3-5 setae; base of R without setae; R1 and R with
setae and spines arranged in one irregular row; shape and proportion of segments of hind
leg as shown in figure G; without calcipala; ratio of length to width of hind basitarsus =
7.2-7.6; claws with well-developed basal tooth (Fig. H); eighth stemite well sclerotized,
with 34-40 hairs on each side (Fig. I), gonapophyses subtriangular; paraproct and cercus as
in figure J; genital fork as in figure K.; spermatheca with small internal spines (Fig. L).
Male: Wing length 2.8-3.0 mm; head coloration as in female; scutum black-gray with
1+1 subtriangular spots anteriorly (Fig. M); with light anterior to specimen 1+1 anterior
gray pollinose spots occupying anterior third; decumbent, silky grayish hair sparsely
covering scutum; abdomen blackish with 1+1 large gray spots on tergite II, smaller on
tergites VII-VIII, VI and IX mostly grayish except for small and short median darker spot;
legs yellowish brown with light gray hair; blackish brown on coxae, apex of femur III,
apex of tibiae, all of basitarsus I, distal third of basitarsi II and III and tarsi; hind legs as
in figure N, basitarsus three times as long as wide; calcipala absent; genitalia very close to
species of Simulium (Pternaspatha) NIGRISTRIGATUM species group; gonostylus with
abundant tubercles (Fig. O).
Pupa: Cocoon shoe shaped with well-developed antero-ventral bridge (Fig. P);
surface generally smooth, but some specimens with small sand granules incorporated;
cocoon closely woven; gill covered basally by walls of cocoon; length of cocoon 3.8-4.0
mm at dorsum, maximum length along base 5.2-6.0 mm, length of body 4.0-4.7 mm, of
gills 2.2-3.3 mm; gills with eight filaments, position of basal branching as in figure Q;
frontoclypeus and thorax with 500-1000 tubercles (Fig. R), those of thorax slightly
pointed, especially on posterior border (Fig. S), others rounded with smooth or slightly
rough surface (Fig. T); trichomes filiform with 4-8 branches, 5-15 frontal and 3-7 facial
trichomes on each side in continuous group; length of frontal trichomes 0.19-0.23 mm;
thoracic trichomes 0.24-0.35 mm, 160-220 in number, with 6-10 filiform branches (Fig.
U); chaetotaxy of abdomen following general pattern shown in Simulium nemorale.
Larva: Maximum length 7.5-8.5 mm; head width 0.7 mm; color dark, greenish
brown, ornamentation and shape in accordance with figure V; integument smooth;
cephalic apotome brown with negative spots without well defined borders (Fig. W);
antennae dark brown; ratio of proximal, media and distal segments = 1:1.8-1.9:1.2-1.3
(Fig. X); cephalic fans with 35-39 combs; mandible with 4-6 secondary teeth arranged in
several rows (Fig. Y); teeth of hypostomium border as in figure Z, with 5-6 lateral
180 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

0.05 mm
0.05 mm

E F 0.0
0.1 mm 2m

B D m

A H

C 0.2 mm
0.1
m
m I
0.2 mm J
L
0.1 mm

0.1 mm
G
K M

O
m
5m

N
0.0

U 0.0
5m
m
R P
0.2 mm

T
0.2
m
m

0.0
1 mm 3m

V
m

Q S
a
W
m
0.1 m

0.1 mm

Z
m
5m

0.03 mm
0.0

X
Y
b c

Figure 35. Simulium (Pternaspatha) diamantinum. A-L: Female; A: scutum; B-C: abdomen in dorsal
and lateral views; D: frons; E: fronto-ocular triangle; F: second palpomere with sensory vesicle; G:
hindleg; H: claw; I: eighth sternite and gonapophyses; J: cercus and paraproct; K: genital fork; L:
spermatheca. M-O Male; M: scutum; N: hind leg; O: gonostylus. P-U : Pupa; P: lateral view; Q: gill;
R: frontoclypeus; S-T: tubercles of thorax pointed and rounded; U: thoracic trichomes. V-Z, a-c :
Larva; V: general aspect in lateral view; W: cephalic apotome; X: antenna; Y: teeth of mandible; Z:
hypostomium and postgenal cleft; a: proleg sclerite; b: anal sclerite; c: rectal papillae.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 181

serrations and 15-17 lateral setae arranged in 1-2 rows; postgenal bridge short, ratio of
hypostomium/postgenal cleft = 1.4-1.5; comb of thoracic proleg with about 43-54 teeth
arranged in 12-14 groups (Fig. a); anal sclerite without scales but with isolated trichomes
as in figure b; anal ring with about 94-100 rows, each composed of about 15-17 hooks;
rectal papillae with 5-7 diverticula on each of the three well developed lobes (Fig. c).
Distribution: Argentina: Mendoza, Neuquén, San Juan.
Bionomics: Simulium diamantinum breeds in creeks with strong current, crystalline and
cold waters on aquatic vegetation at 3000 m altitude. Females show strong anthropophily.
Discussion: Simulium pichi, S. dureti, S. diamantinum, S. horcochuspi and S. deagostinii are
very closely related species that can be distinguished by some morphological characters
and different distributions. Simulium diamantinum can be distinguished from the other
species by several distinctive characters such as the female abdomen with 1+1 small,
narrow and short, black lateral spots on tergites III-V, well isolated from the median black
spot and leaving a wide white transverse band before the posterior border; scutum hair
not appressed; narrower posterior basitarsi; longer pupa gill; greater number of frontal
and thoracic trichomes and longer thoracic trichomes. The larva has a longer medial
antennal segment, more lateral setae on the hypostomium, and a shorter postgenal bridge
in relation to the hypostomial length.

Simulium (Pternaspatha) strigidorsum (ENDERLEIN) (Fig. 36.)


Acropogon strigidorsum ENDERLEIN, 1934b: 279.
Simulium strigidorsum SMART, 1945: 514; VARGAS, 1945a: 199.
Simulium (Pternaspatha) strigidorsum WYGODZINSKY & COSCARÓN, 1967: 74-75; COSCARÓN & WYGODZ-
INSKY, 1972b: 218-219; COSCARÓN, 1991: 121.

Female: Wing length 3.3-3.6 mm; head black; antennae and mouthparts piceous;
occiput, frons, and clypeus gray pollinose; pilosity of antennae very short, of palpi
somewhat longer; occiput with long hair; scutum dark gray to black, with 1+1 light gray,
median, longitudinal stripes arising from 1+1 anterior white spots; narrow, median,
longitudinal line separated from gray stripes on its central half by elongate blackish areas;
stripes and midline confluent on posterior third of sclerite, connected to equally gray though
somewhat darker lateral borders of scutum; width of submedian gray stripes each about
one-eighth of total width of scutum; pubescence of scutum sparse and short, decumbent,
silvery; scutellum and metanotum piceous, silvery pollinose; scutellum with long decumbent
and erect silvery hair; pleural tuft silvery; pleura and sterna piceous, silvery pollinose; wing
veins stramineous, hair and setae black, tufts of hair at base of wings whitish; coloration of
legs of lectotype as in figure A, B, shows more darkened fore legs and femur of hind legs;
coxae and trochanters blackish, gray pollinose; forelegs entirely dark brown (Fig. A); basal
half of tibia, basal three-fourths of first and basal half of second tarsal segment yellowish;
hind legs dark brown; extreme base of femur and proximal half of tibia yellowish, mid legs
brown, basal half of tibia, basal three-fourths of first and basal half of second tarsal
segments yellowish; hind leg dark brown, extreme base of femur and proximal half of tibia
yellowish; basal two-thirds of first and proximal half of second tarsal segment whitish; very
182 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

small dark spot on base of second tarsal segment (Fig. B); dorsal surface of abdomen dull
silvery gray; first segment darkened, its posterior fringe whitish; tergites II-V each with
median velvety black spot (Fig. C); lateral dark spots or areas absent; tergite VI entirely gray;
tergite VII with distinct gray spot, VIII faintly darkened at center; frons wide; fronto-ocular
triangle about as long as wide (Fig. D); shape and portion of antennal segments as shown in
figure E; last segment of maxillary palp more than twice as long as penultimate and longer
than two preceding segments combined (Fig. F); diameter of sensory vesicle about half as
large as diameter of third segment of palp; wings as usual for subgenus; Sc and basal portion
of R bare; R1 with setae and spines arranged in single row, setae of Rs in single row; calcipala
(Fig. G) very small, in shape of small protrusion; claws with small tooth (Fig. H); genitalia as
illustrated in figures I-K; central portion of eighth sternite less strongly pigmented than
lateral areas, only its hind margin narrowly dark; setae of lateral portion of sternite not very
numerous (Fig. I); gonapophyses membranous, subtriangular, beset with microtrichia; cerci
rather short (Fig. J); paraprocts somewhat more intensely pigmented than cerci, their free
extremity very shortly salient; spermatheca subglobular with spiculae on inner surface;
genital fork as shown in figure K.
Male, pupa and larva: Unknown.
Distribution: (?Argentina: Chubut and Mendoza); Peru: Puno.
Bionomics: Unknown.
Discussion: The differences between the types from Peru and the Argentinean
specimens are very slight, and we do not hesitate to consider all specimens as conspecific.

F
0.3 mm
D B
E

I
C

H
K A

J G
Figure 36. Simulium (Pternaspatha) strigidorsum. A-K: Female; A: fore leg; B: hind leg; C: abdomen,
dorsal view; D: fronto-ocular triangle; E: antenna; F: maxillary palp; G: apex of hind basitarsus and
tarsomere I; H: claw; I: portion of eighth sternite and gonapophyses; J: cercus and paraproct; K:
genital fork. (D-F taken from Mendoza specimens; A-C, G-J from type).
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 183

Preimaginal stages from the type habitat could help to elucidate the identity. The color
pattern of the abdomen of the female without blackish lateral areas is unique in the
subgenus. Simulium diamantinum from the same place in Mendoza shows some similarities.
The female abdomen of S. diamantinum has 1+1 blackish lateral spots on tergites III-V, and
the legs are lighter. Several other species of the Simulium NIGRISTRIGATUM species
group are similar in abdominal ornamentation, such as S. pichi, which is smaller (wing length
2.4-2.9 mm), and S. dureti, which has a small subbasal claw tooth. Simulium herreri has smaller
light gray spots on tergites IV-VIII of the female abdomen and a subbasal claw.

Simulium (Pternaspatha) deagostinii COSCARÓN & WYGODZINSKY (Fig. 37.)


Simulium deagostinii COSCARÓN & WYGODZINSKY, 1962: 322-332; VULCANO, 1967: 9.
Simulium (Pternaspatha) deagostinii WYGODZINSKY & COSCARÓN, 1967: 61-62; COSCARÓN & WYGODZIN-
SKY, 1972b: 209; COSCARÓN, 1987: 18; 1991: 117.

Female: Wing length 3.4-3.5 mm; frons and clypeus blackish with gray pollinosity;
scutum blackish gray with 1+1 submedian longitudinal light gray stripes defining dark
bands (Fig. A); abdomen grayish, black dorsally with 1+1 submedian wide silvery gray
spots on tergites II-VII, wider on II, VI-VII (Fig. B); pleural and ventral areas grayish-
brown; legs brown with hair silvery to black, darkened on coxa, apex of femur, tibia and
tarsi as in figures E, F, G; fronto-ocular triangle wider than high (Fig. C); distal palpomere
length shorter than two before combined (Fig. D); calcipala absent (Fig. H), and claws
with small subbasal tooth (Fig. I); eighth sternite homogeneously darkened (Fig. J),
paraproct prominent distally (Fig. K).
Male: Wing length 3.3-3.4 mm; scutum black bordered anteriorly, laterally, and
posteriorly by grayish band; legs dark brown with darkened areas as shown on hind leg
(Fig. L); abdomen velvety black dorsally, grayish ventrally; tergites II, VI, VIII with 1+1
sublateral light gray spots, more developed on tergite II and getting narrower gradually
from VI to VIII; ratio of length/width of hind basitarsus = 3.2; calcipala not developed;
gonocoxite and gonostylus as in figure M, the latter with abundant tuberosity; ventral
plate as in figure N.
Pupa: Cocoon length on dorsum 4.0 mm, at base 4.5-5.0 mm; color light brown,
shoe shaped, strongly projected anteriorly, protecting base of gills (Fig. O); length of
pupal body 3.0 mm, gill 2.4 mm; frontoclypeus and thorax dorsally with abundant
tubercles; cephalic trichomes 5 or 6 in number on each side with 5 or 6 branches (Fig. P);
thorax with 65-95 trichomes (Fig. Q) with 2-8 branches (Fig. R) covering dorsally all free
area; gill with 8 branches (Fig. S).
Larva: Maximum length on dorsum 8.0 mm; color generally light grayish brown;
cephalic apotome light brown, mostly darkened basally (Fig. T); ratio of segments proximal,
medial and distal = 1:1.4:0.85; cephalic fan with about 45 rays; hypostomium with median
teeth at same level as lateral teeth; plate with 11-13 setae on each side, arranged in more than
one row; postgenal cleft deep, postgenal bridge about 1/5 of hypostomium length (Fig. U);
anal sclerite without scales; anal ring with 95 rows of 25-28 hooks each; rectal papillae with
6 diverticula on median lobe and 4 on each lateral lobe (Fig. V).
Distribution: Argentina: from Tierra del Fuego to center of Chubut.
184 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Bionomics: Simulium deagostinii breeds in rivers or large-volume creeks, with fast-


flowing, crystalline waters. Females were captured biting humans and horses.
Discussion: The closest species to S. deagostinii are S. dureti and S. pichi. Differences
from the first species were explained under that species. Comparison with S. pichi shows that
in S. pichi the female abdomen has a narrower median black stripe and tergite VI is totally
light gray, the calcipala is small, the trichomes of the thorax number about 60 on each side,
the tubercles are comparatively few and the lobes of the larval gill are without diverticula.

F
C E
m
3m

m
0.0

m
0.1
D

0.3
A

mm
K

0.3 mm
B

0.1 mm
I 0.02 m
m
G
0.1 mm

J H
N
L
Q
O
M
P

S
T R

0.0
U 5m
m

Figure 37. Simulium (Pternaspatha) deagostinii. A-K: Female; A: scutum; B: abdomen, dorsal view; C:
fronto-ocular triangle; D: maxillary palp; E-G: fore, mid and hind leg, respectively; H: apex of hind
basitarsus and tarsomere I; I: claw; J: eighth sternite and gonapophyses; K: cercus and paraproct. L-
N: Male; L: hind leg; M: gonocoxite and gonostylus; N: ventral plate. O-S: Pupa; O: general aspect;
P: cephalic sclerite extended; Q: portion of thorax with trichomes; R: trichomes and tubercles of
thorax; S: gill. T-V: Larva; T: cephalic apotome; U: head, ventral view; V: rectal papillae.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 185

Simulium (Pternaspatha) cotopaxi WYGODZINSKY & COSCARÓN (Fig. 38.)


Simulium (Pternaspatha) cotopaxi WYGODZINSKY & COSCARÓN, 1979: 19; COSCARÓN, 1991: 103;
COSCARÓN ARIAS, 2003: 198, 204, 208, 215, 219.
Female: Wing length 3.8-4.2 mm; head with eyes black; clypeus, frons and occiput
dark brown, with silver gray pruinosity; antennae and palpi black, their setae from dark
brown to black; scutum black, with variable color pattern according to incidence of light:
in some instances (Fig. A) almost entirely black, with 1+1 triangular anterosubmedian and
1+1 submedian spots on posterior third silver gray, and 1+1 sublateral and one median
longitudinal stripe of same color; with different illumination, scutum with 1+1 anterosub-
median triangular silver spots and 1+1 obliquely longitudinal silvery stripes beginning
somewhat before middle of scutum and connected posteriorly by transverse band (Fig.
B); with still different illumination central portion of scutum almost entirely silvery, with
1+1 anterior and 1+1 submedian spots appearing black; scutellum dark brown, narrowly
margined with lighter along hind border; metanotum velvety grayish brown; setae of
scutum and scutellum from brass to golden colored; pleura blackish; legs grayish brown,
with coxae, apices of femora and tibiae, apex of basitarsi and remaining tarsomeres,
darkened (Figs. H, I, J); setae of legs large and very numerous, producing overall light-
colored appearance of legs; abdomen black, with 1+1 silver gray spots on tergites II-VII,
most conspicuous on segments II and Vl, reduced on Vll (Fig. C); terminal tergites from
dull to slightly polished; ventral surface of abdomen blackish; frons without median
sulcus (Fig. D); frontal angle 125 degrees; fronto-ocular triangle as deep as wide at base
(Fig. E); shape of segments of antennae shown in figure F; setae not unusually elongate or
numerous; maxillary palp as shown in figure G; sensory vesicle of maxillary palp slightly
wider than half diameter of third segment; lacinia and mandibles serrate on both edges;
lacinia with approximately 28 teeth, mandible with 42-47 teeth; Sc with 12-15 hair; R1 with
hair and spiniform setae arranged in one or two irregular rows; shape of legs shown in
figures H-J; calcipala (Fig. K) slightly less than half as wide as basitarsus; pedisulcus
normal; claws with well-developed triangular process subbasally (Fig. L); eighth stemite
more darkened on median area (Fig. M); gonapophyses divergent, their inner margin
conspicuously sclerotized, their disc with very numerous microtrichia; cerci low; para-
procts narrowly and shortly projecting apically, their posterior border emarginated (Fig.
N); genital fork with stem slender, heavily pigmented; spermatheca subglobular, pigment-
ed, except circular area of insertion of spermathecal duct; inner surface of spermatheca
with faintly visible slender spicules.
Male: Wing length 3.6-3.8 mm; head black; eyes dark brown to black; clypeus black
with silvery pruinosity; antennae and palpi black, their hair also black; scutum entirely
black, with certain illumination anterior margin appearing gray with 1+1 short submedian
posteriorly directed gray projections, or with narrow silver gray band along margins of
entire sclerite; setae of scutum and scutellum colored as in female; color of metanotum,
pleura, wings and legs as in female; abdomen (Fig. O) sooty black, with 1+1 silver gray
spots each on segments II, VI, and VII, the one on II largest, the one on Vll smallest; head
strongly hairy (Fig. P); row of hair between eyes; scape and pedicel (Fig. Q) extraordinarily
enlarged, combined about as long as flagellum, and much wider than segments of latter;
186 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

setae of scape and pedicel unusually numerous and long, surpassing level of middle of
flagellum; maxillary palp with setae of third and to lesser degree also fourth palpomere
unusually long and numerous (Fig. R); fifth segment slightly over twice as long as
penultimate; sensory vesicle small; shape and pigment pattern of legs very close to those
of female but more darkened and hind leg with basitarsus strongly widened (Fig. S), about
three times as long as wide; calcipala small, one-fourth as wide as basitarsus apically (Fig.
T); genitalia with gonocoxite and gonostylus subtrapezoidal; gonostylus with distal margin
truncate and slightly concave and smooth; apical spine short, situated on slight projection
of inner apical angle (Fig. U); median sclerite with apical half widened and deeply incised;
endoparameres with numerous spinelike processes; membrane of aedeagus with numer-
ous minute cuticular spicules; ventral plate (Fig. V) wider than long.
Pupa: Cocoon (Fig. W) light brown, slipper shaped, translucent; fabric of cocoon
very uniform, parchment-like, individual threads difficult to perceive; anterior border of
cocoon reinforced, opening of cocoon subcircular; length of cocoon along dorsal surface
3.2-3.7 mm, along ventral surface 4.5-4.8 mm; body pupa 3.7-4.7, and gills 3.0-3.5 mm;
gills brown, darkest on basal half or third, each consisting of six slender filaments; all six
filaments forward directed, of subequal length, forming very tight bundle; head, thorax
and abdomen light brown, with frontoclypeus, antennal sheaths and exposed portion of
thorax dark brown, heavily sclerotized; tubercles of head and thorax delicately and
abundantly spinulose; shape of frontoclypeus of female as in figure Y, and male as in
figure Z; frontoclypeus with 1+1 facial and 2+2 frontal trichomes, from simple to bearing
up to eight branches (Fig. a); thorax (Fig. X) with 5-6+5-6 trichomes, also very small, with
up to eight branches (Fig. b); tergites III and IV with 4+4 simple hooks; apex of abdomen
(Fig. c) without pair of hooks or denticles.
Larva: Length 8.0-8.5 mm; width of head 0.8 mm; general body color light yellow
brown, tinged with greenish; head dark brown; general body shape as shown in figure d,
abdomen gradually widened posteriorly, not abruptly truncate at tip; cephalic apotome as
shown in figure e; spots positive, feeble, darkest along posterior margin; antennae as
shown in figure f, faintly pigmented, third segment darkest; ratio of proximal, medial and
distal segments = 1:1.4-1.7:0.8; cephalic fan with approximately 40 rays; toothing of
mandible as shown in figure g; maxillary palp as shown in figure h; hypostomium with
median and lateral teeth equally prominent (Fig. i); hypostomial setae (Fig. j) in two or
three irregular rows, each group consisting of 12-18 setae; postgenal cleft slightly deeper
than length of postgenal bridge (Fig. k); lateral sclerite of proleg shown in figure l, setae
arranged in three or four groups on prominent tubercles; body of larva glabrous, except
some setae near anal region and isolated ones scattered over rest of body; anal ring with
approximately 90 rows of 16 hooks each; anal sclerite as shown in figure m; rectal papillae
with 11-14 diverticula on each lobe (Fig. n).
Distribution: Ecuador: Cotopaxi, Pichincha, Tungurahua.
Bionomics: The mouthparts of the female of this species are normally developed;
therefore, females are presumed to be haematophagous. Larvae and pupae of Simulium
cotopaxi live in streams 2-5 m wide, with many small rapids and falls; the water temperature
was 11 ºC. Pupae were mostly found attached to the sides and undersurface of large
stones and boulders, as well as on blades of grass trailing in fast-flowing water.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 187

F
G

A B
I J
C
E H
D
K
L
M
N T
Q
O
P R S

W V

X Z
Y
U
f i
a k
b
j
e

c g
d

h
m l n

Figure 38. Simulium (Pternaspatha) cotopaxi. A-N: Female; A-B: scutum with different illumination; C:
abdomen, dorsal view; D: frons; E: fronto-ocular triangle; F: antenna; G: maxillary palp; H-J: fore, mid
and hind leg, respectively; K: apex of hind basitarsus and tarsite I; L: claw; M: eighth sternite and
gonapophyses; N: cercus and paraproct. O-V: Male; O: abdomen in lateral view; P: head; Q: antenna;
R: maxillary palp; S: hind leg; T: apex of hind basitarsus and tarsomere I; U: gonocoxite and
gonostylus; V: ventral plate. W-Z & a-c : Pupa; W: general aspect, lateral view; X: portion of exposed
portion of thorax and gill; Y: female frontoclypeus; Z: male, frontoclypeus; a: frontoclypeus trichome;
b: thoracic trichomes and tubercles; c: abdomen distal portion. d-n: Larva; d: lateral view; e: cephalic
apotome; f: antenna; g: teeth of mandible; h: maxillary palp; i: teeth of hypostomium; j: hypostomium;
k: hypostomium and postgenal cleft; l: proleg lateral sclerite; m: anal sclerite; n: rectal papillae.
188 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Discussion: Simulium cotopaxi is a peculiar species with a variety autopomorphic and


plesiomorphic characters that help separate it from other species, but constitute a
dilemma to place it in either one of the species groups of Pternaspatha. The gonostylus is
without tubercles (plesiomorphic), the gill branches and the enlarged hairs on the
antennae and maxillary palp of the male are characteristic of the NEMORALE group.
On the other hand ,the pupa with multiple filamentous branched trichomes, reduced
number of trichomes on the thorax (5-6 on each side) and female abdomen with 1+1 light
gray stripes on tergites II-VII, are more common in the NIGRISTRIGATUM group.
Some autopomorphies such as ornamentation of the female scutum, the ventral
projection of the paraproct, the elongate male pedicel, hair on the basal segments of the
antenna and maxillary palp, very wide hind basitarsus in the male, and the absence of
terminal hooks on the pupal abdomen, show a specialization probably present in a line of
Pternaspatha ancestors that evolved independently of the rest of the Pternaspatha.

NEMORALE species group

Simulium (Pternaspatha) nemorale EDWARDS (Fig. 39.)


Simulium (Simulium) nemorale EDWARDS, 1931: 152; ORFILA, 1939: 1533.
Simulium nemorale SMART, 1945: 509; VARGAS, 1945a: 168; WYGODZINSKY, 1949: 304; VARGAS & DIAZ,
1951: 138.
Simulium (Pternaspatha) nemorale WYGODZINSKY & COSCARÓN, 1967: 83-88; COSCARÓN & WYGODZIN-
SKY, 1972b: 222; COSCARÓN, 1991: 122-123.

Female: Wing length 3.2-4.0 mm; head black; eyes, antenna, maxillary palp, and
labium dark brown; occiput, frons, and clypeus gray pollinose; pilosity of antenna very
short, that of palp, clypeus, frons, and occiput somewhat longer; hair brass-colored to
black; first and second segments of antennae slightly lighter colored than remainder, and
with longer hair; scutum (Fig. A) black, with 1+1 silvery gray, submedian stripes arising
from 1+1 anterior white spots; median dark stripe divided longitudinally by delicate white
line; submedian stripes connected posteriorly to gray lateral borders of sclerite; appressed
setae of scutum brass-colored, dense, somewhat obliterating pattern; scutellum grayish
brown, spines and hair brass-colored to dark brown; metanotum brown to black, silvery
pollinose; pleural tuft brass-colored; wings hyaline, veins light brown, spines and hair
black; stem of halteres brownish, their knob yellowish; legs light brown with some regions
darkened as shown in hind leg figure J, and male legs (Figs. O, P, Q); their hair silvery to
black; abdomen (Fig. B) black; 1+1 large, silvery gray spots on sides of tergite II, and 1+1
smaller and more medially situated spots on tergite III; tergites IV and V entirely black
except a narrow line laterally on hind margin, very rarely with traces of paired very narrow
gray spots, as shown in figure C; tergite VI white except at sides and forward-pointing,
triangular, central, posterior, dark gray spot; tergites VI-IX black, with lateral light-colored
areas diminishing in size toward posterior segments; hair of abdomen, brass-colored,
rarely with silvery tinge; frons as shown in figure D; fronto-ocular triangle approximately
as long as wide (Fig. E); last segment of maxillary palp (Fig. F) more than twice as long as
penultimate segment, and slightly longer than two preceding segments combined;
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 189

A C
E
D
J K
B
F L
G
I
H
N
O
P M

Q R

S T X

U
W

Y
b
V

d
a c

Figure 39. Simulium (Pternaspatha) nemorale. A-N: Female; A: scutum; B-C: abdomen, of different
specimens dorsal view; D: frons; E: fronto-ocular triangle; F: maxillary palp; G: mandible; H:
lacinia; I: anterior veins of wing; J: hind-leg; K: apex of hind basitarsus and tarsomere I; L: claw; M:
eighth sternite and gonapophyses; N: cercus and paraproct. O-T: Male; O-Q: fore, mid and hind
leg, respectively; R: apex of hind basitarsus and tarsomere I; S: gonocoxite and gonostylus, dorsal
view; T: ventral plate, lateral view. U-Y: Pupa; U: lateral view; V: gill; W: frontoclypeus; X: exposed
portion of thorax with trichomes and tubercles; Y: abdomen chaetotaxy. Z, a- c: Larva; Z: cephalic
apotome; a: antenna; b: teeth of mandible; c: hypostomium.
190 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

diameter of sensory vesicle at least equal to half width of third segment; mandibles (Fig.
G) with about 40 teeth; lacinia (Fig. H) with 25-29; scutum with isolated long hair and
numerous short, appressed setae; Sc with few setae on its base, and from one to eight
setae on its central portion; basal portion of R glabrous; R1 with hair and spiniform setae
arranged in several rows; Rs with one or two irregular rows of setae (Fig. I); shapes and
proportions of hind segments of leg as shown in figure J; calcipala distinct, approximately
as long as wide at its base (Fig. K); claws with subbasal tooth well-developed (Fig. L);
eighth sternite with about 25 long hairs on each side (Fig. M); hind border distinctly
pigmented; gonapophyses subtriangular, beset with microchaetae, their inner border
slightly sclerotized; paraproct and cercus as shown in figure N; genital fork with median
process distinctly sclerotized except its apex which is widened; anterior projections
narrow, with wide, medially directed expansions.
Male: Wing length 3.8 mm; head blackish; eyes, occiput, antenna, and maxillary palp
dark brown, almost black; clypeus black, gray pollinose; occiput, clypeus, and basal
segments of maxillary palp with numerous long hair, latter also in row between eyes;
scutum black, with appressed setae that appear to be golden to black; scutellum light
brown, with brown to black hair; metanotum dark brown, golden pollinose; pleura dark
brown, silvery pollinose; legs brown, with hair golden to black; dark regions distributed as
in figures O, P, Q; abdomen velvety black, its hair golden to black; tergites II and VI with
1+1 silvery gray spots, those of second segment larger and more brilliant, those of sixth
segment darker, with bluish tinge; last segment of maxillary palp more than twice as long
as penultimate; diameter of sensory vesicle smaller than half of diameter of third
segment; wings like those of female, but center of Sc only occasionally with setae, and
hair and spiniform setae on R1 slightly less numerous; hind basitarsus 3.8 times as long as
wide; calcipala well developed, about as long as wide at base (Fig. R); genitalia as shown in
figures S, T; gonocoxite subtrapezoidal, slightly wider than long; gonostylus subquadrate,
distal portion widely concave, smooth (Fig. S); central portion of ventral plate salient
posteriorly at middle, with numerous curved setae; lateral projections and basal margin
distinctly sclerotized (Fig. T); disc of ventral plate with 1+1 translucent areas; en-
doparameres membranous, with strongly pigmented spines.
Pupa: Cocoon shoe shaped (Fig. U), with large, anteroventral bridge; color of
cocoon light brown; surface smooth, closely woven; threads not individualized; rim of
aperture slightly reinforced; gills protected laterally by walls of cocoon, only some of
dorsal filaments free; length of cocoon at dorsum along middle, 4.2 mm; maximum length
along base, 6.0 mm; length of body of pupa, 3.8 mm; length of gill 1.9 mm; gills (Fig. V)
each with eight almost parallel filaments, slightly converging toward their apices; head and
thorax of pupa light brown; head with 3+3 simple, short, frontal, 1+1 long, mostly
branched facial, and 1+1 simple long, ocular trichomes (Fig. W); thoracic region adjacent
to aperture of cocoon with band of tubercles and numerous interspersed, very short and
stout trichomes (Fig. X); several smaller tubercles posteriorly on covered portion of
thorax; disc of thorax on each side with approximately 20 additional trichomes; their size
and exact distribution individually variable, in some cases bifid, in latter case generally
longer than others; toward ventral surface, along band of tubercles, additional trichomes,
similar to those described above, but more delicate; total number of thoracic trichomes,
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 191

approximately 60+60; abdominal tergites light brown, intensity of pigment diminishing


toward posterior segments; chaetotaxy as shown in figure Y.
Larva: Maximum length, 6.0 mm; width of head capsule, 0.37 mm; color of larva
whitish; cephalic apotome light-colored, darker along middle and at hind margin (Fig. Z);
cuticle with isolated short hair, more numerous toward posterior end; antennae as shown in
figure a, light brown, distal segment darkest; medial segment with two slight constrictions,
regions of constrictions with unpigmented areas; ratio of proximal, medial and distal
segments = 1:1.2:0.8; cephalic fan with 37-40 rays; toothing of mandibles as shown in figure
b; two outer, one strong apical, three approximately identical subapical, five internal and two
marginal teeth, first of which much larger than second; anterior border of hypostomium
strongly pigmented (Fig. c); median tooth less prominent than lateral ones; intermediate
teeth gradually increasing in size from II to IV; sublateral tooth barely perceptible; lateral
borders rugose, with four to five very small serrations; hypostomial setae arranged in two
irregular rows, about 14 in each group; disc of hypostomium glabrous; postgenal cleft (Fig.
d) deep, pointed, its depth much larger than postgenal bridge; anal sclerite base with simple
to trifid scales in large numbers; anal ring with about 85 rows, with approximately 15 hooks
each; rectal papillae with three simple lobes.
Bionomics: The pupae were found attached to roots of the bamboo Chusquea culeou
DESVAUX (common name, “caña colihue”) trailing in the water, at about 10 cm under the
surface. They live in torrential creeks at the base of the Andean mountains.
Distribution: Argentina: Chubut, Neuquen, Río Negro, Tierra del Fuego; Chile: Aysen.
Discussion: This species is very similar to S. barbatipes in the female abdominal
ornamentation, but the 1+1 lighter spots of tergite III are smaller in S. nemorale, and in S.
barbatipes the pupa has 6 branches and the thoracic trichomes number about 120.

Simulium (Pternaspatha) barbatipes (ENDERLEIN) (Fig. 40.)


Acropogon barbatipes ENDERLEIN, 1934b: 277; STONE, 1962: 206.
Simulium barbatipes: SMART, 1945: 501; VARGAS, 1945a: 119; VARGAS & DIAZ; 1953b: 141. WYGODZIN-
SKY, 1953b: 321; 1958: 133; STONE, 1962: 206; 1963: 17.
[Simulium (Pternaspatha)] barbatipes STONE, 1963: 1.
Simulium (Pternaspatha) barbatipes WYGODZINSKY & COSCARÓN, 1967: 99; COSCARÓN & WYGODZINSKY,
1972b: 220; COSCARÓN, 1991: 136-137.
Female: Wing length 3.1-3.9 mm; head black; occiput, frons, and clypeus gray
pollinose; palpi and antennae black, latter with two basal segments more or less
conspicuously orange-colored; scutum as usual in group; medium, narrow, longitudinal,
white line distinct; appressed setae of scutum very pale brass-colored; scutellum and
metanotum piceous, former with long, pale, brass-colored or dark hair; pleura gray
pollinose; pleural tuft pale brass-colored; base of halteres dark, remainder yellowish white;
legs light brown, their hair silvery to black; relative intensity of pigmented areas as shown
in figures F, G, H; abdomen dark gray, tergites II-V deep black at center; tergite II with
1+1 large, silvery white spots; tergites III-V with 1+1 small spots, those on III largest,
those of IV and V very small, in some cases difficult to perceive; tergite VI extensively
silver-gray, black at lateral extremities, in some cases very faintly darkened at center; tergite
192 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

VII dark, light gray at sides; tergites VIII and IX entirely dark; hair of abdomen, including
that of basal fringe, silver-colored (Fig. A); frons as shown in figure B, with obsolete
median sulcus; fronto-ocular triangle (Fig. C) slightly wider than long; last segment of
maxillary palp (Fig. D) more than twice as long as penultimate segment; diameter of
sensory vesicle approximately half that of third segment of maxillary palp; structure of
vesicle as shown in figure E; maxillae with about 25 teeth, mandible with 40-47, teeth; Sc
with few setae on central portion; setae and spines on R1 in single row, setae on R in single
series; legs as in figures F, G, H; calcipala small, about as long as wide at base (Fig. I); claws
with small but distinct tooth Fig. J); eighth sternite as in figure K; its lateral portions more
heavily pigmented than central area, latter with posterior dark band; gonapophyses
subtriangular, membranous, beset with microchaetae, inner border faintly sclerotized;
paraprocts and cerci as shown in figure L; cerci much higher than wide, their distal border
rounded; protruding portion of paraprocts subtriangular, their apex rounded; median
process of genital fork strongly sclerotized, its apex widened, light-colored; anterior
projections slightly darker than arms of fork; spermatheca globular, its inner surface with
minute spicules arranged in groups.
Male: Wing length 3.4-4.0 mm; head blackish; eyes, antennae, and palpi dark brown,
almost black, first two and extreme base of third antennal segments lighter, all with silvery
pilosity; palpi, basal two antennal segments, clypeus, line between eyes, and occiput all with
long black hair; scutum as usual for group; appressed setae brass-colored; scutellum brown,
with brass-colored hair; metanotum and pleura dark brown, silvery pollinose; pleural tuft
brass-colored; legs whitish to brown, distribution and intensity of pigment as shown in
figures M, N, O; abdomen velvety black; its hair, including that of basal fringe, silvery;
segment II with 1+1 large, silvery white spots, segments VI and VII with 1+1 smaller ones
(Fig. P); last segment of maxillary palp more than twice as long as penultimate; diameter of
sensory vesicle approximately half that of third palpomere; wings like those of female, but
Sc devoid of setae; hind basitarsus 3.4-3.9 times as long as wide; calcipala slightly shorter
than wide at base (Fig. Q); genitalia very similar to those of Simulium nemorale.
Pupa: Cocoon slipper shaped (Fig. R), with very narrow anteroventral bridge; rarely
shoe shaped; color of cocoon brown; surface smooth, closely woven, threads not
distinctly individualized; rim of aperture slightly reinforced; only extreme base of gills
protected by lateral walls of cocoon, all filaments free; length of cocoon at dorsum, 3.0-
3.6 mm; maximum length along base, 4.2-5.1 mm; length of pupa, 3.5-4.0 mm; length of
gill, approximately 2.0 mm; gills (Fig. S) each consisting of six closely parallel filaments;
three primary branches arising from short basal trunk, each branch dividing into two
filaments not far from its base; exact level of branching somewhat variable; head and
thorax of pupa dark brown; head with 3+3 branched frontal, 3+3 branched facial, and
1+1 branched ocular, trichomes (Fig. T); thorax showing zone adjacent to aperture of
cocoon with numerous tubercles of various sizes and numerous trichomes of different
structure; short, simple or bifurcate, spatulate, flattened, in some cases curved, in others
hair like, with from one to six long and delicate filaments (Figs. U, V); total number of
trichomes, 110-130+110-130; chaetotaxy of abdomen like that of S. nemorale.
Larva: Maximum length, 8.0 mm; maximum width of head capsule, 0.7 mm; larva
with extensive hypodermal pigment on dorsal and on anterior half of ventral surface of
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 193

B D F
J
E

A C
G

H
K

L I
M
R

P
N Q
T

W
Z
U
S
a
X

V
c Y
b

Figure 40. Simulium (Pternaspatha) barbatipes. A-L: Female; A: abdomen, dorsal view; B: frons; C:
fronto-ocular triangle; D: maxillary palp; E: sensory vesicle of maxillary palp; F-H: fore, mid and
hind leg, respectively; I: apex of hind basitarsus and tarsomere I; J: claw; K: eighth sternite and
gonapophyses; L: cercus and paraproct. M-Q: Male; M-O: fore, mid and hind leg, respectively; P:
abdomen, dorsal view; Q: apex of hind basitarsus and tarsomere I. R-V: Pupa; R: lateral view; S: gill;
T: frontoclypeus; U: exposed portion of thorax and trichomes; V: trichomes and tubercles of
thorax. W-Z & a-c: Larva; W: cephalic apotome; X: antenna; Y: teeth of mandible; Z: hypostomium
a: cephalic capsule in ventral view; b: anal sclerite; c: rectal papillae.
194 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

body; cephalic apotome yellowish, with dark pattern as shown in figure W; body
integument glabrous except some short setae at base of anal sclerite; antennae dark
brown, distal segment darkest; ratio of proximal, medial and distal segments = 1:1.2:0.72
(Fig. X); cephalic fan with approximately 40 rays; toothing of mandible as shown in figure
Y; hypostomium with median tooth a little more prominent than lateral ones; lateral
borders with four or five small serrations; hypostomial setae arranged in two irregular
series, their number about 10 in each group (Fig. Z); postgenal cleft (Fig. a) triangular, its
depth larger than length of hypostomial bridge; anal sclerite as shown in figure b, its base
with numerous small scales; anal ring with about 80 rows, each composed of 13-14 hooks;
rectal papillae (Fig. c) consisting of three primary lobes, middle one with six, lateral ones
with two diverticula each.
Distribution: Argentina: Jujuy, Mendoza, Neuquén, Salta, Tucuman; Chile: Acon-
cagua, O’Higgins, Santiago; Bolivia: La Paz; Peru: Chocó, Cuzco, Junin, Puno.
Bionomics: Simulium barbatipes breeds in torrential high-mountain streams of
relatively low temperatures (10-12 °C). It is also found in irrigation ditches with fast-
flowing water. The larvae and pupae are attached to vegetation trailing in the current or to
the surface of rocks.
Discussion: Simulium barbatipes is one of the most wide-ranging species of the
Pternaspatha group. Although there is some individual variation, no characters were found
that would separate the various populations that were examined.

Simulium (Pternaspatha) philippii COSCARÓN (Fig. 41.)


Simulium (Pternaspatha) philippii COSCARÓN, 1976a: 29-33; 1991: 125-126.
Female: Wing length 2.9-3.4 mm; head blackish, antenna, palp and proboscis dark
brownish; scutum as in Simulium nemorale; legs light brown with apex of segments and
most of tarsi dark brown; abdomen grayish brown, blackish on median area of tergites,
with 1+1 whitish gray submedian spots on tergites III-V, larger on tergite II and extending
sublaterally on tergites VI-VIII (Fig. A); fronto-ocular triangle deep (Fig. B); lacinia with
about 20 teeth and mandible with 30-34 teeth; maxillary palp as in figure C, with enlarged
sensory vesicle with abundant sensilla (Fig. D); short calcipala (Fig. E); claw without
subbasal tooth (Fig. F); eighth sternite and gonapophyses similar to those of Simulium
stelliferum; paraprocts relatively elongated, not acuminate distally (Fig. G).
Male: Wing length 3.1 mm; antenna and palp coloration as in female; thorax and legs
with golden to grayish brown pilosity; abdomen with 1+1 silvery spots on tergites II, VI
and VII, the two latter smaller; calcipala small (Fig. H); ratio of length/width of hind
basitarsus = 3.5; gonostylus without tubercles (Fig. I).
Pupa: Cocoon length on dorsum 2.5-2.8 mm; base 3.0-3.2 mm; gills 2.4-2.8 mm;
pupa body 3.0 mm; cocoon slipper shaped, scarcely projected ventrally and with
reinforced border (Fig. J); gill with 6 branches disposed as shown in figures L, M; head and
thorax with abundant tubercles; on frontoclypeus with about 1200, those of thorax bigger
than on head; frontoclypeus with 2+2 frontals and 1+1 facial trichomes relatively short,
uni- or biramous (Fig. K); thorax with 60-85 trichomes on each side, filiform, with 1-3
branches. (Figs. M, N); abdominal chaetotaxy as in Simulium nemorale.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 195

Larva: Length 7.5 mm; color light grayish brown-green; head dark brown; cephalic
apotome more darkened on base (Fig. O), central spots negative; antenna with distal
segment more darkened (Fig. P); ratio of proximal, medial and distal segments = 1:1.2-
1.3:0.8; cephalic fan with 42-52 rays; toothing of mandible as in figures Q, R;
hypostomium with apical border well pigmented, with median teeth as high as lateral
teeth; hypostomial setae 7-11 in number on each side and postgenal bridge as long as
hypostomium length (Fig. S); anal sclerite without scales; anal ring with 83-93 rows with
13-15 hooks in each; rectal papillae with 3 single lobes (Fig. U); comb of thoracic proleg
sclerite with about 17 groups of setae of different length (Fig. T).
Distribution: Chile: O’Higgins, Tarapaca.
Bionomics: Simulium philippii breeds in creeks with high current and cold waters
(about 10 °C) in the southern and temperate areas of the northern region. The feeding
habits of the females are unknown.

E
C H
B
D
F
L

A G
I M

N
K

S P
Q

O
U
R T
Figure 41. Simulium (Pternaspatha) philippii. A-G: Female; A: abdomen, dorsal view B: fronto-ocular
triangle; C: maxillary palp; D: sensory vesicle of maxillary palp; E: apical portion of hind basitarsus
and tarsomere I; F: claw; G: cercus and paraproct. H-I: Male; H: apical portion of hind basitarsus and
tarsomere I; I: gonostylus. J-N: Pupa; J: lateral view; K: trichomes and tubercles of frontoclypeus; L:
basal portion of gill; M: exposed portion of thorax with trichomes and gill; N: trichomes and
tubercles of thorax. O-U: Larva; O: cephalic apotome; P: antenna; Q-R: teeth of mandible in different
specimens; S: hypostomium and postgenal cleft; T: lateral sclerite of proleg; U: rectal papillae.
196 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Discussion: On the basis of the abdominal ornamentation, the closest species to S.


philippii is S. nigristrigatum, but in S. nigristrigatum the light spots on tergites III-V are larger,
the calcipala is absent, the subbasal claw teeth are well developed, the pupal gill has 8
filaments, and the thoracic trichomes are multiramous.

Simulium (Pternaspatha) herreri WYGODZINSKY & COSCARÓN (Figs. 42 A-Z, a-f.)


Simulium (Pternaspatha) herreri WYGODZINSKY & COSCARÓN, 1967: 104-109; COSCARÓN, 1991: 130.
Female: Wing length 2.8-3.5 mm; head black; eyes piceous; antennae, palpi, and
labium black, first two antennal segments tinged with gray; occiput, frons, and clypeus
gray pollinose; pilosity of antennae very short; basal segments of palpi, clypeus, frons, and
occiput with some long dark hair; scutum as usual for group; dark regions velvety black;
appressed setae of scutum brass-colored, not very dense, not obliterating pattern;
scutellum grayish brown, with long hair brass-colored to brown; metanotum, pleura, and
sterna piceous, silvery pollinose; legs light brown, pigment areas as shown in figures F, G,
H; abdomen (Fig. A) black; tergite II with 1+1 large, silvery white spots, 1+1 smaller ones
on tergite III, 1+1 very small ones each on tergites IV and V; tergite VI with transverse,
silvery white, posteriorly emarginated band on central two-fourths; tergites VII and VIII
grayish or silvery white on sides; tergites II-VI narrowly banded with grayish on lateral
portions of hind margin; hair of fringe of first segment yellowish white, that of rest of
abdomen brass-colored to very dark; frons as shown in figure B; fronto-ocular triangle
about as long as wide (Fig. C); last segment of maxillary palp (Fig. D) slightly more than
twice as long as penultimate segment, and as long as third and fourth segments combined;
diameter of sensory vesicle about equal to half diameter of third palpomere, its surface
structure as shown in figure E; lacinia with 26 or 27 teeth, mandible with 36-40, teeth;
scutum with appressed setae on disc; posterior declivity with erect or forwardly curved
longer hair; Sc with three or four hairs on central portion; R1 with hair and spiniform setae
arranged in one row, in some cases becoming double on apical portion of vein; leg as
shown in figures F-H; calcipala (Fig. I) small but distinct, about as long as wide; claws with
well-developed tooth (Fig. J); eighth sternite pigmented, central posterior portion as in
figure K, gonapophyses subtriangular, rounded apically, their inner border slightly
sclerotized, their disc with very numerous microtrichia; paraprocts and cerci as shown in
figure L; projecting portion of paraprocts semi elliptical; cerci with distal border truncate.
Male: Wing length 3.0-3.2 mm; head black; eyes purple; occiput, antennae, and palps
black, clypeus black, gray pollinose; occiput, clypeus, and basal maxillary segments with
numerous very long hair, latter also arranged in row between eyes; scutum as usual for
group, deep velvety black, on its anterior three-fourths a very delicate, median, longitudi-
nal line; hair brass-colored; scutellum black, its hair brass-colored; metanotum, pleura,
sterna, as in female; legs yellowish brown, their hair brass-colored to black; relative
intensity of pigment areas as shown in figures M, N, O; abdomen (Fig. M) velvety black,
hair silvery gray; segment II with 1+1 very large, silvery white spots closely approaching
but not meeting dorsally; segment V in some individuals very faintly iridescent, at certain
angle showing small, silvery white area dorsally at center; VI entirely silvery white except
small dark area dorsally at center posteriorly, VII with 1+1 medium-sized, silvery white
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 197

D F
C
B G
E
A
N
K J
H

O
M
I

L
P
† R
T
W

Q S
V

c U
a
b X
m
5m
0.0

Z e
d f

Figure 42. Simulium (Pternaspatha) herreri. A-L: Female; A: abdomen, dorsal view B: frons; C: fronto-
ocular triangle; D: maxillary palp; E: sensory vesicle of maxillary palp; F-H: fore, mid and hind leg,
respectively; I: apical portion of hind basitarsus and tarsomere I; J: claw; K: portion of eighth sternite
and gonapophyses; L: cercus and paraproct. M-S: Male; M: abdomen in lateral view; N: antenna; O:
maxillary palp; P: anterior veins of wing; Q: hind leg; R: apical portion of hind basitarsus and tarsite
I; S: gonostylus. T-Y: Pupa; T: lateral view; U: gill; V: frontoclypeus; W: frontoclypeus tubercles with
high magnification; X: exposed portion of thorax and trichomes; Y: thoracic trichomes with high
magnification. Z, a-f: Larva; Z & a: cephalic apotome in different specimens; b: antenna; c: teeth of
mandible; d: hypostomium; e: hypostomium and postgenal cleft; f: rectal papillae.
198 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

spots widely separated above; IX faintly iridescent; eyes separated by series of long hairs;
scape and pedicel with long hair (Fig. N) as also basal segments of maxillary palp (Fig. O);
diameter of sensory vesicle of palp half that of third palpomere; wings without setae on
central portion of R1 with one row of setae intermixed with hair (Fig. P); shapes and
proportion of segments of hind legs as shown in figure Q; hind basitarsus 3.2 times as
long as wide; calcipala (Fig. R) small but distinct, slightly wider than long; legs with
numerous long setae; gonocoxite wider than long; gonostylus subquadrate, distal portion
widely concave, not tuberculate (Fig. S); ventral plate with 1+1 distinct, translucent areas.
Pupa: Cocoon (Fig. T) slipper shaped, with short anteroventral bridge; color light
brown, surface smooth, closely woven, threads not individualized under medium magnifica-
tion; rim of aperture not or only feebly reinforced; length of cocoon at dorsum, 2.5-3.0 mm;
maximum length along base, 4.0 mm; average length of pupa, 3.5 mm; length of gills 2.5-3.0
mm; gills (Fig. U) each with six filaments, dorsal and median primary branches and their
filaments forming tight bundle, ventral primary branch and its filaments sharply curved
ventrally and backward, filaments almost entirely hidden by cocoon; head and thorax of
pupa light brown; head with 1+1 fields of tubercles at base of clypeus (Figs. V, W), 2+2
frontal and 1+1 facial trichomes, all simple, facial trichomes much longer than frontal
trichomes; thoracic region adjacent to aperture of cocoon with band of tubercles that
extends to midline ventrally and attains base of gill; most of remainder of exposed surface
of thorax covered with trichomes in shape of elongate, lanceolate setae (Figs. X, Y), their
length, 0.07-0.10 mm, their total number approximately 250+250; few fine hair near
insertion of gills; chaetotaxy of abdomen on tergite I with 2+2 lateral setae; tergite II with
3+3 relatively small, equidistant spines (acrostichal, dorsal and dorsocentral), and in some
cases also dorsolateral one, one or two anterior dorsocentral and approximately 3+3 lateral
setae also present; tergites III and IV with 4+4 large, spinelike setae, dorsolateral separated
from dorsocentral by relatively wide space; tergites III and IV also with 1+1 anterior
dorsocentral and 2+2 lateral setae; tergite V with 1+1 small, dorsal and 1+1 dorsolateral
setae, as well as with 1+1 small, anterolateral groups of small, scale like structures; tergites
VI-VIII at anterior border with 1+1 rows of spiniform, simple, bifid, or trifid tubercles, last
transformed laterally into small, scale like structures arranged in several irregular rows and
occasionally with 1+1 dorsocentral setae; rows of tubercles on tergites VI and VII widely
separated from one another, those of VIII closely approximated, almost confluent; numbers
of tubercles in rows 6-13+6-13 on tergites VI and VII, 10-13+10-13 on VIII; anterior
border of tergite IX in some cases with 1+1 groups of one to three tubercles; sternite III
with 1+1 lateral and 1+1 ventral setae, central region with transverse area of numerous small
and very delicate, scale like structures; sternite IV with 3+3 or 4+4 ventral and lateral setae,
and central area of scale like structures as on III; sternites V-VII with 2+2 spines before
posterior margin; spines of each pair close to one another on fifth, separated by wider space
on both sixth and seventh segments; spines of V generally bifid, but in some cases
ventrolateral one simple; on VI and VII, generally ventral spine bifid and ventrolateral one
simple, but in some both simple; sternites V-VII each with 1+1 suboval fields of scale like
structures; VIII with one transverse group of scales near anterior margin.
Larva: Maximum length, 7.0 mm; width of head capsule, 0.60-0.65 mm; color
variable; dark hypodermal pigment generally present on dorsal and ventral of anterior
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 199

half of body and on dorsal surface of posterior half; cephalic apotome with two different
basic color patterns, as illustrated in figures Z, a; cuticle with isolated short hair, becoming
more numerous toward posterior end; antennae as shown in figure b; ratio of proximal,
medial and distal segments = 1:1.1-1.25:1.0-1.08; cephalic fan with approximately 40 rays;
toothing of mandibles as shown in figure c; hypostomium with median tooth as
prominent as lateral ones; central intermediate tooth smallest; lateral serration well
developed, irregular in shape; hypostomial setae arranged in two irregular rows, with
average of 11 setae in each group (Fig. d); postgenal cleft rather wide (Fig. e), its depth not
much larger than length of postgenal bridge; anal sclerite with scales at its base small and
few in number; anal ring with approximately 90 rows, of about 15 hooks each; central lobe
of rectal papillae with 1+1, lateral lobes each with one secondary diverticula (Fig. f).
Distribution: Peru: Lima.
Bionomics: The larvae and pupae S. herreri live at 2300-2500 m altitude on trailing grass
leaves and rocks in a small stream and irrigation ditches, at 1-3 cm below the water surface.
Pupae frequently form small aggregations. The temperature in the watercourses was 15 °C.
Discussion: Simulium herreri, S. albilineatum, and S. yacuchuspi are very similar. The
male of S. herreri differs from males of the two other species by the absence of paired
light-colored spots on tergite III. The female of S. herreri is distinguished by the shape of
the paraprocts and the light-colored central portion of the eighth sternite. The pupa is
characterized by the sharply divergent filaments of the ventral primary branch of the gill,
the conspicuous areas of tubercles at the base of the frontoclypeus, and the lanceolate
trichomes of the thorax. The larvae of S. herreri have the postgenal bridge about as long as
the postgenal cleft and in S. albilineatum and S. yacuchuspi it is much shorter, about a half as
long as the postgenal cleft.

Simulium (Pternaspatha) yacuchuspi WYGODZINSKY & COSCARÓN (Fig. 43.)


Simulium (Pternaspatha) yacuchuspi WYGODZINSKY & COSCARÓN, 1967: 110-115; COSCARÓN, 1991: 127-128.
Female: Wing length 3.5-3.8 mm; pilosity and color of head and thorax as in
Simulium herreri; legs light brown, darkened on apices of femur of tibia and tarsi; their hair
silvery to black; abdomen (Fig. A) black; tergite II with 1+1 large, silvery white spots,
tergite III with 1+1 smaller ones, tergites IV and V each with 1+1 very small ones; tergite
VI entirely silvery white, very faintly darkened posteriorly at center; VII black on central
half, white on lateral fourths; tergite VIII black, silvery white only at lateral extremities;
tergite IX grayish black; tergites IV and V narrowly banded with grayish on lateral portion
of hind border; hair of basal fringe and of remainder of abdomen silvery; frons as shown
in figure B; fronto-ocular triangle about as long as wide; maxillary palp like that of
Simulium herreri, but tubercles of sensory vesicle (Fig. C) relatively larger and somewhat
less numerous; lacinia with 22-27 teeth, mandible with 42-48, teeth; shape of hind leg as in
figure D; calcipala distinct (Fig. E), about as long as wide; claws with very small tooth (Fig.
F); eighth sternite and gonapophyses like those of Simulium albilineatum; paraprocts and
cerci as shown in figure G; projecting portion of paraprocts rather narrowly subtriangular.
Male: Wing length 3.0-3.5 mm; color of head and its appendages like that of
Simulium herreri; color of thorax like that of S. herreri, but knob of halteres yellow; legs pale
200 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

yellow-brown with darkened area as in female but more extended on hind basitarsus, their
hair silvery white to black; abdomen (Fig. H) velvety black; hair, including that of basal
fringe, generally silvery, rarely brass-colored; segment II with 1+1 very large, silvery white
spots closely approaching but not meeting dorsally; segment III with 1+1 small, silvery
white spots laterally at hind margin; segment VI almost entirely silvery white except small
dark area at center behind; VII with 1+1 medium-sized, lateral, silvery white spots;
segment IX faintly iridescent; shape and proportions of segments of hind legs as in figure
I; hind basitarsus 2.7-3.0 times as long as wide; calcipala (Fig. J) very small but distinct,
about as long as wide; genitalia much like those of Simulium herreri.
Pupa: Cocoon slipper shaped (Fig. K) with well-developed anteroventral bridge;
color light brown, translucent, surface smooth, texture closely woven, threads not
perceptible under low magnification; rim of aperture feebly reinforced; length of cocoon
at dorsum along middle, 2.8-3.4 mm; maximum length along base, 4.1 mm; length of body
of pupa 3.2-3.4 mm; length of gills 2.2-2.5 mm, viz., half as long as cocoon, and two-
thirds as long as body of pupa; gills (Figs. L, M) each with six filaments arising from three
primary branches; dorsal and median primary branches and their filaments lying closely
together, ventral primary branch sharply ventrally bent, its ventral filament curved
backward under cocoon, its dorsal filament generally curved forward, but not closely
associated with remaining filaments; all primary branches with division into filaments not
far removed from base of gill; division of dorsal primary branch closest to, and either that
of median or ventral branch most remote from, base of gill; dorsal primary branch
thickest, ventral primary branch thinnest; dorsal filament of dorsal primary branch
conspicuously thicker on basal portion than any other; head and thorax of pupa dark
brown; head (Fig. N) with (1-)2+2(-1) frontal and 1+1 facial trichomes, latter frequently
longer, and generally bifid; thoracic region adjacent to aperture of cocoon with band of
not very numerous tubercles, becoming evanescent toward sides of thorax; smaller
tubercles also occupying large area of dorsal portion of part of thorax covered by
cocoon; exposed surface of thorax with large and very variable number of simple (rarely
bifid) trichomes, longest and most delicate near band of tubercles on lateral portion of
thorax, diminishing in size toward dorsum and lateral disc of thorax (Figs. O, P);
trichomes mostly slender, hair like (Fig. Q), but in some specimens trichomes on disc
becoming very short and pointed, almost spinelike; length of trichomes varying from 0.02
to 0.16 mm., most frequently around 0.1 mm; total number of trichomes varying from
50+50 to 130+130; type and number of trichomes invariably closely similar on both
halves of thorax in one individual; chaetotaxy of abdomen like that of Simulium herreri, but
spiniform tubercles on tergite VIII generally forming continuous row.
Larva: Maximum length, 7.0 mm; width of head capsule, 0.65-0.70 mm; larvae color
brownish green (in alcohol) on whole body except posterior fifth of ventral surface; larvae
appearing black in life; head very variable in color, extremes of pattern of cephalic
apotome shown in figures R, S.; dark form prevalent; cuticle with few scattered short hair;
antennae as shown in figure T, ratio of proximal, medial and distal segments = 1:1.1-
1.3:0.75-0.9; cephalic fan with 40-46 rays in large fan; toothing of mandibles (Fig. U)
much like that of Simulium herreri; structure and pigmentation of hypostomium as shown
in figure V; lateral serrations well developed, irregular in shape; hypostomial setae
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 201

D
B

F
C
E

A G
H K
O

J
N
0.05
mm

L
M

Q
P
T R

U
X S

W V

Figure 43. Simulium (Pternaspatha) yacuchuspi. A-G: Female; A: abdomen, dorsal view B: frons; C:
sensory vesicle of maxillary palp; D: hind leg; E: apical portion of hind basitarsus and tarsomere I;
F: claw, in different position; G: cercus and paraproct. H-J: Male; H: abdomen in lateral view; I:
hind leg; J: apical portion of hind basitarsus and tarsomere I. K-Q: Pupa; K: lateral view; L-M: gill;
N: frontoclypeus; O-P: exposed portion of thorax with different number of trichomes; Q: thoracic
trichomes with high magnification. R-X: Larva; R-S: cephalic apotome in different specimens; T:
antenna; U: teeth of mandible; V: portion of hypostomium; W: hypostomium and postgenal cleft;
X: rectal papillae.
202 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

arranged in two irregular rows, with average of 15 setae in each group; region of insertion
of setae distinctly less pigmented than remainder of disc of hypostomium; postgenal cleft
relatively narrow, pointed apically, its depth larger than length of postgenal bridge (Fig.
W); anal sclerite with small scales at its base and few in number; anal ring with 81-89 rows,
each composed of 12-20 hooks; central and lateral lobes of rectal papillae each with 16-20
very long and slender secondary lobules arranged in two to three irregular rows (Fig. X).
Distribution: Peru: Junin.
Bionomics: The larvae and pupae of S. yacuchuspi were found at 3750 m altitude,
attached to leaves and stems of grasses trailing in a small stream, near the surface; the
stream was not more than 50 cm wide and not more than 10 cm deep, and the
temperature of the water was 10 °C.
Discussion: Simulium yacuchuspi shows great variation in the number and size of the
trichomes of the pupal thorax. This species is closely related to S. herreri, S. albilineatum,
and S. albicinctum. The larva differs from that of S. herreri and S. albilineatum (that of S.
albicinctum is unknown) mainly by the large number of diverticula of the rectal papillae.
The pupa of S. yacuchuspi differs from that of S. herreri (with which it shares the backward-
directed ventral filament of the ventral primary branch of the gill) by the thicker
lanceolate-shaped thoracic trichomes in S. herreri, and from that of S. albilineatum by the
virtual absence of bifid trichomes on the thorax and the presence of the above-
mentioned backward-directed ventral filament.
The adults are more difficult to distinguish with certainty. The male of S. yacuchuspi
differs from that of S. albicinctum by the presence of 1+1 white spots on abdominal tergite
III, and a less strongly widened hind basitarsus. It differs from that of S. herreri by the
presence of the spots and the absence of such spots from segment V, but it cannot be
separated from the male of S. albilineatum. The female of S. yacuchuspi can be recognized by
the structure of its paraprocts. It differs from that of S. albilineatum by the white spots on
tergite V which are much smaller than those of tergite III, and from that of S. herreri also by
the dark central portion of the eighth sternite, the poorly developed tooth of the claw, and
the almost completely white tergite VI. The female of S. yacuchuspi is rather similar to that of
S. albicinctum, but the latter lacks white spots on tergites IV and V, tergite VI is entirely white,
the dark area at the base of the posterior tibia is more extended, and the calcipala is wider.

Simulium (Pternaspatha) albilineatum (ENDERLEIN) (Fig. 44.)


Pternaspatha albilineata ENDERLEIN, 1936: 115.
Simulium albilineata SMART, 1945: 500.
Simulium albilineatum VARGAS, 1945a: 109.
Simulium (Pternaspatha) albilineatum WYGODZINSKY & COSCARÓN, 1967: 95-99; COSCARÓN, 1991: 126-127.
Female: Wing length, 3.5-3.7 mm; color of head and thorax like that of Simulium
herreri; legs yellowish brown, their hair from silvery to black; color pattern, illustrated in
figures C, D, E; abdomen (Fig. A) black; tergite II with 1+1 large, silvery white spot; tergites
II and V with 1+1 smaller ones; tergite IV either entirely black or with 1+1 minute whitish
spots; tergite VI silvery white darkened only at lateral extremities; tergite VII dark at middle,
silvery white on lateral thirds, dark region widened posteriorly; tergite VIII black, silvery only
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 203

at lateral extremities; tergite IX silvery to black; tergites II and III narrowly banded with
grayish on lateral portions of hind margin (not shown in illustration); hair of fringe of first
segment yellowish white, that of rest of abdomen brass-colored to very dark; fronto-ocular
triangle (Fig. B) about as long as wide; lacinia with 22-28 teeth, mandible with 40-45 teeth;
shape of legs as shown in figures C-E; calcipala (Fig. F) small but distinct, somewhat
pointed, about as long as wide; subbasal projection of claw very small (Fig. G); eighth
sternite with central portion lighter than lateral portions (Fig. H); gonapophyses subtriangu-
lar, rounded or pointed apically, their inner border slightly sclerotized; paraprocts with
projecting portion angular; cerci with distal border rounded.
Male: Wing length 3.5-3.6 mm; color of head, thorax and legs as in female but the
latter more darkened; abdomen (Fig. I) velvety black; hair, including that of basal fringe,
light brass-colored; segment II with 1+1 very large, silvery white spots closely approach-
ing but not meeting dorsally; segment III with 1+1 small, silvery white spots laterally at
hind margin, these spots in some cases faint; segment VI almost entirely silvery white
except small dark area at center posteriorly; VII with 1+1 medium-sized, silvery white
spots widely separated above; IX faintly iridescent; shape of legs as shown in figures J-L;
hind basitarsus 3.0-3.2 times as long as wide; calcipala (Fig. M) small but distinct, slightly
wider than long; legs with numerous long setae as illustrated; genitalia as for genus in
general shape, but gonostylus distinctly wider than long (Fig. N).
Pupa: Cocoon (Fig. O) like wall pocket or slipper shaped, with very short antero-
ventral bridge; color light brown, translucent; surface smooth, closely woven, threads not
individualized under medium magnification; rim of aperture feebly reinforced; length of
cocoon at dorsum 3.5-4.0 mm; maximum length along base, 5.0 mm; length of pupa, 3.0-3.5
mm; length of gill 2.5-3.0 mm; gills each with six filaments, filaments spreading and forming
fanlike structure; ventral primary branch and its two filaments somewhat bent downward
(Fig. P); head and thorax of pupa dark brown; frontoclypeus without tubercles; head with
2+2-3+3 short frontal, single or bifid, and 1+1 longer facial, trichomes (Fig. Q), latter
generally bifid; thorax on region adjacent to aperture of cocoon with band of tubercles,
becoming evanescent toward sides of thorax; tubercles also occupying large area of dorsal
portion of that part of thorax covered by cocoon; exposed surface of cocoon with large
number of simple or bifid trichomes (Fig. R), their length, 0.05-0.08 mm, their total number,
250-300+250-300; trichomes also present on region anterior to insertion of gill.
Larva: Maximum length, 7.0 mm.; maximum width of head capsule, 0.7 mm; dark
hypodermal pigment on dorsal and ventral surfaces of whole body; pattern of cephalic
apotome as shown in figure S; cuticle with isolated short setae, slightly more numerous on
caudal portion; antennae light brown, subapical segment darkest; ratio of proximal,
medial and distal segments = 1:1.25:1; cephalic fan with about 40 rays; hypostomium
median tooth as prominent as lateral ones; lateral serrations well developed, irregular in
shape; hypostomial setae arranged in two irregular rows, with 10-12 setae in each group;
postgenal cleft rather wide, much deeper than hypostomial bridge length (Fig. T); anal
sclerite, with well-developed scales at base of sclerite; anal ring with approximately 65-70
rows, each composed of about 17 hooks; each lobe of rectal papillae (Fig. U) with
diverticula arranged in two rows, from four to six diverticula on lateral, and from six to
eight diverticula on central lobe.
204 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Distribution: Peru: Cuzco, Junin.


Bionomics: The larvae and pupae of Simulium albilineatum were found on blades of
heavy aquatic vegetation trailing in the stream. The temperature of the water was 10 °C,
and the altitude was about 3700-4000 m.
Discussion: Simulium albilineatum imagoes are similar to those of S. yacuchuspi and S.
herreri but both have more reduced whitish areas on tergites III and V. The more
convergent filaments of the gills and the structure of the thoracic trichomes distinguish

C D
B E

M
F
A

I G

H J
L

K
O
N
U

S R
T
Q
Figure 44. Simulium (Pternaspatha) albilineatum. A-H: Female; A: abdomen, B: fronto-ocular triangle;
C-E: fore, mid and hind leg, respectively; F: apical portion of hind basitarsus and tarsomere I; G:
claw; H: eighth sternite and gonapophyses. I-N: Male; I: abdomen dorsal view; J-L: fore, mid and
hind leg, respectively; M: apical portion of hind basitarsus and tarsomere I; N: gonocoxite and
gonostylus. O-R: Pupa; O: lateral view; P: gill; Q: frontoclypeus and trichomes; R: exposed portion
of thorax and trichomes. S-U: Larva; S: cephalic apotome; T: hypostomium and postgenal cleft; U:
rectal papillae.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 205

the pupa of S. albilineatum from the pupae of S. herreri and yacuchuspi; in the latter two, one
or two filaments of the gills sharply diverge from the others, and the thoracic trichomes
are thicker. Simulium bordai has very similar pupal characters but the thoracic trichomes
have 2 to 6 branches and the number is lower (120-170) on each side.

Simulium (Pternaspatha) bordai COSCARÓN & WYGODZINSKY (Fig. 45.)


Simulium (Pternaspatha) bordai COSCARÓN & WYGODZINSKY, 1972b: 221-222; COSCARÓN, 1991: 124-125.
Female, Male and Larva: Unknown.
Pupa: Cocoon as in Simulium albilineatum, viz., slipper shaped, with short anteroven-
tral bridge, only slightly raised from substrate; cocoon light brown, its surface smooth,
with threads not clearly perceptible; rim of aperture distinctly reinforced; length of
cocoon dorsally, 4.5 mm; maximum length as seen from below, 5.6 mm; gill about 2.5 mm.
long, with six filaments arising from three very short primary branches (Figs. A, B), with
two filaments to each branch; filaments approximately parallel, only slightly diverging at
base; dorsal filament of dorsal branch conspicuously wider on basal portion than
remaining filaments; head and thorax dark brown; frontoclypeus (Fig. C) without
tubercles; 3+3 frontal, 1+1 facial and 1+1 ocular trichomes, in shape of long bifid or
trifid, very rarely simple, setae; facial trichomes inserted in distinct depression delimited
by conspicuously sclerotized ridges; disc of thorax without tubercles, the latter restricted
to area adjacent to aperture of cocoon and at insertion of gills (Fig. D); trichomes
occupying disc of thorax, in shape of setae with three to six branches (Figs. E-F), from
rather short and stout to long and slender, rather uniform or varied; number of trichomes
approximately 120-170+120-170; their average length 0.09-0.11 mm; onchotaxy of
abdomen as usual for subgenus.

C B

Figure 45. Simulium (Pternaspatha) bordai. A-F: Pupa; A-B: gills; C: frontoclypeus; D: exposed
portion of thorax; E: trichomes of disc of thorax; F: trichomes and tubercles of posterior margin
of exposed portion of thorax.
206 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Distribution: Bolivia: La Paz to Yungas.


Bionomics: Simulium bordai breeds in creeks of fast-flowing, crystalline and cold
water, at about 4000 m altitude.
Discussion: Simulium bordai is described and named even though only its pupa is
known, because the latter is conspicuously different from that of any other described
species. The pupa of S. bordai is similar in many ways with that of S. albilineatum, but the
latter differs by the greater number of thoracic trichomes (250-300) that are simple or
bifid, and the gill branches diverging immediately from the base.

Simulium (Pternaspatha) quechuanum COSCARÓN & WYGODZINSKY (Fig. 46.)


Simulium (Pternaspatha) quechuanum COSCARÓN & WYGODZINSKY, 1972b: 232-234; COSCARÓN, 1991:
135-136.
Female: Wing length 3.2-3.7 mm; color of head with appendages, thorax, wings, and
halteres as in Simulium herreri; color pattern of legs as illustrated for S. hectorvargasi; abdomen
(Fig. A) black, with 1+1 large silvery white spots on terga II, VI, and VII; terga III and V, and
in some specimens also IV, with 1+1 smaller silver-white spots; posterior margins of these
terga very narrowly white at sides; terga VIII and IX dark gray; frons and fronto-ocular
triangle as in Simulium herreri; frontal angle 115 degrees; lacinia with 24-26 teeth, mandible
with approximately 35 teeth; wings as in S. hectorvargasi, shapes and proportions of leg
segments as in S. hectorvargasi; calcipala very small but distinct, wider than long as in male
(Fig. C); claws with distinct tooth (Fig. B); genitalia similar to those of S. herreri.
Male: Wing length 3.5 mm; color and structure extremely similar to those of
Simulium hectorvargasi, but calcipala smaller (Fig. C); apex of gonostylus smooth.
Intersex (= specimen with male and female characters): General aspect, color, and
most morphological characters as in females, but genital fork shorter and stouter; legs
with hind basitarsus distinctly widened, and claws of all legs as in male.
Pupa: Cocoon (Figs. D, E) slipper shaped; anteroventral bridge distinct, rather
salient; cocoon dark brown; surface smooth, but with individual threads distinctly
perceptible with moderate magnification; rim of aperture distinctly reinforced; length of
cocoon dorsally, 3.5 mm; maximum length along base, 5.0-5.5 mm; length of body 4.0
mm; of gills 2.0-2.2 mm; gills (Fig. F) each composed of six filaments arising from three
main branches; dorsal and median primary branches and their filaments lying closely
together, ventral primary branch sharply ventrally bent, its ventral filament curved
backward under cocoon, its dorsal filament generally curved forward but not closely
associated with remaining filaments; all primary branches with division close to base of
gill, all about equidistant from base; head and thorax light brown; frontoclypeus (Fig. G)
without tubercles; 2+2 frontal trichomes, extremely varied in shape, often different in
same specimen, from very short, divided, spine-shaped, to elongate bristle-like, generally
divided. 1+1 facial trichomes, also extremely varied in shape, from spinelike (although not
as short as some frontal trichomes) to elongate hair like, with two branches, or simple;
facial trichomes situated in depression delimited by distinct ridges; 1+1 ocular trichomes,
hair like, with two branches; thorax with tubercles forming band along area adjacent to
aperture of cocoon (Fig. F); trichomes of thorax very similar to those of Simulium
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 207

barbatipes, from lanceolate, simple or divided, to elongate hair like (Fig. H), but most
frequently lanceolate; number of thoracic trichomes from 120+120 to 180+180; onchot-
axy of abdomen as usual for subgenus.
Larva: Length of mature larva, 6.5-7.0 mm; width of head capsule, 0.6 mm; color
much as in Simulium hectorvargasi, although somewhat darker; general structure of antennae as
in S. hectorvargasi; ratio of proximal, medial and distal segments = 1:1.7:0.9-1.1; cephalic fan
with 35-38 rays; toothing of mandibles generally with first marginal tooth much over twice
as long as second (Fig. I), although in some individuals only twice as long as second;
structure and pigmentation of hypostomium much as in S. hectorvargasi; hypostomial setae
arranged in two or three irregular series, with 11-16 setae in each group; postgenal cleft (Fig.
J) 2.0-2.5 times as deep as postgenal bridge; anal sclerite spicules or scales not perceptible;
anal ring with 80-85 rows with about 14 hooks each; rectal papillae with three simple lobes.
Distribution: Argentina: Jujuy, San Juan; Chile: Tarapaca.
Bionomics: Simulium quechuanum is a species restricted to the puna or high areas
(3000-4300 m altitude). It is found in streams having cold, crystal clear water, on leaves of
trailing grasses, on aquatic plants such as Potamogeton, and occasionally on the surface of
rocks. The intersexes mentioned in the above description are the first-ever reported from
the subgenus Pternaspatha; they were bred from pupae collected west of Putre. Mermithids
were here found to parasitize several larvae, and might explain the presence of intersexes.
Discussion: Simulium (P.) quechuanum resembles S. barbatipes in the kind and
arrangement of the thoracic trichomes of the thorax, but in S. barbatipes all branches of

G
B C

A
E

I
H J
F

Figure 46. Simulium (Pternaspatha) quechuanum. A-C: Female; A: abdomen, dorsal view B: claw; C:
apical portion of hind basitarsus and tarsite I. D-H: Pupa; D-E: dorsal and lateral views; F: gill and
exposed portion of thorax with trichomes; G: frontoclypeus and trichomes; H: trichomes of
thorax. I-J: Larva; I: marginal teeth; J: hypostomium and postgenal cleft.
208 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

the gill lie close together. The diverging filaments of the ventral primary branch, as
observed in S. quechuanum, are also found in S. herreri and S. yacuchuspi, but the thoracic
trichomes are very different in these species. The adults of S. quechuanum are extremely
similar to those of S. hectorvargasi; the two species are sympatric. The females can be
distinguished by the presence in S. quechuanum of 1+1 white spots on abdominal tergum V,
and the absence of these spots in S. hectorvargasi, and the slightly less-developed calcipala
in S. quechuanum. This latter character is the only one to separate males of the two species.
The very different pupae, distinguished easily by the number of filaments of gills and by
the thoracic trichomes, are sufficient to recognize each species. There seem to be no clear-
cut characters to distinguish the larvae of both species.

Simulium (Pternaspatha) hectorvargasi COSCARÓN & WYGODZINSKY (Fig. 47.)


Simulium (Pternaspatha) hectorvargas COSCARÓN & WYGODZINSKY, 1972b: 222-225; COSCARÓN, 1991:
129-130.
Female: Wing length 3.3-3.5 mm; color of head with appendages, and thorax as in
Simulium herreri; legs brown, their pattern as shown in figures C, D, E.; pattern of abdomen
as shown in figure A; frons and fronto-ocular triangle much as in S. herreri; frontal angle
115 degrees; maxillary palp as illustrated (Fig. B); sensory vesicle half as wide as third
palpomere, its structure as in S. herreri; lacinia with 25-29 teeth, mandible with 38-40 teeth;
Sc glabrous or with one to four setae on central portion; basal section of R glabrous; setae
and spines on R1 and setae on Rs arranged in one irregular row; shapes and proportions of
leg segments as shown in figures C, D, E; calcipala (Fig. F) distinct but small, wider than
high; claws with tooth well developed (Fig. G); genitalia much as in S. herreri; cercus and
paraproct as shown in figure H.
Male: Wing length 3.3-3.5 mm; color of head and its appendages and of thorax and
its appendages very similar to that of Simulium herreri; abdomen (Fig. I) velvety black
dorsally, especially intense on terga III-V; terga II, VI, and VII with 1+1 large sublateral
silvery white spots; tergum III with 1+1 very small whitish spots, not easily perceptible in
every specimen; terga III-V narrowly lighter along hind border; chaetotaxy of wings as in
female, but Sc invariably lacking hair; shapes and proportions of segments of hind leg as
shown in figure J; calcipala small as female; hind basitarsus 3.1-3.5 times as long as wide;
genitalia much as in Simulium nemorale; apex of gonostylus smooth (Fig. K).
Pupa: Cocoon (Figs. L, M) wall slipper shaped, with extremely short anteroventral
bridge, not raised from substrate; cocoon dark brown, closely woven, surface smooth but
with threads distinctly perceivable with moderate magnification; rim of aperture distinctly
reinforced; length of cocoon dorsally 3.7-4.2 mm, maximum length along base, 4.5 mm;
length of body pupa, 3.5-3.8 mm; length of gills 2.5-2.8 mm; gills (Fig. N) each with eight
filaments arising from three main branches, more or less parallel-sided or spreading on
basal half; ventral branch divided rather close to its base; median and dorsal branches
divided at slightly larger distance from their base and again at considerable and rather
variable distance; head and thorax light brown; frontoclypeus (Fig. O) without tubercles;
2+2 frontal trichomes, in shape of very short cones; 1+1 facial trichomes in shape of long
setae; 1+1 ocular trichomes in shape of short spines (Fig. P); facial trichomes inserted
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 209

basad of faint but perceptible transverse ridge; disc of thorax without tubercles, the latter
restricted to area adjacent to aperture of cocoon and at insertion of gills (Fig. Q);
trichomes occupying large part but not all of free surface of thorax, mostly simple hair
like, very rarely bifid (Fig. R); their number approximately 150+150, their length, 0.9-0.12
mm; onchotaxy of abdomen as usual for subgenus.
Larva: Maximum length of mature larva, 6.0-6.5 mm; width of head capsule, 0.5-
0.6 mm; head light brown; cephalic apotome as shown in figure S; in some cases more
G
B

D
J

A E
C

H F

O
I L

P
N
M

Q
R

V T U
S
Figure 47. Simulium (Pternaspatha) hectorvargasi. A-H: Female; A: abdomen, dorsal view; B: maxillary
palp; C-E: fore, mid and hind leg, respectively; F: apical portion of hind basitarsus; G: claw; H:
cercus and paraproct. I-K: Male; I: abdomen, dorsal view; J: hind leg; K: gonostylus. L-R: Pupa; L-
M: cocoon in lateral and dorsal view; N: gill; O: frontoclypeus and trichomes; P: frontal trichome
enlarged; Q: exposed portion of thorax with trichomes; R: trichomes of thorax. S-V: Larva; S:
cephalic apotome; T: antenna; U: teeth of mandible; V: hypostomium and postgenal cleft.
210 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

light colored; antennae (Fig. T) with distal segment darkest; medial with two constric-
tions accompanied by unpigmented areas; ratio of proximal, medial and distal segments
= 1:1.2-1.4:0.6-0.7; cephalic fan with approximately 45 rays; toothing of mandible as
shown in figure U; hypostomium with median tooth as prominent as lateral ones;
hypostomial setae in one or two irregular rows, with 8-15 setae in each group; postgenal
cleft (Fig. V), about twice as deep as length of postgenal bridge; anal sclerite with some
simple spicules; anal ring with 75-80 rows, composed of 15-18 hooks each; rectal
papillae with three simple lobes.
Distribution: Chile: Tarapaca.
Bionomics: Simulium hectorvargas larvae and pupae were collected on blades of grass
and aquatic vegetation in fast-flowing streams with crystal-clear water. The temperature
of the water was very low; ice formed in the mornings in shallow spots. The area in which
this species was collected is typical puna.
Discussion: Simulium albicinctum, S. herreri and S. yacuchuspi show female abdominal
ornamentation very similar to that of S. hectorvargasi, but can be differentiated by the
presence of the grayish tergite VI in the first species, and the other two have 1+1 small
light spots on tergites IV and V. The latter species has a pupal gill with 6 branches. A
distinctive character of S. hectorvargasi is to have vestigial frontal trichomes.

Simulium (Pternaspatha) albicinctum (ENDERLEIN) (Fig. 48.)


Acropogon albicinctus ENDERLEIN, 1934b: 277.
Simulium albicinctus SMART, 1945: 500; VARGAS, 1945a: 109.
Simulium (Pternaspatha) albicinctum WYGODZINSKY & COSCARÓN, 1967: 94-95; COSCARÓN, 1991: 128-129.
Female: Wing length, 3.6 mm; head black; antennae and mouthparts piceous;
occiput, frons, and clypeus gray pollinose; long hair on occiput, frons, clypeus, basal
antennal segments, and basal segments of maxillary palpi; scutum black, median and
submedian stripes dull gray, not contrasting strongly with black areas; pubescence
relatively dense; a few suberect to decumbent, brass-colored, long hairs present;
scutellum, metanotum, pleura, and sterna like those in male; legs brown to yellowish-
brown with coxae very dark, and distal portion of femur, tibia and tarsi darkened; dorsal
surface of abdomen (Fig. A) velvety black; tergite II with 1+1 silvery white spots, dorsal
dark area very small; tergite III with 1+1 smaller spots, their distance from each other
about subequal to their diameter; tergites IV and V entirely black; tergite VI entirely
silvery white; tergite VII black at center, extensively silvery white at sides; tergite VIII
light-colored at sides, otherwise dark but not black; tergite IX entirely dark; hair of fringe
of first segment whitish, hair of posterior segments dark; frons wide; fronto-ocular area
about as long as wide; hind legs as shown in figure B; calcipala small but distinct (Fig. C);
claws with small tooth (Fig. D); genitalia not well known.
Male: Wing length, 3.1 mm; head black; eyes, antennae, and palpi piceous;
clypeus black, silvery pollinose; conspicuous dark long hair on occiput, between eyes,
on clypeus, basal two segments of antennae, and basal segments of maxillary palpi;
scutum velvety black; median longitudinal line not clearly perceptible; pre-scutellar
region and sides grayish at certain angle; anterior border with 1+1 distinct, submedi-
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 211

an, silvery white spots; scutum with rather sparse, long, suberect and decumbent,
brass-colored setae; scutellum dark brown, slightly shining, with long and very long,
erect and decumbent, dark or light-colored hair; metanotum black, slightly shining;
pleura and sterna piceous, with gray pollinosity; pleural tuft silvery; coxae black,
grayish pollinose; trochanter of forelegs dark, femur stramineous, with apical fifth
piceous, tibiae and tarsi piceous, with central portion of tibiae somewhat lighter;
pattern of mid legs like that of fore legs, but first tarsal segment somewhat lighter at
base; pattern of hind legs as shown in figure F; dorsal surface of abdomen (Fig. E)
velvety black; long hair of fringe of first segment, and hair of other segments,
whitish; tergites II and VI each with 1+1 large, silvery white spots, dark dorsal
portion small; at certain angle entire sixth tergite silvery white; tergite VII with 1+1
smaller, silvery white spots; dorsal dark spot large; tergite VIII dark, with small,
lateral, silvery spots visible only at certain angle; wings: Sc and basal section of R
glabrous; setae and spines on R1 arranged in single row, setae on Rs in single row with
numerous long hairs on dorsal surface of most segments; basitarsus slightly less than
three times as long as wide; calcipala small but distinct (Fig. G); genitalia not
examined in detail; gonostylus apparently smooth apically.
Distribution: Peru: High Peru.
Bionomics: Unknown.
Discussion: Simulium albicinctum is only known from the type material, a female and
a male; both were designated types by ENDERLEIN. The female is close to S. yacuchuspi, but
can be differentiated by the presence of 1+1 small submedian light grayish spots on
tergites IV and V, and a median posterior small darker grayish spot on tergite VI. The
males in both species are also characterized by an unusually wide posterior basitarsus but
S. yacuchuspi has 1+1 small light grayish spots on tergite III.

B F

A D

E
Figure 48. Simulium (Pternaspatha) albicinctum. A-D: Female; A: abdomen, dorsal view; B: hind leg; C:
apical portion of hind basitarsus and tarsomere; D: claw. F-G: Male; F: hind leg; G: apical portion
of hind basitarsus. and tarsomere I.
212 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Simulium (Pternaspatha) luchoi COSCARÓN & WYGODZINSKY (Fig. 49.)


Simulium (Pternaspatha) luchoi COSCARÓN & WYGODZINSKY, 1972b: 225-228; COSCARÓN, 1991: 134.
Female: Wing length approximately 3.5 mm; color of head, thorax and wings as
described for Simulium herreri; color pattern of fore and mid legs as illustrated for male, of
hind legs as shown in figure D; abdomen (Fig. A) dark gray; first tergum black; terga II-
VIII velvety black at center, extension of black areas as illustrated, that on second tergum
widened anteriorly; terga II, III, VI, VII, and VIII with 1 +1 sublateral silver-white spots,
large on II and VI, smaller on III, VII, and VIII; fronto-ocular triangle as shown in figure
B; frontal angle 115 degrees; sensory vesicle of maxillary palp as shown in figure C; lacinia
with 22-23 teeth; mandibles not examined; wings as usual for subgenus; chaetotaxy of
wings as in Simulium hectorvargasi; hind legs as shown in figure D; calcipala distinct but
small, subtriangular (Fig. E); claws with distinct tooth (Fig. F); central portion of eighth
sternum darker than peripheral areas; paraproct relatively elongated ventrally (Fig. G).
Male: Wing length 3.3 mm; color of head, thorax, and abdomen much as in Simulium
herreri; color pattern of legs as shown in figures J, K, L; structure of head as in S. herreri,
antennae (Fig. H) and maxillary palp (Fig. I) with long setae; last palpomere more than
twice as long as penultimate; shapes and proportions of segments of legs as in figures J,

B D
K

C F
E J
L

A
G M
I

P
O
R

N S Q

Figure 49. Simulium (Pternaspatha) luchoi. A-G: Female; A: abdomen, dorsal view; B: fronto-ocular
triangle; C: second palpomere with sensory vesicle; D: hind-leg; E: apical portion of hind
basitarsus, with small calcipala; F: claw; G: cercus and paraproct. H-N: Male; H: antenna; I:
maxillary palp; J-L: fore, mid and hind leg, respectively; M: apical portion of hind basitarsus, with
small calcipala; N: gonocoxite and gonostylus. O-S: Pupa; O: gill; P: frontoclypeus; Q: exposed
portion of thorax, with trichomes and tubercles; R: tubercles, high magnification; S: trichomes,
high magnification.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 213

K, L; calcipala present but very small (Fig. M); hind basitarsus slightly less than three times
as long as wide; abdomen with numerous long hairs; gonocoxite and gonostylus as in
figure N; the latter smooth.
Pupa: Cocoon slipper shaped, anteroventral bridge distinct, slightly elevated from
substrate; general aspect of cocoon much as in Simulium quechuanum; cocoon light brown,
translucent; surface smooth, but with individual threads distinctly perceptible; rim of
aperture faintly reinforced; length of cocoon dorsally, 3.3 mm, maximum length at base, 4.0-
4.5 mm; length of body, 3.5 mm; length of gill 2.0 mm; gill composed of eight filaments
arranged as shown in figure O; all filaments lying close together except ventral most
filament, which is sharply ventrally and backwardly bent, situated inside cocoon; head and
thorax light brown; frontoclypeus (Fig. P) without tubercles, with 2+2 frontal, 1+1 facial,
and 1+1 ocular trichomes; facial and ocular trichomes much longer than frontal trichomes;
all trichomes slender, hair like, simple, or with one or two branches; facial trichomes situated
in depression delimited by conspicuous ridges; thorax (Fig. Q) with disc smooth; tubercles
forming band adjacent to aperture of cocoon; these tubercles frequently somewhat pointed
(Fig. R); thorax with 30+30 to 50+50 trichomes (Fig. S), most shortly lanceolate, simple or
divided, with some simple or branched hair like setae interspersed.
Larva: Unknown.
Distribution: Chile: Tarapaca.
Bionomics: Simulium luchoi has been collected at 4100 m altitude.
Discussion: The female of Simulium luchoi is very similar to that of S. hectorvargasi, but S.
luchoi has smaller 1+1 light grayish spots on abdominal tergite III and the darkened area of
tergites III-V is homogeneous. It also can be separated by the larger, rounded calcipala present
in S. hectorvargasi. The males of S. luchoi are very close in structure and color to the males of S.
albicinctum, S. hectorvargasi, and S. quechuanum. Minor characters of the structure and coloring of
the hind legs, such as that in S. luchoi showing 3/5 of the hind femur whitish and in the other
species light brown, may make specific identification possible. The pupa of S. luchoi can be
distinguished by the structure and number of its thoracic trichomes from the other species
with 8 gill branches; that in S. luchoi has one or two of these branches sharply diverging.

Simulium (Pternaspatha) prodexargenteum (ENDERLEIN) (Fig. 50.)


Pternaspatha prodexargenteum ENDERLEIN, 1936: 116.
Simulium (Pternaspatha) prodexargenteum WYGODZINSKY & COSCARÓN, 1967: 109-110; COSCARÓN &
WYGODZINSKY, 1972b: 228-232; COSCARÓN, 1991: 123-124.
Female: Wing length 2.9-3.2 (3.3-3.7) mm; color of head and thorax and their
appendages much as in Simulium nemorale; pattern of legs as shown in figures G-I; general
color of abdomen gray, showing terga II-V velvety black at center; terga II and III with
1+1 large, sublateral, silver-white spots; IV and V at sides of posterior border narrowly
silver-white; VI almost entirely white dorsally, slightly darker at center and at sides, silver-
gray sublaterally; VIII and IX dark gray (Fig. A); frons as shown in figure B; frontal angle
115 degrees; fronto-ocular triangle as shown in figure C; shapes and proportions of
antennal segments as shown in figure D; maxillary palp (Fig. E), sensory vesicle over half
as wide as third palpomere, with abundant tubercles (Fig. F); lacinia with 28-29 teeth,
214 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

mandible with 37-43 teeth; Sc with two to five scattered setae; setae and spines on R1 and
setae on Rs arranged in single row; shapes and proportions of leg segments as shown in
figures G-I; calcipala obsolescent or absent (Fig. J); claws with distinct tooth (Fig. K);
eighth sternum, gonapophyses, genital fork, and spermatheca as in Simulium dureti.
Male: Wing length 3.0 (3.1) mm; scutum velvety black with very narrow median
longitudinal, white line; color pattern of legs very close to female, hind legs as in figures O, P;
abdomen velvety black, with silvery spots as shown in figure L; antenna showing scape and
pedicel with very long hair (Fig. M); maxillary palp with very long setae on basal segments (Fig.
N); diameter of sensory vesicle less than half width of third segment of palp (Fig. O);
chaetotaxy of wing veins as in female; shapes and proportions of hind leg segments as shown
in figures P-Q showing wide basitarsus; calcipala obsolescent (Fig. R) or absent; hind basitarsus
3.0-3.6 times as long as wide; genitalia as in Simulium herreri; apex of gonostylus smooth.
Pupa: Cocoon (Figs. S, T) wall-pocket shaped, but with very short anteroventral
bridge; color light brown; when observed in liquid, shows leaden sheen; cocoon closely
woven, threads not individualized, texture somewhat parchment-like; surface of cocoon
smooth, in some specimens with adhering foreign particles; rim of aperture slightly
reinforced; length of cocoon dorsally along middle, 2.5-3.0 (3.2-3.5) mm; maximum
length along base 3.2-3.8 (4.2-4.5) mm; length of body 2.5-2.7 (3.6-3.7) mm; length of
gills 1.8 (2.5) mm, viz., distinctly shorter than both cocoon and pupa proper; gills from
light to dark brown, each with eight filaments (Fig. U); filaments of dorsal and median
branches and dorsal filament of ventral branch diverging at base, approaching near apex;
lower filament of ventral branch sharply diverging from rest of filaments, ventrally and
backwardly curved; head and thorax of pupa from light to dark brown; frontoclypeus
(Fig. V) without tubercles, and with 3+3 simple, hair like, short frontal, 1+1 in some cases
bifid facial, and 1+1 simple hair like ocular trichomes; facial trichomes inserted in, or
basad of, area of low irregular ridges; thorax with tubercles restricted to margin of
exposed area (Fig. W), attaining base of gill; trichomes numbering approximately 10+10
(14+14), in shape of short simple or bifid setae.
Larva: Length 4.5-4.7 (6.7) mm; width of head capsule, 0.48 (0.70) mm; general
body shape as shown in figure X; color of larva greenish brown, head light brown;
cephalic apotome with pattern very faint, and distinctly darkened near hind margin (Fig.
Y); antennae dark brown (Fig. Z); ratio of proximal, medial and distal segments = 1:1.4-
1.8:0.9-1.0; cephalic fan with 30-40 (39-50) rays; toothing of mandible as shown in figure
a; anterior border of hypostomium strongly pigmented; median tooth as prominent or
more prominent than lateral teeth; marginal serrations distinct; hypostomial setae
arranged in one or two very irregular rows (Fig. b), with 8-12 (14-17) setae in each group;
postgenal cleft (Fig. c) pointed, deep, its depth equal to five times length of postgenal
bridge; anal sclerite much as in Simulium dureti; perianal hair few in number; scales at base
of anal sclerite from inconspicuous to distinct; anal ring with approximately 78 (95) rows,
with 14-15 (16-17) hooks each; rectal papillae with three simple lobes.
Distribution: Argentina: Salta; Bolivia: La Paz; Peru (High Peru).
Bionomics: Larvae of Simulium prodexargenteum live in small creeks with strong
current, crystalline and cold waters at great altitudes (3200-4700 m), flowing over aquatic
vegetation and stones.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 215

Discussion: The material studied is homogeneous regarding morphological features but


heterogeneous regarding some meristic characters. All the known localities that correspond to
the “puna” area are close together. No geographical pattern of variability emerges; it may be

D B
C F

E
K
G H
A I

R
L
J
P
M
Q
O
S

N
W
T

X V

Z
a
U

b c
Y
Figure 50. Simulium (Pternaspatha) prodexargenteum. A-K: Female; A: abdomen, dorsal view; B: frons;
C: fronto-ocular triangle; D: antenna; E: maxillary palp; F: sensory vesicle of maxillary palp; G-I:
fore, mid and hind leg, respectively; J: apical portion of hind basitarsus and tarsomere I. K: claw; L-
R: Male; L: abdomen in lateral view; M: antenna; N: maxillary palp; O: sensory vesicle of maxillary
palp;. S-W: Pupa; S-T: dorsal and lateral views; U: portion of thorax with gill; V: frontoclypeus; W:
portion of P-Q: hind legs; R: apical portion of hind basitarsus thorax with trichomes and tubercles.
X-Z, a-c: Larva; X: lateral view; Y: cephalic apotome; Z: antenna; a: teeth of mandible; b: portion of
hypostomium; c: hypostomium and postgenal cleft.
216 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

significant, though, that the largest measurements were obtained from the lot collected at the
highest altitude. The differences between the populations examined are only meristic, and such
differences by themselves do not necessarily indicate specific differences. The abdominal
ornamentation of females shows similarities with S. nemorale but this species has 1+1 whitish
submedian longitudinal stripes on tergites IV and V, the calcipala well developed, tubercles on
the frontoclypeus and gill branches not much opened basally.

Simulium (Pternaspatha) annulatum PHILIPPI (Fig. 51.)


Simulium annulatum PHILIPPI, 1865: 634; KERTESZ, 1902a: 285; KNAB, 1914a: 18; SILVA FIGUEROA,
1917: 30; PINTO, 1931: 727; EDWARDS, 1931: 153; SMART, 1945: 500; VARGAS, 1945a: 112;
STUARDO ORTIZ, 1946: 42.
Simulium (Pternaspatha) annulatum WYGODZINSKY & COSCARÓN, 1967: 89-94; COSCARÓN & WYGODZ-
INSKY, 1972b: 219-220; COSCARÓN, 1991: 133-134.
Simulium varipes PHILIPPI, 1865: 634; KERTESZ, 1902a: 291; SILVA FIGUEROA, 1917: 30; PINTO, 1931:
730; SMART, 1945: 515; VARGAS, 1945a: 205; VARGAS & DIAZ, 1951: 139; STUARDO ORTIZ, 1946:
41; STONE, 1963: 5.
Simulium (Simulium) varipes EDWARDS, 1931: 152.
Friesia varipes ENDERLEIN, 1929: 327; 1930: 92.
Dasypelmoza varipes ENDERLEIN, 1934b: 275.
Female: Wing length 3.2 mm; color of head and its appendages, and of thorax,
halteres, and wings like that in Simulium nemorale; legs light brown, with darker regions: mid
and hind coxae; apical halves of trochanters; apices of all femora and tibiae; entire fore
tarsus, apex of first tarsal segment of mid tarsus as well as all other segments entirely, apical
third of hind basitarsus, apical two-thirds of second and entire third, fourth, and fifth
segments of hind tarsus; abdomen (Fig. A) black on tergites I-V, dark gray on remainder;
silvery gray areas distributed as follows: narrow line on posterior border of tergite I; 1+1
large lateral spots on tergite II; 1+1 very delicate lines on lateral portions of posterior
borders of tergites III-V; 1+1 large lateral spots, in some cases meeting medially and thus
occupying entire tergite, on tergites VI-IX; ventral surface of abdomen almost entirely gray
pollinose; hair of abdomen, including that of basal fringe, silvery; frons as shown in figure B,
median sulcus obsolete; fronto-ocular triangle slightly wider than long; last segment of
maxillary palp (Fig. C) almost twice as long as penultimate segment; base of cibarium with
well-sclerotized lateral projections; lacinia with approximately 25 teeth, mandible with about
48 teeth; scutum with abundant decumbent setae, longer erect ones on posterior declivity; Sc
with 15-30 setae and several rows of setae also on basal sector of R; setae and spines on R1
arranged in several rows; hair on Rs, in single row (Fig. D); calcipala (Fig. E) well developed,
about as long as wide; claws with well-developed tooth (Fig. F); eighth sternite (Fig. G) with
central portion lighter than lateral areas; gonapophyses rounded apically, beset with
microchaetae, their inner margins more strongly sclerotized; cercus with distal margin
truncate to slightly rounded; projecting portion of paraprocts subtriangular (Fig. H); median
process of genital fork strongly sclerotized, extreme apex widened.
Male: Wing length 3.0 mm; scutum black, with golden-colored, decumbent setae; fore
legs dark brown, mid and hind legs light brown, darkened area almost black, distributed very
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 217

similar to female; setae brown to black; abdomen velvety black dorsally, with golden-colored
hair; tergites II, VI, and VII with 1+1 lateral silvery gray spots; tergite IX dark gray pollinose;
sternites grayish to dull brown; last segment of maxillary palp (Fig. I) more than twice as
long as penultimate segment, and longer than two preceding segments combined; diameter
of sensory vesicle approximately half that of third segment; wings as in female, but hairs on
basal portion of R slightly less numerous; basitarsus of hind legs 4.2 times as long as wide,
calcipala well developed, slightly wider than long; genitalia similar to those of Simulium
nemorale, but the 1+1 translucent, areas of basal plate larger.

B
D

C E
A
G
F
J

H
M N
O
I
m
5m
0.0

L
Q
K

R
T

S U
P
Figure 51. Simulium (Pternaspatha) annulatum. A-H: Female; A: abdomen, dorsal view; B: frons; C:
maxillary palp; D: anterior veins of wing; E: apical portion of hind basitarsus and tarsite I; F: claw;
G: eighth sternite and gonapophyses; H: cercus and paraproct. I: Male, maxillary palp. J-O: Pupa; J:
lateral view; K-L: gills of different specimens; M: frontoclypeus and trichomes; N: exposed portion
of thorax and trichomes; O: thoracic trichomes. P-U: Larva; P: cephalic apotome; Q: antenna; R:
teeth of mandible; S: hypostomium; T: cephalic capsule, ventral view; U: rectal papillae.
218 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Pupa: Cocoon slipper shaped (Fig. J), with elongate anteroventral bridge; color of
cocoon light to dark brown; closely woven, individual threads perceptible; rim of aperture
slightly reinforced; gills protected laterally by cocoon at their extreme base only; length of
cocoon at dorsum 3.3 mm; at base, 4.5 mm; length of pupa body 3.6 mm; length of gill
2.5 mm; gills (Fig. K) with six (exceptionally eight as Fig. L) closely parallel filaments; head
and thorax of pupa distinctly pigmented, more strongly so on frontal region of head and
anterior dorsal portion of thorax; trichomes of head and thorax simple, hair like; head
(Fig. M) with 3+3 frontal, 1+1 facial, and 1+1 ocular, trichomes; region of thorax
adjacent to aperture of cocoon with band of small tubercles, dorsal exposed portion of
thorax covered with long trichomes (Fig. N), approximately 150+150 in number, their
length (Fig. O), 0.14-0.17 mm; abdomen of pupa much like that of Simulium nemorale,
spines on tergite II relatively smaller.
Larva: Maximum length, 6.5 mm; maximum width of head capsule, 0.66 mm; color
greenish brown; head brown; cephalic apotome darkened on median line and 1+1 lateral,
sometimes darkened also on anterior area (Fig. P); body integument with isolated delicate
hairs, latter numerous at base of anal sclerite; antennae brown, pigment most intense on
third segment (Fig. Q); ratio of proximal, medial and distal segments = 1:1.3:0.9; cephalic
fan with approximately 40 rays; toothing of mandibles as shown in figure R; anterior
border of hypostomium (Fig. S) distinctly pigmented, median tooth less prominent than
lateral ones; intermediate teeth gradually increasing in size from inner to outer ones;
sublateral teeth very small; lateral borders of hypostomium serrated, with five small teeth;
hypostomial setae with 7-13 setae in each group; postgenal cleft deep subtriangular, longer
than postgenal bridge (Fig. T); anal sclerite without scales at base; anal ring with about 85
rows, each composed of 13-14 hooks; rectal papillae consisting of three primary lobes,
each bearing three pairs of long secondary lobules (Fig. U).
Distribution: Argentina: Chubut over Andean area, Neuquen, Rio Negro; Chile:
Central and southern region from Santiago to Llanquihue.
Bionomics: Simulium annulatum breeds in rivers with fast-flowing, sometimes
torrential crystalline waters. Females are anthropophilic and are known by the regional
inhabitants as “petros”, the araucanian name.
Discussion: Simulium annulatum differs from all other species of Pternaspatha by the
presence of hair on the basal portion of the radial vein, but agrees with Pternaspatha in all
other essential characters. The pupa shows great similarity with that of S. caprii, but has
fewer trichomes on the thorax (about 100). The female has similar ornamentation but the
posterior rim of tergites III-V has no narrow grayish band as in S. caprii.

Simulium (Pternaspatha) caprii WYGODZINSKY & COSCARÓN (Fig. 52.)


Simulium (Pternaspatha) caprii WYGODZINSKY & COSCARÓN, 1967: 79-83; COSCARÓN & WYGODZINSKY,
1972b: 222; COSCARÓN, 1991: 132.
Female: Wing length 3.1-3.5 mm; head blackish; antenna, maxillary palp, and labium
brown, with eyes and basal segment of palp darkest; first and second antennal segments
somewhat more light-colored than remainder; antenna with short hair, that on two basal
segments distinctly longer; hair of frons, clypeus, occiput, and palpi silvery to black; color
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 219

pattern of scutum very similar to that of Simulium nemorale, with decumbent, brass-colored
to black hair. Median longitudinal light-colored line formed by hair only, not by gray
pigment as in remaining species of the group; scutellum grayish brown, with long silvery
to black hair. Metanotum and pleura dark brown, with velvety violaceous pollinosity;
pleural tuft pale brass-colored; legs brown, with silvery to black hair; following regions
dark: all coxae and trochanters, small spot on base and large one on apex of all femora,
base and apex of all tibiae, entire fore tarsus, apical half of basitarsus and all remaining
segments of mid tarsus, extreme base and apex of basitarsus and second segment, as well
as entire third to fifth segments, of hind tarsus; abdomen (Fig. A) piceous; tergites II-V
velvety black at center, dark gray at sides; tergite II with 1+1 large, silvery white spots;
tergites VI-IX dark, with 1+1 sublateral, silvery white spots decreasing progressively in
size toward posterior segments; ventral surface of abdomen brown; frons as shown in
figure B, with obsolete median sulcus; fronto-ocular triangle approximately as wide as
long; last segment of maxillary palp (Fig. C) much longer than two preceding segments
combined, and three times as long as penultimate segment; diameter of sensory vesicle
approximately equal to half of diameter of segment; structure of vesicle as shown in
figure D; lacinia with about 27 teeth, mandible with about 45 teeth; Sc with approximately
10 setae on central and apical regions; spines and setae on R1 in two very irregular rows;
setae on Rs in one row, in some places in two irregular rows; calcipala well developed,
slightly wider at base than long (Fig. E); claws with short tooth (Fig. F); eighth sternite
with about 30 long setae on each side (Fig. G); gonapophyses subtriangular, with abundant
microtrichiae; inner borders more sclerotized; paraproct and cercus as shown in figure H
with median process well sclerotized.
Male: Wing length 3.3 mm; color of head, thorax, wings, and halteres like that of
Simulium nemorale; legs light brown; darkened in some areas as hind leg (Fig. I); following
dark: all coxae brown; trochanters except base; spot at base of fore and mid, and on apex
of all femora; base and apex of tibiae of all legs; entire fore tarsus, tarsus of mid legs
except basal two-fifths of basitarsus, base and apex of hind basitarsus, basal scale and
apical half of second segment, and entire third to fifth segments, of hind tarsus; abdomen
black, with 1+1 silvery gray spots on tergites II, VI, VII, and VIII, most conspicuous on
second, becoming progressively smaller and fainter on posterior segments; last segment
of maxillary palp longer than two preceding segments combined and sensory vesicle
smaller than half diameter of third segment; wings like those of female, but Sc with not
more than seven setae, and spines and hair on R1 and Rs slightly less numerous; segments
of hind legs as shown in figure I; hind basitarsus narrow, 3.8-4.0 times as long as wide;
calcipala distinct, about as long as wide at its base (Fig. J); gonostylus subquadrate; apical
border widely concave, glabrous (Fig. K).
Pupa: Cocoon (Fig. L) with relatively small, anteroventral bridge, nearly shoe shaped;
color of cocoon dark brown; surface relatively rough, threads individualized; rim of
aperture distinctly reinforced; gills protected by walls of cocoon at their extreme base
only. Maximum length of cocoon dorsally along middle, 4.2 mm; maximum length along
base, 5.2 mm; length of body of pupa 2.5 mm, of gill 2.1 mm; gill (Fig. M) with eight
filaments; head and thorax of pupa dark brown; trichomes of head (Fig. N) exceptionally
long; simple or bifid (Fig. O), until trifid in facial (Fig. P); clypeus without tubercles; thorax
220 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

with band of tubercles adjacent to aperture of cocoon, with numerous interspersed


trichomes (Fig. Q); disc of thorax lacking tubercles but possessing about 100 slender, hair
like, simple or bifid trichomes; latter (Fig. R) with average length, 0.14 mm; chaetotaxy of
abdomen like that in Simulium nemorale.
Larva: Maximum length, 7.5 mm; maximum width of head capsule, 0.7 mm; color
of larva dark greenish brown; cephalic apotome brown, with darker spots (Fig. S); some
curved, short setae at base of anal sclerite; antennae as shown in figure T; brown,
subapical segment darkest; ratio of proximal, medial and distal segments = 1:1.3:0.9;
cephalic fan with 35-40 rays in large fan; toothing of mandible as shown in figure U: two

J
B C
F
I
E
A D

O
G N
K
P
L

U M

Q
mm

V
S
0.05

W R
T

Figure 52. Simulium (Pternaspatha) caprii. A-H: Female; A: abdomen, dorsal view B: frons; C:
maxillary palp; D: sensory vesicle of maxillary palp; E: apical portion of hind basitarsus and
tarsomere I; F: claw; G: eighth sternite and gonapophyses; H: cercus and paraproct. I-K: Male; I:
hind leg; J: apical portion of hind basitarsus and tarsomere I; K: gonostylus. L-R: Pupa; L: lateral
view; M: gill; N: frontoclypeus; O-P: frontal and facial trichomes, respectively; Q: exposed portion
of thorax with trichomes and tubercles; R: thoracic trichomes. S-W: Larva; S: cephalic apotome; T:
antenna; U: teeth of mandible; V: hypostomium; W: rectal papillae.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 221

external, one strong apical, three subapical, six internal, and two marginal teeth, internal
ones variable in size; anterior border of hypostomium strongly pigmented (Fig. V);
median tooth about as prominent as lateral ones; intermediate teeth gradually increasing
in size from center to side; sublateral tooth hardly perceptible; lateral border with six to
seven very small serrations; hypostomial setae in groups of 12-15, arranged in two
irregular series; disc of hypostomium with few short and simple bifid setae; postgenal
cleft deep, pointed, about twice as deep as postgenal bridge; anal sclerite with numerous
scales at base; anal ring with approximately 85 rows, with 13-15 hooks each; rectal papillae
(Fig. W) with three primary lobes, lateral ones each with three or four and median one
with six to eight, diverticula.
Distribution: Argentina: Chubut.
Bionomics: The larvae and pupae of S. caprii were found attached to stems of
grasses trailing in clear streams and on branches of trees that had fallen into the water, as
well as on stones.
Discussion: Simulium caprii is similar to S. nemorale but it is much smaller. The female
of S. caprii differs from that of S. nemorale by the more conspicuous pilosity on the scutum
which also forms a line along the middle of the sclerite (this line is formed by light-
colored pigment in S. nemorale as in all other species of the subgenus), and the central
portion of the eighth sternite, which is darker than the lateral areas. The most obvious
differences are found in the chaetotaxy of the head and thorax of the pupae, with longer
and more abundant trichomes present in S. caprii.

Simulium (Pternaspatha) stelliferum COSCARÓN & WYGODZINSKY (Fig. 53.)


Simulium (Pternaspatha) stelliferum COSCARÓN & WYGODZINSKY, 1972b: 234-239; COSCARÓN, 1991: 137-139.
Simulium (Pternaspatha) schoenemanni WYGODZINSKY & COSCARÓN, 1967: 89 (misidentification).
Female: Wing length 2.9-3.1 mm. (mode, 3.1 mm.); color of head and its
appendages blackish, antennae, palpi and labrum piceous; frons and clypeus grayish
pollinose; scutum as in Simulium nemorale; legs brownish with darker area distributed as
shown in figures F-H; general color of abdomen gray; pattern as shown in figure A; terga
II-V velvety black at center; tergum II with 1+1 large sublateral silver-white spots; terga
III-V narrowly silver-white along hind margin except on median black area; terga VI and
VII each with 1+1 large silver-white spots, those of VII separated by larger dark median
area than those of VI; terga VIII and IX silver-gray; frons as shown in figure B; frontal
angle 115 degrees; fronto-ocular triangle very slightly deeper than wide (Fig. C). Maxillary
palp as illustrated in figure D; sensory vesicle (Fig. E) half as wide as third palpomere;
lacinia with 26-28 teeth, mandible with 42-45 teeth; Sc with 12-14 hairs occupying most of
length of vein; basal section of R glabrous; setae and spines on R1 and setae on Rs
arranged in one or two irregular rows; shapes and proportions of leg segments as shown
in figures F, G, H; calcipala well developed (Fig. I); claws with tooth well developed (Fig.
J); eighth sternum (Fig. K) with central and lateral areas well pigmented; gonapophyses
with inner margins distinctly pigmented; cerci and paraprocts as shown in figure L.
Male: Wing length 2.7-2.9 mm. (mode, 2.8 mm); color of head and its appendages
much as in Simulium nemorale, thorax black, legs as female but slightly darker; abdomen
222 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

(Fig. M) grayish, velvety black dorsally except on segments VIII and IX which are dark
gray; terga II, V, and VI with 1+1 large sublateral silver-white spots; terga III-VII narrowly
lighter along hind margin; scape and pedicel with conspicuous long hair. Maxillary palp
with basal segments with long hair; chaetotaxy of wing veins as in female, but Sc only with
8-10 hairs; shapes and proportions of segments of hind legs as shown in figure N; hind
basitarsus narrow, 4.3-5.5 times as long as wide; calcipala well developed, not quite so long
as wide; gonostylus smooth distally (Fig. O).
Pupa: Cocoon slipper shaped, but with very short anteroventral bridge (Fig. P);
cocoon brown, its surface smooth, closely woven, individual threads not perceptible; rim
of aperture slightly reinforced; length of cocoon dorsally, 3.7-3.8 mm; along base, 4.5-4.7
mm; length of pupa 3.5-3.7 mm; gills 2.5-2.8 mm; gills (Fig. Q) each with six filaments
arising from three main branches; ventral and dorsal primary branches divided very close
to their bases; median primary branch slightly longer; dorsal and median branches and
filaments closely parallel to each other; ventral branch diverging somewhat from
remaining branches, its filaments much narrower than remaining, ventrally but not
backwardly directed; surface structure of filaments as usual for subgenus; head and thorax
of pupa brown, darkest on frontal region and exposed portion of thorax; frontoclypeus
(Fig. R) with very numerous smooth tubercles, and with 2+2 frontal, 1+1 facial and 1+1
ocular trichomes; shape of these trichomes extremely variable, even on opposite sides in one
individual; frontal trichomes generally rather short, with two or three branches, but
occasionally longer; facial trichomes generally hair like, bifid; ocular trichomes short,
inconspicuous; facial trichomes inserted in distinct depression delimited by irregular carinae;
exposed portion of thorax covered with very numerous tubercles similar to those of head,
and several hundred trichomes (Fig. S). Most thoracic trichomes roughly stellate, as shown in
figure T, with three to six branches, the latter short, flattened, pointed apically, slightly
curved; some trichomes long, hair like, branched; such trichomes more numerous in lower
portion of exposed area; onchotaxy of abdomen as usual for subgenus.
Larva: Length 8.5-8.8 mm; width of head capsule, 0.70-0.75 mm; general body
shape of larva as in figure U; color of larva light greenish brown; head brown; anterior
half or two-thirds of cephalic apotome (Fig. V) conspicuously lighter than rest of head,
which is dark brown; in some specimens, pigment on cephalic apotome reduced to narrow
dark band along basal margin (Fig. W); antenna (Fig. X) light brown, third segment
darkest; ratio of proximal, medial and distal segments = 1:1.2:0.8; cephalic fan with 37-39
rays; toothing of mandible as shown in figure Y; anterior border of hypostomium strongly
pigmented. Median tooth of hypostomium (Fig. Z) almost as prominent as lateral teeth;
marginal serration distinct but not very pronounced; hypostomial setae arranged in one or
two irregular rows, with 12-15 setae in each group; postgenal cleft pointed or narrowly
rounded apically (Fig. a); postgenal bridge shorter than hypostomium length; sclerite of
thoracic proleg with about 15 setal groups of different length (Fig. b); anal sclerite as
shown in figure c, characterized by very extensive sclerotized areas between anterior and
posterior arms; base of anal sclerite lacking scale like cuticular structures, at most with
small number of spicules; anal ring with 81-83 rows composed of 14-15 hooks each;
rectal papillae (Fig. d) with three primary lobes, either simple or each with not more than
five lobules.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 223

B D
G
E H

L
C
A
I J
F
K

M V
R
X
S
U
W

a Y

T
b
Q
Z d

Figure 53. Simulium (Pternaspatha) stelliferum. A-L: Female; A: abdomen, dorsal view; B: frons; C:
fronto-ocular triangle; D: maxillary palp; E: sensory vesicle of maxillary palp; F-H: fore, mid and
hind leg, respectively; I: apical portion of hind basitarsus and tarsomere I; J: claw; K: eighth sternite
and gonapophyses; L: cercus and paraproct. M-O: Male; M: abdomen, dorsal view; N: hind leg; O:
gonostylus. P-T: Pupa; P: lateral view; Q: gill; R: frontoclypeus and trichomes; S-T: thoracic
trichomes and tubercles. U-Z, a-d: Larva; U: lateral view; V-W: cephalic apotome in different
specimens; X: antenna; Y: teeth of mandible; Z: anterior portion of hypostomium; a: hypostomium
and postgenal cleft; b: lateral sclerite of thoracic proleg; c: anal sclerite; d: rectal papillae.
224 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Distribution: Argentina: Río Negro isolated on the east low Patagonian steppe;
Chile: from Santiago to Tarapaca on the lowlands.
Bionomics: Simulium stelliferum has a comparatively wide ecological spectrum. It is
found mainly in fast-flowing streams but also occasionally in irrigation ditches, at altitudes
from 200 to 2500 m, and at water temperatures of 6-17 °C. The larvae and pupae were
found attached to vegetation, such as grass or small branches of other plants trailing in the
current, but also, in some cases, to the surface of rocks. The streams in which specimens
were collected were mostly clear, but sometimes S. stelliferum was obtained in slightly
turbid waters containing abundant diatoms and other organic matter; in the latter case, the
current was exceedingly fast.
Discussion: The adults of S. stelliferum are most similar to those of S. caprii from
which they can be distinguished by tergites III-V being entirely black in the latter. The
pupa differs from all other species of the subgenus by the unique structure of the
trichomes of the thorax. The larva of S. stelliferum lacks peculiarities, and varies in some
key characters; consequently, it had to be placed in the key more than once.

Species inquirenda

Simulium (Pternaspatha) schoenemanni (ENDERLEIN)


Acropogon schoenemani ENDERLEIN, 1934b: 277.
Simulium schoenemanni SMART, 1945: 513; VARGAS, 1945a: 195; STUARDO ORTÍZ, 1946: 141.
Simulium (Pternaspatha) schoenemanni WYGODZINSKY & COSCARÓN, 1967: 89.
WYGOZINSKY & COSCARÓN (1967) tentatively identified certain Chilean specimens of
Pternaspatha as Simulium schoenemanni. Careful re-examination led them to conclude that the
females fail to agree with the characters given by ENDERLEIN (1934b) for the female type,
the only specimen known of S. schoenemanni, especially as to the color pattern of the
abdomen. The type of S. schoenemanni could not be found at the Zoological Museum in
Berlin, where it was deposited, according to ENDERLEIN (1934b), nor at the Staatliches
Museum fûr Tierkunde, Dresden, where the type might have been deposited by error. The
species formerly identified by WYGODZINSKY & COSCARÓN as S. schoenemanni was described
later as Simulium(Pt.) stelliferum COSCARÓN & WYGODZINSKY, 1972b, but S. schoenemanni itself
continues to be an enigma.

Simulium (Notolepria) ENDERLEIN (Figs. 54-56, 139.)


Type-species: Simulium exiguum ROUBAUD, 1906b: 108.

Female: Species of small size, wing length 2.2-2.5 mm; coloration blackish; scutum
black, bordered laterally and posteriorly by a silvery pollinose area; scutum and scutellum
generally with bright golden hair grouped and simulating scales; abdomen black, tergites
V-VIII with cereous sheen; fronto-ocular triangle absent or reduced; cibarium with basal
portion smooth, scarcely sclerotized, without reinforcement; wings with Sc and R bare;
legs with petaloid hair; claws with or without subbasal teeth; calcipala well developed,
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 225

about as long as wide; hind basitarsus with normal width; gonapophyses subtriangular
with internal border concave, and well sclerotized; paraprocts about as long as wide with
posterior border acuminate and generally with thin elongated process; genital fork with
branches well sclerotized with stem about two times longer than posterior branches and
scarcely widened apically; spermatheca ovoidal.
Male: Wing length 2.0-2.3 mm; coloration black; scutum velvety black, bordered by
pollinose grayish area; hind basitarsus not inflated; abdomen with II, VI, VII with 1+1
light grayish spots, smaller on VIII tergite; gonostylus half as long as gonocoxite,
subtriangular and with reduced obsolescent apical spine; ventral plate rather acuminate
distally, nearly as long as wide, sometimes with median constriction.
Pupa: Cocoon slipper shaped, generally slightly projected basally, weave not too
thick, with visible threads; gill with 6-8 branches; frontoclypeus and exposed portion of
thorax with abundant tubercles; frontoclypeus with 2 frontal and facial trichomes on each
side, with one to 3-6 branches; thorax with 5-6 trichomes on each side with 3-6 branches;
abdomen with apical spur on last tergite.
Larva: Cephalic apotome darkened on base, generally with median and lateral spots
not well defined; antennal medial segment longer than proximal and this about 1/3
hypostomium length. Mandible with several rows of teeth, and two marginal teeth thin,
curved and with second tooth small; latero-mandibular process not evident; thoracic
proleg lateral sclerite with teeth arranged in about 9 groups; anal ring with 52-76 rows of
about 15 hooks each; rectal papillae with 0-12 lobules on each lobe.
Distribution: Northern Argentina to Mexico, primarily in the intertropical occi-
dental area (Fig. 139).
Discussion: This subgenus is made up of 10 species: 5 with scarce information and
3 of which are considered species inquirendae.

KEYS TO SIMULIUM (NOTOLEPRIA) SPECIES

Females
1 Scutum and scutellum with cylindrical hairs distributed homogeneously, without
bright metallic scales; with some light, showing 1+1 anterior subtriangular light
grayish spots; paraproct with small distal process (Fig. 56 E) ........ Simulium llutense
– Scutum and scutellum with flattened hairs (Fig. 5 B) arranged in groups (Fig. 5 A)
appearing as bright metallic scales; with some light, not showing 1+1 anterior
grayish subtriangular spots; paraproct with distal process generally well developed
(Figs. 54 I, Q, X) ................................................................................................................ 2
2(1) Paraproct with reduced distal process (Fig. 54 X); posterior branches of genital
fork long and thin and scarcely sclerotized ....................................................................
................................................. Simulium gonzalezi, Simulium subexiguum, Simulium sp. A
– Paraproct with well-developed distal process (Figs. 54 I, Q); posterior branches of
genital fork relatively short and wide, and well sclerotized ...................................... 3
3(2) Paraproct distal process relatively short and hooked; eighth sternite well
sclerotized medially and distally ................................................... Simulium cuasiexiguum
226 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

– Paraproct distal process relatively long (Figs. 54 I, Q); eighth sternite not well
sclerotized medially and distally (Fig. 54 F) ................................................................. 4
4(3) Paraproct process thick at base (Fig. 54 I); genital fork stem comparatively shorter
in relation to posterior branches (Fig. 54 H) ................................... Simulium exiguum
– Paraproct process thin at base (Fig. 54 Q); genital fork stem comparatively longer
in relation to posterior branches (Fig. 54 R) ............................. Simulium paraguayense

Males
1 Scutum with normal hairs arranged homogeneously; ventral plate very short
(Fig.56 G) ................................................................................................... Simulium llutense
– Scutum with flattened hairs arranged in groups appearing as bright scales; ventral
plate more elongated (Figs. 54 O, P, U, W) .................................................................. 2
2(1) Gonostylus subtriangular (Figs. 54 M, N); ventral plate without (Fig. 54 O) or with
(Fig. 54 P) lateral constriction ...........................................................................................
.................................................. Simulium exiguum, Simulium subexiguum, Simulium sp. A
– Gonostylus subquadrate (Figs. 54 S, T); ventral plate with well-defined lateral
constriction .......................................................................................................................... 3
3(2) Ventral plate with strong mediolateral constriction appearing like wide median
spatulate process (Figs. 54 U-W) ................ Simulium gonzalezi, Simulium paraguayense
– Ventral plate with lateral constriction appearing as narrow distomedial process ..
.............................................................................................................Simulium cuasiexiguum

Pupae
1 Gills with 6 branches (Fig. 55 L) .................................................................................... 2
– Gills with 8 branches (Fig. 55 K) .................................................................................... 3
2(1) Frontal and facial trichomes simple ............................................ Simulium cuasiexiguum
– Frontal trichomes with 1-3 branches, facial trichomes with 3-5 branches ..............
................................................................................................................... Simulium gonzalezi
3(1) Cocoon extended ventrally and closed anteriorly (Fig. 56 H) ........ Simulium llutense
– Cocoon slightly extended ventrally and open anteriorly (Figs. 55 A, I) ................. 4
4(4) Cocoon not extended ventrally (Fig. 55 I) ................................. Simulium paraguayense
– Cocoon with short ventral projection (Fig. 55 A) ....................................................
.................................................. Simulium exiguum, Simulium subexiguum, Simulium sp. A

Larvae
Larvae of Simulium cuasiexiguum, Simulium subexiguum and Simulium sp. A are unknown.
Simulium gonzalezi larvae are not included due to lack of information.

– Cephalic apotome with well-developed positive spots (Fig. 56 K); rectal papillae
without diverticula; anal sclerite without scales ................................. Simulium llutense
– Cephalic apotome with negative or scarcely evident spots (Fig. 55 C); rectal
papillae with 4-12 diverticula on each lobe (Fig. 55 G); anal sclerite with abundant
scales (Fig. 55 H) ............................................. Simulium exiguum, Simulium paraguayense
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 227

CLAVES DE ESPECIES DE SIMULIUM (NOTOLEPRIA)


Hembras
1 Pelos del escudo y escutelo cilíndricos distribuidos uniformemente sin brillo
metálico; con cierta iluminación presenta 1+1 manchas grisáceo claro; paraprocto
con pequeña prolongación distal (Fig. 56 E) ..................................... Simulium llutense
– Pelos del escudo y escutelo achatados (Fig. 5 B) dispuestos en grupos (Fig. 5 A)
con aspecto de escamas de brillo metálico; con cierta iluminación no presenta 1+1
manchas anteriores grisáceas; paraproctos con prolongación distal generalmente
bien evidente (Figs. 54 I, Q, X) ...................................................................................... 2
2(1) Paraprocto con prolongación distal reducida (Fig. 54 X); ramas posteriores de la
horquilla genital larga, delgada y poco esclerotizadas ..................................................
................................................. Simulium gonzalezi, Simulium subexiguum, Simulium sp. A
– Paraprocto con prolongación distal bien desarrollada (Fig. 54 I, Q); ramas
posteriores de la horquilla genital relativamente cortas y bien esclerotizadas ...... 3
3(2) Prolongación distal del paraprocto relativamente corto y en forma de gancho;
octavo esternito fuertemente esclerotizado mediano y distalmente .........................
............................................................................................................. Simulium cuasiexiguum
– Prolongación distal del paraprocto relativamente larga (Fig. 54 I, Q); octavo
esternito no muy esclerotizado mediano y distalmente (Fig. 54 F) ......................... 4
4(3) Prolongación del paraprocto grueso en la base (Fig. 54 I); tallo de la horquilla genital
comparativamente más corto en relación a las ramas posteriores (Fig. 54 H) ...........
.................................................................................................................... Simulium exiguum
– Prolongación del paraprocto fino en la base (Fig. 54 Q); horquilla genital
comparativamente más larga en relación con las ramas posteriores (Fig. 54 R) ....
............................................................................................................. Simulium paraguayense

Machos
1 Escudo con pelos normales dispuestos homogéneamente; placa ventral muy corta
(Fig. 56 G) .................................................................................................. Simulium llutense
– Escudo con pelos achatados dispuestos en grupos con aspecto de escamas; placa
ventral más alargada (Figs. 54 O, P, U, W) .................................................................... 2
2(1) Gonostilo subtriangular (Figs. 54 M, N); placa ventral sin(Fig. 54 O) ó con (Fig. 54
P) contracción lateral ........... Simulium exiguum, Simulium subexiguum, Simulium sp. A
– Gonostilo de forma subcuadrada (Figs. 54 S, T); placa ventral con notoria
contracción lateral .............................................................................................................. 3
3(2) Placa ventral con fuerte contracción medio-lateral, mostrándose como una
prolongación espatulada mediana (Figs. 54 U-W) ........................................................
............................................................................ Simulium gonzalezi, Simulium paraguayense
– Placa ventral con lateral contracción, mostrándose como una prolongación
medio-distal ...................................................................................... Simulium cuasiexiguum

Pupas
1 Branquias con 6 ramas (Fig. 55 L) ................................................................................. 2
228 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

– Branquias con 8 ramas (Fig. 55 K) ................................................................................. 3


2(1) Tricomas frontales y faciales simples .......................................... Simulium cuasiexiguum
– Tricomas frontales con 1-3 ramas, faciales con 3-5 ramas .......... Simulium gonzalezi
3(1) Capullo bien extendido ventralmente y cerrado anteriormente (Fig. 56 H) ...........
...................................................................................................................... Simulium llutense
– Capullo escasamente extendido ventralmente y abierto anteriormente (Figs. 55 A, I) ..... 4
4(3) Capullo no extendido ventralmente (Fig. 55 I) ......................... Simulium paraguayense
– Capullo con una corta prolongación ventral (Fig. 55 A) .............................................
.................................................. Simulium exiguum, Simulium subexiguum, Simulium sp. A

Larvas
Larvas de Simulium cuasiexiguum, Simulium subexiguum y Simulium sp. A no se conocen y la de
Simulium gonzalezi no se incluye por escasa información.

– Apotoma cefálico con manchas positivas bien evidentes (Fig. 56 K); papilas rectales
sin divertículos; esclerito anal sin escamas ........................................... Simulium llutense
– Apotoma cefálico con manchas negativas o escasamente evidentes (Fig. 55 C);
papilas rectales con 4-12 divertículos sobre cada lóbulo (Fig. 55 G); esclerito anal
con abundantes escamas (Fig. 55 H) .......... Simulium exiguum, Simulium paraguayense

Simulium (Notolepria) exiguum ROUBAUD (Figs. 5D, 54 A-P; 55 A-H.)


Simulium exiguum ROUBAUD, 1906b: 108-109; WYGODZINSKY, 1951a: 214-217; TIDWELL et al., 1980:
119, 121, 126.
Simulium (Notolepria) exiguum DALMAT, 1955: 112; COSCARÓN & WYGODZINSKY, 1975: 227-287;
COSCARÓN, 1987: 20; SHELLEY et al., 1989: 89; COSCARÓN, 1991: 147-148; SHELLEY et al., 2000:
177. COSCARÓN ARIAS, 2003: 195, 198, 204, 209, 215, 219.
Wilhelmia exigua ENDERLEIN, 1921a: 200.
Simulium delpontei PATERSON & SHANNON, 1927: 739.
Simulium glaucophthalmun KNAB, 1914c: 123.
Simulium urubambanum ENDERLEIN, 1934b: 288.
Female: Wing length 1.8-2.5 mm; general color black; antennae dark brown; frons
nacreous; scutum and scutellum with bright golden to greenish scales, formed by flat, grouped
hairs (Figs. 5 A, B); pleurae with gray pruinosity; legs yellowish brown with darkened areas on
first and hind coxae, 3/4 of hind femora, apex of tibiae, tarsi, as shown in male figures 54 J-L;
abdomen blackish matte with tergites II-IV brown blackish and V-VII with cereous sheen;
fronto-ocular triangle not developed (Fig. 54 A); cibarium with basal portion as in figure 54 B;
wing chaetotaxy of anterior veins as in figure 54 C. Median claw with subbasal tooth variable
on fore leg (Figs. 5 E, 54 D), present on median, and hind legs (Fig. 54 E); legs with cylindrical
and flattened hair (Fig. 5 D); eighth sternite with median area slightly darkened and with about
15 setae on each side (Fig. 54 F); gonapophyses without setae, and sclerotized on internal
margin; spermatheca ovoidal with insertion to spermathecal ducts membranous (Fig. 54 G);
genital fork with well-sclerotized branches ovoidal (Fig. 54 H); cerci hemispherical distally,
paraprocts subtriangular with acuminate distal posterior process (Fig. 54 I).
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 229

Male: Wing length 1.5-2.2 mm; scutum velvety black, bordered posteriorly by gray
pollinosity with pilosity similar to scales, as in female; legs as shown in figures 54 J-L;
length/width ratio of hind basitarsus = 7.1-7.4; abdomen velvety black with silvery
tergites II and VI and 1+1 lateral spots of this color on tergite VII; genitalia as shown in
figure M; gonostylus subtriangular with distal spur obsolescent (Fig. 54 N); ventral plate
subtriangular without lateral decolletage (Figs. 54 O, P). Median sclerite with median
depression and endoparameres with large hooks (Fig. 54 M).
Pupa: Cocoon length on dorsum 2.0-2.3 mm; on base 2.0-2.6 mm; body of pupa
1.9-2.2 mm; gills 1.2-1.8 mm; color brown, slipper shaped (Fig. 55 A); eight gill branches
emerging from three primary branches, very similar to those of Simulium paraguayense (Fig.
55 K); frontoclypeus and thorax with abundant platelets; frontal and facial trichomes with
2-3 branches, thoracic with 2-6 branches; abdominal chaetotaxy as in figure 55 B.
Larva: Maximum length 3.6-4.4 mm; color light gray-greenish, head light brown;
cephalic apotome light brown with basal median portion darkened (Fig. 55 C); antenna as
in figure 55 D, ratio of proximal, medial and distal segments = 1:1.1:1; mandible with thin
and curved marginal teeth, as in Simulium paraguayense (Fig. 55 E); cephalic fan with about
40 rays; hypostomium well pigmented on anterior margin; setae 3 or 4 in number disposed
in one row; postgenal cleft deep, rounded anteriorly (Fig. 55 F), postgenal bridge about 1/
3 of hypostomium length; thoracic proleg sclerite with 9-10 groups of spines; rectal
papillae with 4-8 diverticula on each lobe (Fig. 55 G); anal sclerite with abundant scales
arranged in two to four rows (Fig. 55 H); anal ring with 70-76 rows with 13-15 hooks each.
Distribution: Argentina: Jujuy, Salta, Tucumán; Bolivia: Beni, Cochabamba, Tarija;
Brazil: Amazonas, Distrito Federal, Goiás, Mato Grosso, Minas Gerais, Roraima, São
Paulo; Colombia: Cauca, Magdalena, Meta, Valle; Ecuador: Bolivar, Bucay, Cañar,
Cotopaxi, El Oro, Esmeraldas, Guayas, Imbabura, Loja, Los Rios, Manabi, Napo, Pastaza,
Pichincha, Puyo; Guatemala; Mexico: Morelos; Panama: Darien; Peru: Huanuco, Junin,
Lima; Venezuela: Amazonas, Anzoategui, Aragua, Barinas, Bolivar, Carabobo, Cojedes,
Distrito Federal, Guarico, Lara, Mérida, Miranda, Monagas, Sucre, Tachira, Trujillo,
Yaracuy, Zulia.
Bionomics: Simulium exiguum breeds in creeks and rivers with fast currents, in
crystalline and temperate water flowing over aquatic vegetation and branches of trees or
rocks, sometimes up to 2 m deep. Females bite humans and animals, both domestic and
wild, such as tapirs. Simulium exiguum is a vector of Onchocerca volvulus in Ecuador,
Colombia, Venezuela, and Brazil, and of Mansonella ozzardi in Argentina.
Discussion: At least five species, Simulium paraguayense, S. cuasiexiguum, S. subexiguum,
S. sp. A and S. gonzalezi, are very close to Simulium exiguum. The first species shows a few
differences, such as in the female paraprocts, gonostylus and ventral plate, and some
variations in ornamentation, size of legs, and proportion of larval antennae, as explained
for that species. Simulium gonzalezi also shows differences in paraproct shape and especially
in the number of pupal gill branches. Simulium cuasiexiguum is also differentiated by the 6
gill branches of the pupa. Simulium subexiguum has a short distal posterior process on the
paraproct (similar to S. gonzalezi). As shown in Simulium sp. A, some of these differences
are considered by SHELLEY et al. (1989) as possible variants of Simulium exiguum. It is also
possible to see great variation in size and female feeding habits (SHELLEY et al., 2000),
230 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

B
C

A P

J
O

K
D
M
E N

S
G
L

F T

U
Q
I
H
X
R
V
Figure 54. A-P: S. (Notolepria) exiguum; A-I: Female; A: fronto-ocular triangle; B: Basal portion of
cibarium; C: wing anterior portion of chaetotaxy; D: claw without subbasal tooth; E: claw with
subbasal tooth; F: eighth sternite and gonapophysis; G: spermatheca; H: genital fork; I: cercus and
paraproct. J-P: Male; J-L: fore, mid and hind leg, respectively; M: genitalia showing gonocoxite,
gonostylus, ventral plate, endoparameres and median sclerite; N: gonostylus; O, P: ventral plate of
different specimens. Q-W: S. (N.) paraguayense; Q-R: Female; Q: cercus and paraproct; R: genital
fork; S-W: Male; S-T: gonostylus in different position; U-W: ventral plate in different position. X: S.
(N.) gonzalezi, cercus and paraproct.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 231

common in species with large geographic distributions. As indicated by the presence of at


least 4 cytospecies in Ecuador (CHARALAMBOUS et al., 1993), it is probable that there are
more morphological differences in the range of S. exiguum, and these small variations
could correspond to sibling species. We prefer to maintain S. exiguum as a species, pending
further information, such as cytological or molecular, to clarify its identity.

Simulium (Notolepria) paraguayense SCHROTTKY (Figs. 54 Q-W; 55 I-K.)


Simulium paraguayense SCHROTTKY, 1909: 63; LUTZ, 1909: 144-145; 1910: 235; SURCOUF & GONZÁLEZ
RINCONES, 1911: 291-292; LUTZ et al., 1918: 165; LUTZ & NUÑEZ TOVAR, 1928: 43;
WYGODZINSKY, 1953a: 298-301.
Simulium (Eusimulium) paraguayense LUTZ, 1909: 144; ORFILA, 1939: 1531.
Eusimulium paraguayense PINTO, 1931: 678; LANE & PORTO, 1939: 169, 175; BRICEÑO IRAGORRI, 1941:
114; LANE & VULCANO, 1943: 432.
Simulium (Notolepria) paraguayense COSCARÓN & WYGODZINSKY, 1975: 283-287; COSCARÓN, 1987: 20;
1991: 148-150; SHELLEY et al., 2001: 485.
Female: Size, ornamentation and morphology very similar to those of Simulium
exiguum but differentiated by the following: claw subbasal tooth always present on all legs;
hind leg with half of distal portion of femur, apex of basitarsus, and second and third
tarsomeres more darkened; paraproct distal process thinner acuminate apically and more
elongated (Fig. 54 Q); genital fork stem proportionally more elongated and posterior
branches more sclerotized (Fig. 54 R).
Male: Very similar to Simulium exiguum. Maximum wing length 2.5 mm; gonostylus
subquadrate to subovoidal with very small spur (Figs. 54 S, T); ventral plate with lateral
constriction at about half length (more or less evident in accordance with position, (Figs.
54 U, V) and with small median elevation in lateral view (Fig. 54 W).
Pupa: The small differences from Simulium exiguum are the more translucent cocoon,
permitting the pupal body to be seen(Fig. 55 I) and more abundant platelets on the
frontoclypeus and thorax (Figs. 55 J-K).
Larva: Greatly similar to S. exiguum; showing only small differences in antennal
segment ratios: 1:1.2:1.4; anal ring with 52-62 rows 11-13 hooks each, and presence of one
latero-mandibular process.
Distribution: Argentina: Misiones; Brazil: Bahia, Minas Gerais, Rio de Janeiro, São
Paulo (cited by LUTZ); Paraguay; Venezuela: Aragua, Distrito Federal (cited by LUTZ &
NUÑEZ TOVAR).
Bionomics: Simulium paraguayense breeds in swift, large river up to 1 m deep, flowing
over Podostemaceae. Females show great anthropophily and their bites are very painful.
Discussion: Simulium paraguayense has only small differences, as shown in the
description, that permit it to be separated from S. exiguum. These could be considered
population differences, but S. exiguum has been known until now from 4 cytotypes only in
Ecuador, showing the presence of several species in the same morphotypic species. This is
the motive for not synonymizing S. paraguayense with S. exiguum. Simulium incertum LUTZ, with
8 pupal gill branches, probably is a synonym of S. paraguayense, which LUTZ also suspected.
Material collected by us very close to the type area corresponds to S. paraguayense.
232 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

D
B

L
C

G
F H

Figure 55. A-H: S. (Notolepria) exiguum; A-B: Pupa; A: general aspect; B: abdominal chaetotaxy. C-
H: Larva; C: cephalic apotome; D: antenna; E: dental portion of mandible; F: hypostomium and
postgenal cleft; G: rectal papillae; H: anal sclerite with rows of scales on superior area and some
hair on inferior. I-K: S. (N.) paraguayense; I-K: Pupa; I: general aspect; J: frontoclypeus; K: dorsal
portion of thorax with trichomes and gill. L: S. (N.) gonzalezi; Pupa, dorsal portion of thorax
showing tubercles, trichomes and gill.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 233

Simulium (Notolepria) gonzalezi VARGAS & DÍAZ NÁJERA (Figs. 54 X, 55 L.)


Simulium gonzalezi VARGAS & DÍAZ NÁJERA, 1953b: 235; SHELLEY & ARZUBE, 1985: 329.
Simulium (Notolepria) gonzalezi VARGAS & DÍAZ NÁJERA, 1957b: 158, 338; COSCARÓN, 1987: 20;
SHELLEY et al., 1989: 88; IBÁÑEZ-BERNAL, 1992: 263; CHARALAMBOUS et al., 1997: 19; SHELLEY et
al., 2002: 135; COSCARÓN ARIAS, 2003: 195, 198, 204, 208, 215, 219.
Simulium gonzalesi VULCANO, 1967: 12 (incorrect spelling).
Simulium exiguum KNAB, 1913: 154; VARGAS, 1941a: 115; 1941b: 213; 1942a: 229; 1943b: 363-365.
Simulium (Notolepria) exiguum VARGAS et al., 1946: 171; DALMAT, 1955: 112-116.
Female: Wing length 1.5-2.2 mm; color and morphology of head, scutum, abdomen
and appendages as in Simulium exiguum; claws without basal tooth, sometimes restricted to
hind leg; paraproct with distal process more reduced than in S. exiguum (Fig. 54 X).
Male: Wing length 1.1-2.0 mm; color and morphology very similar to that of
Simulium exiguum; gonostylus subquadrate, very close to that of S. paraguayense; ventral
plate very close to that of S. exiguum; some males dichoptic with external morphology
similar to that of female, with narrow frons and more elongated antennal segments;
abdominal ornamentation similar to that of S. exiguum.
Pupa: Cocoon length dorsally 1.4-2.0 mm, along base 1.4-2.5 mm, pupa body 1.9-2.4
mm, gills 1.0-1.8 mm. Cocoon slipper shaped, with reinforced rim; gill with 6 branches
(Fig. 55 L); chaetotaxy of cephalothorax and abdomen as in Simulium exiguum.
Larva: Maximum length 4.2 mm; coloration and morphology as in Simulium exiguum;
negative cephalic apotome pattern of Simulium gonzalezi also found in some specimens of
S. exiguum (CHARALAMBOUS et al., 1997).
Distribution: Belize: Cayo, Stan Creek, Toledo; Ecuador: Esmeraldas; Guatemala;
Mexico: Colima, Chiapas, Guerrero, Hidalgo, Jalapa, Michoacan, Nayarit, Oaxaca, Potosi,
San Luis, Tabasco, Tamaulipas, Veracruz.
Bionomics: Simulium gonzalezi breeds in fast-flowing small creeks to rivers, from 1 to
1500 m in altitude in water of about 20 °C, on aquatic vegetation. Females are
haematophagous, biting humans, domestic mammals and birds. This species is recognized
as a vector of Onchocerca volvulus in Mexico and Guatemala.
Discussion: Several Simulium (Notolepria) species, such as S. exiguum, S. paraguayene, S.
cuasiexiguum, S. subexiguum and Simulium sp. A, are very similar to S. gonzalezi; it is possible
to differentiate the first two by the 8 gill branches and more elongate distal process on the
paraproct. The small differences in the ventral plate and gonostylus, as well as the negative
spots on the cephalic apotome of larvae of S. gonzalezi, are included in the variation of
characters of the other two species. All compared species (except S. cuasiexiguum) have
pupal gills with 8 branches.

Simulium (Notolepria) cuasiexiguum SHELLEY, LUNA-DIAS, MAIA-HERZOG & LOWRY


Simulium (Notolepria) cuasiexiguum SHELLEY et al., 2000: 178; SHELLEY et al., 2001: 483; SHELLEY et al.,
2002: 154.
Female: Wing length 1.5-2.1 mm; coloration and morphology similar to Simulium
exiguum, S. paraguayense and S. gonzalezi; the great differences are in the bigger amplitude of
234 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

sclerotized area of eighth sternite, the thinner and curved distal paraproct process, and
the insertion of spermathecal duct to sclerotized spermatheca well, that can reach to the
basal portion of spermathecal duct.
Male: Wing length 1.4-1.8 mm; as in the female, it is very similar to the other tree
species in this subgenus; it shows differences in the more acuminate ventral plate distal
process and with Simulium paraguayense and S. gonzalezi by the subquadrate gonostylus.
Pupa: Cocoon length dorsally 2.1-2.7 mm, ventrally 2.2-3.1 mm; body pupa 2.0-2.1
mm; gills 1.3-1.5 mm; gill has 6 branches with similar arrangement as Simulium gonzalezi.
Larva: Unknown.
Distribution: Brazil: Goiás, Mato Grosso.
Bionomics: Simulium cuasiexiguum breeds in fast flowing rivers, over aquatic vegeta-
tion, in grasslands and gallery forests. Females have zoophilic haematophagous habits.
Discussion: As was shown in the description, S. cuasiexiguum is very similar to S.
exiguum, S. paraguayense and S. gonzalezi. The pupal gills with 8 branches allow S.
cuasiexiguum to be separated from the first two species. The shorter distal process of the
paraproct and the subtriangular gonostylus differentiate S. gonzalezi from S. cuasiexiguum.

Simulium (Notolepria) subexiguum FIELD


Simulium (Notolepria) subexiguum FIELD, 1967: 189; IBÁÑEZ-BERNAL, 1992: 286.
Female: Similar to Simulium exiguum; differs in paraproct, which has very short distal
posterior process; also very similar to Simulium gonzalezi, S. llutense and Simulium sp. A.
Male: Similar to Simulium exiguum in external aspects; gonostylus and ventral plate
included in variations of S. exiguum.
Pupa: Gill with 8 branches, as in Simulium exiguum, S. paraguayense, S. llutense and
Simulium sp. A; without any difference from those species.
Larva: Unknown.
Distribution: Mexico: Chiapas; Panama: Canal Zone.
Bionomics: The species has been collected, according to the species author, on
“trailing grass”.
Discussion: The paraproct morphology of S. subexiguum allows it to be differentiat-
ed from S. exiguum. The male subtriangular gonostylus and pupal gills with 8 branches
separate it from S. gonzalezi.

Simulium (Notolepria) sp. A IBÁÑEZ-BERNAL


Simulium (Notolepria) sp. A IBÁÑEZ-BERNAL, 1992: 279.
IBÁÑEZ-BERNAL gave a detailed description of this species based on pharate adults. It
has similarities with S. cuasiexiguum, S. exiguum and S. subexiguum in the adults and also the
pupal gill with 8 branches. It can be differentiated from S. exiguum by the short paraproct
posterior process and other small anatomical differences, some of them included among
the variation in S. exiguum. Compared with S. gonzalezi, it shows differences in the
paraproct distal elongation, which is greater in Simulium sp. A, and in the pupal gill with 8
branches. It is practically the same as S. subexiguum. As explained by IBÁÑEZ-BERNAL, it will
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 235

be necessary to study more material of all stages. Maybe Simulium sp. A is a variation of S.
gonzalezi with 8 branches, as suggested by DALMAT (1955) in Guatemala for S. exiguum with
8 and 6 branches (IBÁÑEZ-BERNAL, 1992: 286). We have seen the same situation in Simulium
(Pternaspatha) annulatum with 6 and 8 gill branches.
Distribution: Mexico: Chiapas.
Bionomics: Simulium sp. A lives in the same places as S. gonzalezi.

Simulium (Notolepria) llutense COSCARÓN & MATTA (Fig. 56.)


Simulium (Notolepria) llutense COSCARÓN & MATTA, 1982: 49; COSCARÓN, 1987: 20; 1991: 150.
Female: Wing length 2.1-2.5 mm; head blackish, frons and clypeus dark brown;
antennae, palpi and proboscis dark grayish-brown with scape and pedicel brown;
scutum, scutellum black with abundant pilosity brownish or greenish-gray; with some
light incidence 1+1 anterior submedian subtriangular grayish spots can be observed,
changing light position these spots appear blackish. Metanotum velvety black; pleurae
blackish with gray pollinosity; wing veins yellowish brown with hair and setae dark
brown; legs yellowish brown with fore tarsomere and distal portion of median and
posterior tarsomeres blackish; abdomen matte black, with cereous sheen on VI-IX
tergites; fronto-ocular triangle slightly wider than long (Fig. A). Maxillary palp sensory
organ about half palpomere length; cibarium basally as in figure B; legs without
petaloid hair; calcipala well developed; claws without subbasal teeth; basal portion of
R and Sc without hair or setae; eighth sternite (Fig. C) with about 70 setae;
gonapophyses with internal border well sclerotized; genital fork with well-sclerotized
branches (Fig. D); paraproct with truncate distal border and short distal posterior
process (Fig. E).
Male: Wing length 1.8 mm; head and scutum black, the latter with golden hair, and
with some light with anterior fourth grayish; leg coloration as in female, but darker,
especially hind legs; pleurae blackish with gray pollinosity; abdomen black with 1+1 light
silvery spots on tergites II, VI-VIII; tergite IX grayish black; length/width ratio of hind
basitarsus = 7; gonostylus subtriangular, smoothly emarginated apically with apical spur
obsolescent (Fig. F); ventral plate shorter than wide at base (Fig. G); endoparameres with
strong hooks.
Pupa: Cocoon length at base 2.7-3.0 mm, at dorsum 1.8-2.5 mm; gills slipper shaped
with thick weave strong, projected ventrally (Fig. H); frontoclypeus (Fig. I) and exposed
area of thorax with regular quantity of smooth tubercles; frontoclypeus with 1+1 facial
and 2+2 frontal trichomes simple or bifid, thorax dorsally with 7 multirramous trichomes;
gill with 8 branches, primary branches with short basal trunk (Fig. J).
Larva: Maximum length 4.6-4.8 mm; color light grayish-green, more intense on
dorsum; head yellowish brown; cephalic apotome with isolated median, sublateral and
subbasal positive spots (Fig. K); antenna longer than stem of cephalic fan (Fig. L); ratio of
proximal, medial and distal segments = 1:1.2:1.1. Median tooth of hypostomium at same
level as lateral teeth; with 4 setae on each side (Fig. M); postgenal cleft deep; with
postgenal bridge shorter than half of hypostomium length. Mandible with 6 internal
teeth, second marginal teeth shorter than first tooth by half; cephalic fan with about 25
236 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

A
E

G
F

H
L

I
J

Figure 56. S. (Notolepria) llutense. A-E: Female; A: fronto-ocular triangle; B: cibarium, basal portion;
C: eighth sternite and gonapophysis; D: genital fork; E: cercus and paraproct. F-G: Male; F:
gonocoxite and gonostylus; G: ventral plate. H-J: Pupa; H: general aspect; I: frontoclypeus; J:
portion of thorax with trichomes and gill. K-M: Larva; K: head dorsal view; L: antenna; M:
hypostomium and postgenal cleft.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 237

rays; anal sclerite with some hair, scales not evident; anal ring with about 74 rows with 13-
14 hooks each; rectal papillae with 3 simple lobes.
Distribution: Chile: Tarapacá.
Bionomics: Simulium llutense breeds in small creeks of the Chilean desert area, which
flow over aquatic vegetation. The feeding habits of the females are unknown.
Discussion: Similarity in adult genitalia, cibarial shape, pupal gill and chaetotaxy, as
well as in the larvae, allow S. llutense to be included in Notolepria, but the species shows
some differences from other species of this subgenus, such as the absence of grouped
hairs simulating scales on the scutum, 1+1 subtriangles for the anterior silvery spots on
the female scutum; a small fronto-ocular triangle in the female and shorter ventral plate
paraprocts, which are very similar to those of S. gonzalezi.
Simulium spinifer KNAB from Peru, of uncertain position, shows some similarities
in the female description, but the male genitalia, as well as the other stages, do not
allow us to decide if they are the same species; therefore, we consider S. spinifer
species inquirenda.

Species inquirendae

Simulium (Notolepria) blantoni FIELD


Simulium (Notolepria) blantoni FIELD, 1967a: 192.
This species is known only from a female that is very close to Simulium gonzalezi and
S. subexiguum. The absence of a male and pupa do not allow it to be adequately
differentiated from the two previously mentioned species.
Distribution: Panama: Bocas del Toro.

Simulium incertum LUTZ


Simulium incertum LUTZ, 1910: 259; PINTO, 1931: 755.
LUTZ described this species without illustrations. It is not easy to identify, and he
says at the end of his description that it might be S. paraguayense SCHROTTKY. PINTO
(1931) published a pupal gill (Fig. 8 j) from the type area, and considered it to be that
of S. incertum, but it is not similar to the PINTO description of S. paraguayense that
agrees with our interpretation of this species. It is not possible to be sure what S.
incertum might be.
Distribution: Brazil: Minas Gerais, São Paulo.

Simulium paranense SCHROTTKY


Simulium paranense SCHROTTKY, 1909: 63; VULCANO, 1967: 18.
Simulium (Notolepria) paranense COSCARÓN, 1987: 20; CROSSKEY & HOWARD, 1997: 53.
The description is not sufficient to permit identification of this species.
Distribution: Paraguay: Alto Paraná.
238 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Simulium (Chirostilbia) ENDERLEIN (Figs. 57-63, 140.)


Type-species Chirostilbia flavifemur ENDERLEIN, 1921b: 79 = Simulium pertinax KOLLAR, 1832: 117.
Female: Species of medium size, wing length 2.4-3.6 mm; general coloration
blackish with soft pollinosity gray blackish, but can be black-grayish to greenish yellow;
some illumination shows 1+1 silvery submedian vittae changing to black with other
lighting; scutellum brownish, metanotum blackish; abdomen blackish to grayish green to
brown yellowish; frons medium wide; fronto-ocular triangle variable from deep to flat;
cibarium basally with median area smooth and cornuae branches well sclerotized; wing
generally with hair on basal sector of R; claws with or without basal tooth; gonapophysis
subtriangular, nearly two times longer than wide at base; genital fork with anterior branch
about one quarter longer than each posterior branch.
Male: Wing length 2.2-3.5; scutum black to brown yellowish or reddish, bordered
laterally and posteriorly by silver band, enlarged on pronotal angles; abdomen blackish to
gray greenish with 1+1 grayish lateral spots on tergites II, V-VIII; gonocoxite subtrapezoi-
dal to subquadrate, longer than gonostylus; gonostylus subtriangular to subtrapezoidal,
curved, acuminate distally, without apical spur, or very reduced (like small seta); ventral
plate wider than long, with slight curvature in lateral view and not acuminate apically.
Median sclerite wide with strong median concavity; parameres with robust hooks.
Pupa: Cocoon slipper to shoe shaped, compact with resistant weave, evident threads
compact or with apertures and festoons; frontoclypeus and exposed portion of thorax
with abundant to no tubercles; head with 2+2 frontal and 1+1 facial trichomes; thorax
dorsally with 5-6 trichomes on each side; gills each with 8-10 to 120 branches, sometimes
relatively thick; terminal spines relatively short.
Larva: Body cuticle without hair or scales; cephalic apotome variable, generally with
subtriangular darker medium basal area with median longitudinal, submedian and
subbasal spots positive or negative; antenna proximal segment shorter than medial
segment and this shorter than distal. Mandible with 8-9 internal teeth; second marginal
tooth generally about half of first; cephalic fan with about 45 rays; postgenal cleft deep
with postgenal bridge about half of hypostomium length; lateral sclerite of thoracic
proleg with about 25 teeth, arranged in groups; anal sclerite with single hairs and
sometimes with scales; anal ring with 65-190 rows with 12-24 hooks each; rectal papillae
with 4-24 diverticula on each lobe.
Distribution: Simulium (Chirostilbia) is a South American taxon, distributed predom-
inantly in the intertropical non-Andean region (Fig. 140).
Discussion: This subgenus is composed of two species groups:
· PERTINAX species group: imagoes blackish; female claw with subbasal tooth;
male gonostylus subquadrate, about as long as wide at base and without median
distal diagonal crest; ventral plate about as long as wide, generally curved latero-
apically; pupa generally with abundant tubercles on frontoclypeus and thorax, and
cocoon slipper to shoe shaped; larval cephalic apotome basally with darker
subtriangular.
· SUBPALLIDUM species group: imagoes brown grayish to green yellowish
especially females; female claw generally without subbasal tooth; male gonostylus
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 239

subtriangular, longer than wide at base with median distal diagonal crest; ventral
plate wider than long not curved apically; pupal frontoclypeus and thorax without
tubercles or these very scarce; cocoon slipper shaped; larval cephalic apotome
without basally darker subtriangular area.

KEYS TO SIMULIUM (CHIROSTILBIA) SPECIES

Females
Not included: Simulium friedlanderi (unknown female) and S. empascae (scarce information).

1 General coloration blackish; scutum and abdomen black to dark reddish brown;
claw with subbasal tooth (Fig. 57 M) ...................... PERTINAX species group......2
– General coloration grayish brown to yellowish green; scutum yellowish brown to
lead gray; abdomen yellowish, grayish brown to greenish gray; claw without
subbasal tooth (Fig. 60 O) ................................ SUBPALLIDUM species group......9
2(1) Scutum blackish, without submedian silvery vittae joined to posterior grayish area
.................................................................................................................... Simulium pertinax
– Scutum blackish to reddish brown, with 1+1 submedian silvery anterior
subrectangular spots or vittae joining to posterior grayish area in some incident
light (Figs. 57 F, G, J, O) ................................................................................................... 3
3(2) Scutum with 1+1 submedian silvery anterior subrectangular small spots ...............
............................................................................................................. Simulium bifenestratum
– Scutum with 1+1 submedian silvery vittae joining posterior grayish area (Figs. 57
F, G, J, O) ............................................................................................................................. 4
4(3) Scutum blackish; wing length 2.0-3.0 mm .................................................................... 5
– Scutum dark reddish brown to light yellowish brown; wing length 2.5-3.6 mm
................................................................................................................................................ 7
5(4) Sc without hair; wing length 2.0-2.2 mm; scutum with pile silvery ..........................
.....................................................................................................................Simulium serranus
– Sc with hair; wing length 2.2-3.0 mm; scutum with pile golden or with lead gray
tones ...................................................................................................................................... 6
6(5) Scutum with abundant pile, with lead gray tones ................. Simulium spinibranchium
– Scutum with scarce pile, with golden tones ................................... Simulium laneportoi
7(4) Scutum light yellowish brown, with light grayish to yellow pile ................................
...............................................................................................................Simulium riograndense
– Scutum dark grayish brown or reddish brown, with golden to silvery pile .......... 8
8(7) Cercus with curved distal border; paraproct about 1.5 times longer than wide at
base (Fig. 57 N); scutum dark grayish brown ............................... Simulium distinctum
– Cercus with straight distal border; paraproct about 2.5 times longer than wide at
base (Fig. 57 U); scutum dark reddish brown .................................... Simulium obesum
9(1) Scutum light yellowish brown with light brown posterior area; pleurae yellowish
brown .................................................................................................. Simulium subpallidum
– Scutum dark grayish leaden, with blackish brown posterior area; pleurae dark
grayish ................................................................................................................................. 10
240 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

10(9) Wings with hair on basal sector of R; abdomen greenish with dark brown tergal
plates (Fig. 60 E) ................................................................................. Simulium acarayense
– Wings without hair on basal sector of R; abdomen brownish with dark grayish
brown tergal plates ............................................................................... Simulium papaveroi

Males
Simulium empascae and S. friedlanderi are not included due to lack of information.

1 Gonostylus subquadrate, about as long as wide at base and without median distal
diagonal crest (Figs. 58 B, F, G, I, J, L, O); ventral plate about as long as wide,
generally curved apicolaterally (Figs. 58 A, C, H, K, M, P) ........................................
......................................................................................... PERTINAX species group......2
– Gonostylus subtriangular, longer than wide at base, with median distal diagonal
crest (Figs. 61 D, F, H, I) ventral plate wider than long, not curved apically (Figs.
61 E, J, K) ............................................................ SUBPALLIDUM species group......5
2(1) Scutum dark grayish brown ............................................................................................. 3
– Scutum black ....................................................................................................................... 4
3(2) Wing length 2.4-3.3 mm; scutum blackish; gonostylus with relatively short process
on external distal side (Fig. 58 J); ventral plate relatively elongated (Fig. 58 K) ....
................................................................................................................. Simulium distinctum
– Wing length 3.3-3.5 mm; scutum reddish brown to yellowish; gonostylus with
well-developed process on external distal side (Fig. 58 L), ventral plate relatively
short (Fig. 58 M) ...............................................................................Simulium riograndense
4(2) Wing length 3.2-3.5 mm; gonostylus with lateral external and posterior sides
concave (Fig. 58 O) .......................................... Simulium obesum, Simulium bifenestratum
– Wing length 2.0-3.1 mm, gonostylus with lateral external and posterior sides from
convex to straight (Figs. 58 B, F, G) ................................................................................
....... Simulium pertinax, Simulium laneportoi, Simulium serranus, Simulium spinibranchium
5(1) Scutum yellowish brown to reddish brown ................................ Simulium subpallidum
– Scutum black ....................................................................................................................... 6
6(5) Wings with hair on basal sector of R; gonostylus only a little longer than wide at
base (Fig. 61 D) ................................................................................... Simulium acarayense
– Wings without hair on basal sector of R; gonostylus nearly 2 times longer than
wide at base (Fig. 61 F) ....................................................................... Simulium papaveroi

Pupae
1 Gill with 8 filaments; cocoon generally not covering gill branches and with anterior
portion smooth, without festoons (Figs. 59 A, D, G, J; 62 A, C, D, G) ....................... 2
– Gill with 10-120 filaments; cocoon covering gill and anterior portion with
festoons (Figs. 59 L, O, S) ................................................................................................ 8
2(1) Cocoon covering branches (Fig. 59 J); primary and secondary branches emerging
as a bundle (Fig. 59 K) ......................................................................... Simulium empascae
– Cocoon not covering gill (Figs. 59 A, D, G; 62 A, C, D, G); primary and secondary
branches emerging isolated, not as bundle (Figs. 59 C, F, H; 62 B, F, I) ............... 3
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 241

3(2) Gill filaments thick basally (Fig. 59 H) ................................... Simulium spinibranchium


– Gill filaments not thick basally ........................................................................................ 4
4(3) Frontoclypeus and exposed portion of thorax with abundant tubercles (Figs. 59 B,
E) ........................................................................................................................................... 5
– Frontoclypeus and exposed portion of thorax without or with few tubercles (Figs.
62 E, H) ................................................................................................................................ 6
5(4) Gill length about as long as pupal body length (Fig. 59 D) and arranged in 1 plane
(Fig. 59 F); frontoclypeus and thorax with only moderate abundance of tubercles
(Fig. 59 E) ................................................................................................ Simulium serranus
– Gill length about half of pupal body length (Fig. 59 A), and arranged in more
than 1 plane (Fig. 59 C); frontoclypeus and thorax with very abundant tubercles
(Fig. 59 B) .............................................................. Simulium pertinax, Simulium laneportoi
6(4) Gill branches slightly grouped basally, especially dorsal branches (Fig. 62 I);
primary and secondary dorsal branches thicker than ventral branches; cephalic
and thoracic trichomes simple or bifid (Fig. 62 H) ...................... Simulium papaveroi
– Gill branches open (Figs. 62 B, F); primary and secondary dorsal branches no
thicker than ventral branches; cephalic and thoracic trichomes with 3-6 branches
(Fig. 62 E) ............................................................................................................................ 7
7(6) Gill filaments approximately straight and disposed in 1 plane (Figs. 62 C, D, F);
sometimes rounded tubercles present on frontoclypeus and thorax dorsally (Fig.
62 E) ...................................................................................................... Simulium acarayense
– Gill filaments with slight curvature and arranged like basket (Figs. 62 A, B);
without tubercles on frontoclypeus or thorax, or if present, with conical shape
.............................................................................................................. Simulium subpallidum
8(1) Gill with about 120 branches; thorax covered totally with trichomes (Fig. 59 U)
...................................................................................................................... Simulium obesum
– Gill with 10-11 branches; thorax with 5-6 trichomes on each side ......................... 9
9(8) Gill with 11 branches very thick and with strong curvature (Fig. 59 R) ..................
................................................................................................................ Simulium friedlanderi
– Gill with 10 branches filamentous and without strong curvature (Figs. 59 N, Q)
.............................................................................................................................................. 10
10(9) Cocoon with 1+1 anterolateral apertures; frontoclypeal margin elevated as a
crown ................................................................................................. Simulium bifenestratum
– Cocoon without 1+1 anterolateral apertures (Figs. 59 L, O); frontoclypeal margin
not elevated ....................................................................................................................... 11
11(10) Frontoclypeus and thorax with regular quantity of tubercles uniformly distributed
(Fig. 59 M); cocoon length basally 3.0 mm, dorsally 2.8 mm, gills 1.0 mm ............
................................................................................................................. Simulium distinctum
– Frontoclypeus and thorax with scarce quantity of tubercles, distributed on
frontoclypeus near facial trichomes (Fig. 59 P) and on exposed portion of thorax;
cocoon length basally 4.2-4.7 mm; dorsally 4.5-4.7 mm; gills 1.7-2.0 mm ..............
...............................................................................................................Simulium riograndense
242 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Larvae
Simulium obesum larva not included (unknown).

1 Postgenal cleft subtriangular (Fig. 63 F) to dome shaped (Fig. 63 C); anal ring with
85-204 rows of hooks; maximum length 6.0-11.8 mm ............................................. 2
– Postgenal cleft subovoidal (Fig. 63 Q); anal ring with 60-81 rows of hooks;
maximum length 5.0-5.8 mm .............................................................................................
......................................... Simulium subpallidum, Simulium acarayense, Simulium papaveroi
2(1) Anal ring with 145-204 rows of hooks; maximum length 8.1-11.8 mm ................ 3
– Anal ring with 76-122 rows of hooks; maximum length 5.6-7.5 mm .................... 6
3(2) Anal ring with 201-204 rows of hooks; rectal papillae with 4-6 diverticula on each
lobe ........................................................................................................... Simulium empascae
– Anal ring with 145-190 rows of hooks; rectal papillae with 7-26 diverticula on
each lobe .............................................................................................................................. 4
4(3) Postgenal cleft dome shaped; cephalic apotome darkened totally on basal 1/3 and
continued anteriorly as triangle nearly reaching frontal area ...... Simulium bifenestratum
– Postgenal cleft subtriangular with blunt apex; cephalic apotome darkened only on
basal 1/3 .............................................................................................................................. 5
5(4) Anal ring with 170-190 rows of hooks; maximum length 8.5-9.0 mm ....................
................................................................................................................. Simulium distinctum
– Anal ring with 145-152 rows of hooks; maximum length 9.5-10.5 mm ..................
...............................................................................................................Simulium riograndense
6(2) Cephalic apotome darkened totally on basal 1/3 and continued anteriorly as
triangle nearly reaching frontal area (Fig. 63 A) .......................................................... 7
– Cephalic apotome darkened only on middle of base (Fig. 63 D) or basal 1/4 and
slightly continued on midlongitudinal area ................................................................... 8
7(6) Postgenal cleft dome shaped (Fig. 63C); anal ring with 88-100 rows of hooks
.................................................................................................................... Simulium pertinax
– Postgenal cleft subtriangular; anal ring with 76-86 rows of hooks ...........................
..........................................................................................................Simulium spinibranchium
8(6) Cephalic apotome darkened on basal 1/4 and slightly continued on
midlongitudinal area, median longitudinal and submedian spots positive...............
.................................................................................................................. Simulium laneportoi
– Cephalic apotome darkened only at mid base, median longitudinal and submedian
spots negative (Fig. 63 D) ................................................................................................ 9
9(8) Anal ring with 85-88 rows of hooks; rectal papillae with 15-18 diverticula on each
lobe .............................................................................................................Simulium serranus
– Anal ring with 105-122 rows of hooks; rectal papillae with 20-24 diverticula on
each lobe .............................................................................................. Simulium friedlanderi
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 243

CLAVES DE ESPECIES DE SIMULIUM (CHIROSTILBIA)


Hembras
Simulium friedlanderi (hembra desconocida) y S. empascae (escasa información) no se incluyen.

1 Coloración general negruzca; escudo y abdomen negro a castaño rojizo oscuro;


uña con diente subbasal (Fig. 57 M) .............................. PERTINAX sp. grupo......2
– Coloración general castaño grisáceo a verde amarillento; escudo castaño
amarillento a gris plomizo; abdomen desde amarillo, castaño grisáceo a gris
verdoso; uña sin diente subbasal (Fig. 60 O) ...........................................................
........................................................................................SUBPALLIDUM sp. grupo......9
2(1) Escudo negruzco, no mostrando con ninguna luz bandas plateadas submedianas
llegando al área posterior grisácea ...................................................... Simulium pertinax
– Escudo negruzco a castaño rojizo presentando con alguna luz 1+1 manchas
plateadas submedianas subrectangulares o bandas llegando al área posterior
grisácea (Figs. 57 F, G, J, O) ............................................................................................ 3
3(2) Escudo con 1+1 mancha plateada subrectangular pequeña anterior submediana
............................................................................................................. Simulium bifenestratum
– Escudo con 1+1 banda plateada submediana, uniéndose al área grisácea posterior
(Figs. 57 F, G, J, O) ............................................................................................................ 4
4(3) Escudo negruzco; longitud alas 2,0-3,0 mm ................................................................ 5
– Escudo castaño rojizo oscuro a castaño amarillento claro; longitud de las alas 2,5-
3,6 mm .................................................................................................................................. 7
5(4) Sc sin pelos; longitud ala 2,0-2,2 mm; pilosidad del escudo plateada ......................
.....................................................................................................................Simulium serranus
– Sc con pelos; longitud ala 2,2-3,0 mm; pilosidad del escudo dorado a gris plomizo
................................................................................................................................................ 6
6(5) Escudo con pilosidad abundante con tonalidad gris plomizo ...................................
..........................................................................................................Simulium spinibranchium
– Escudo con pilosidad relativamente escasa, con tonalidad dorada ...........................
.................................................................................................................. Simulium laneportoi
7(4) Escudo castaño amarillento claro, con pilosidad grisácea clara a amarillenta ........
...............................................................................................................Simulium riograndense
– Escudo castaño grisáceo oscuro o castaño rojizo con pilosidad dorada a plateada
................................................................................................................................................ 8
8(7) Cerco con borde distal curvo; paraprocto aproximadamente 1,5 veces más largo
que ancho en la base (Fig. 57 N); escudo castaño grisáceo oscuro ..........................
................................................................................................................. Simulium distinctum
– Cerco con borde distal recto; paraprocto aproximadamente 2,5 veces más largo que
ancho en la base (Fig. 57 U); escudo castaño rojizo oscuro .............. Simulium obesum
9(1) Escudo castaño amarillento claro, con fondo castaño claro; pleuras castaño
amarillento ......................................................................................... Simulium subpallidum
– Escudo gris plomizo oscuro con fondo castaño negruzco; pleuras grisáceo oscuro
.............................................................................................................................................. 10
244 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

10(9) Alas con pelos en sector basal de R; abdomen verdoso con placas tergales castaño
oscuro (Fig. 60 E) ............................................................................... Simulium acarayense
– Alas sin pelos en sector basal de R; abdomen castaño con placas tergales castaño
grisáceo oscuro ..................................................................................... Simulium papaveroi

Machos
Simulium empascae y S. friedlanderi no se incluyen por escasa información.

1 Gonostilo subcuadrado, aproximadamente tan largo como ancho en la base y sin


cresta mediana distal (Figs. 58 B, F, G, I, J, L, O); placa ventral aproximadamente
tan larga como ancha, generalmente curvada latero apicalmente (Figs. 58 A, C, H,
K, M, P) ................................................................................ PERTINAX sp. grupo......2
– Gonostilo subtriangular, más largo que ancho en la base, con una cresta medio
longitudinal (Figs. 61 D, F, H, I); placa ventral más ancha que larga, no curvada
apicalmente (Figs. 61 E, J, K) ..................................SUBPALLIDUM sp. grupo......5
2(1) Escudo castaño grisáceo oscuro ..................................................................................... 3
– Escudo negro ...................................................................................................................... 4
3(2) Longitud ala 2,4-3,3 mm; escudo negruzco; gonostilo con relativamente corta
prolongación sobre el costado distal externo (Fig. 58 J); placa ventral
relativamente alargada (Fig. 58 K) ................................................... Simulium distinctum
– Longitud ala 3,3-3,5 mm; escudo castaño rojizo a amarillento; gonostilo con una
prolongación bien desarrollada sobre el costado distal externo (Fig. 58 L); placa
ventral relativamente corta (Fig. 58 M) ........................................Simulium riograndense
4(2) Longitud ala 3,2-3,5 mm; gonostilo con el costado lateral externo y posterior
cóncavos (Fig. 58 O) ....................................... Simulium obesum, Simulium bifenestratum
– Longitud ala 2,0-3,1 mm; gonostilo con el costado lateral externo y posterior
desde convexo a recto (Figs. 58 B, F, G) ........................................................................
....... Simulium pertinax, Simulium laneportoi, Simulium serranus, Simulium spinibranchium
5(1) Escudo castaño amarillento a castaño rojizo ............................. Simulium subpallidum
– Escudo negro ...................................................................................................................... 6
6(5) Alas con pelos en sector basal de R; gonostilo solamente un poco más largo que
ancho en la base (Fig. 61 D) ............................................................. Simulium acarayense
– Alas sin pelos en sector basal de R; gonostilo cerca 2 veces más largo que ancho
en la base (Fig. 61 F) ........................................................................... Simulium papaveroi

Pupas
1 Branquias con 8 filamentos; capullo generalmente no cubriendo las ramas de las
branquias y con porción anterior lisa y sin festones (Figs. 59 A, D, G, J; 62 A, C, D,
G) ........................................................................................................................................... 2
– Branquias con 10-120 filamentos; capullo cubriendo las branquias; porción
anterior con festones (Figs. 59 L, O, S) ......................................................................... 8
2(1) Capullo cubriendo las ramas (Fig. 59 J); ramas primarias y secundarias emergiendo
agrupadas como un ramo (Fig. 59 K) ............................................... Simulium empascae
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 245

– Capullo no cubriendo la branquia (Figs. 59 A, D, G; 62 A, C, D, G); ramas


primarias y secundarias emergiendo aisladas no como un ramo (Figs. 59 C, F, H;
62 B, F, I) ............................................................................................................................. 3
3(2) Filamentos branquiales gruesos basalmente (Fig. 59 H) ........ Simulium spinibranchium
– Filamentos branquiales no gruesos basalmente .......................................................... 4
4(3) Frontoclípeo y porción expuesta del tórax con abundantes tubérculos (Figs. 59 B, E)
................................................................................................................................................ 5
– Frontoclípeo y porción expuesta del tórax sin o con escasos tubérculos (Figs. 62
E, H) ..................................................................................................................................... 6
5(4) Largo de la branquia aproximadamente como el largo del cuerpo (Fig. 59 D) y
ordenadas en un plano (Fig. 59 F); frontoclípeo y tórax con solamente una
moderada abundancia de tubérculos (Fig. 59 E) ..............................Simulium serranus
– Largo de las branquias aproximadamente la mitad del largo del cuerpo (Fig. 59 A)
y ordenadas en más de un plano (Fig. 59 C); frontoclípeo y tórax con muy
abundantes tubérculos (Fig. 59 B) ...................... Simulium pertina, Simulium laneportoi
6(4) Ramas de las branquias relativamente agrupadas basalmente, en especial las
ramas dorsales (Fig. 62 I); ramas primaria y secundaria dorsal más gruesas que la
ventral; tricomas cefálicos y torácicos simples o bífidos (Fig. 62 H) ....................
.................................................................................................................. Simulium papaveroi
– Ramas de la branquia abiertas (Figs. 62 B, F); ramas primaria y secundaria dorsal
no más gruesas que la ventral; tricomas cefálicos y torácicos con 3-6 ramas (Fig.
62 E) ..................................................................................................................................... 7
7(6) Filamentos de la branquia aproximadamente rectos y dispuestos en un plano
(Figs. 62 C, D, F); tubérculos redondeados a veces presentes en frontoclípeo y
dorsalmente en tórax (Fig. 62 E) ..................................................... Simulium acarayense
– Filamentos de la branquia con una pequeña curvatura y dispuestos en conjunto
como un cesto (Figs. 62 A, B); sin tubérculos en frontoclípeo y tórax, si existe
alguno tiene forma de cono ............................................................ Simulium subpalidum
8(1) Branquia con alrededor de 120 ramas; tórax totalmente cubierto con tricomas
(Fig. 59 U) .................................................................................................. Simulium obesum
– Branquia con 10-11 ramas; tórax con tricomas de 5-6 ramas sobre cada costado
................................................................................................................................................ 9
9(8) Branquia con 11 ramas muy gruesas y con fuerte curvatura (Fig. 59 R) .................
................................................................................................................ Simulium friedlanderi
– Branquia con 10 ramas filamentosas y sin fuerte curvatura (Figs. 59 N, Q) ...... 10
10(9) Capullo con 1+1 agujeros anteriores; áreas marginales del frontoclípeo elevadas
semejando una corona ................................................................... Simulium bifenestratum
– Capullo sin 1+1 agujeros anteriores (Figs. 59 L-O); área marginal del frontoclípeo
no elevada .......................................................................................................................... 11
11(1) Frontoclípeo y tórax con regular cantidad de tubérculos uniformemente
distribuidos (Fig. 59 M); largo del capullo en la base 3,0 mm, en el dorso 2,8 mm,
branquias 1,0 mm ................................................................................ Simulium distinctum
– Frontoclípeo y tórax con escasa cantidad de tubérculos distribuidos en
frontoclípeo cercanos a los tricomas faciales (Fig. 59 P) y en la porción expuesta
246 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

del tórax; largo del capullo en la base 4,2-4,7 mm, en el dorso 4,5-4,7 mm,
branquias 1,7-2,0 mm .......................................................................Simulium riograndense

Larvas
La larva de Simulium obesum desconocida.

1 Hendidura postgenal subovoidal (Fig. 63 Q); anillo anal con 60-81 hileras de
ganchos; longitud máxima 5,5-8,0 mm ............................................................................
......................................... Simulium subpallidum, Simulium acarayense, Simulium papaveroi
– Hendidura postgenal subtriangular (Fig. 63 F), con forma de domo (Fig. 63 C);
anillo anal con 85-204 hileras de ganchos; longitud máxima 6,0-11,8 mm ........... 2
2(1) Anillo anal con 145-204 hileras de ganchos; longitud máxima 8,1-11,8 mm ....... 3
– Anillo anal con 76-122 hileras de ganchos; longitud máxima 5,6-7,5 mm ............ 6
3(2) Anillo anal con 201-204 hileras de ganchos; papilas rectales con 4-6 divertículos
sobre cada lóbulo .................................................................................. Simulium empascae
– Anillo anal con 145-190 hileras de ganchos; papilas rectales con 7-26 divertículos
en cada lóbulo ..................................................................................................................... 4
4(3) Hendidura postgenal en forma de domo; apotoma cefálico oscurecido totalmente
sobre 1/3 basal y continuando adelante como un triángulo llegando cerca del área
frontal ................................................................................................ Simulium bifenestratum
– Hendidura postgenal subtriangular pero con ápice romo; apotoma cefálico
oscurecido solo en el 1/3 basal ....................................................................................... 5
5(4) Anillo anal con 170-190 hileras de ganchos; longitud máxima 8,5-9,0 mm ............
................................................................................................................. Simulium distinctum
– Anillo anal con 145-152 hileras de ganchos; longitud máxima 9,5-10,5 mm .........
...............................................................................................................Simulium riograndense
6(2) Apotoma cefálico oscurecido totalmente sobre 1/3 basal y continuando hacia
delante como un triángulo, llegando cerca del área frontal (Fig. 63 A) ................. 7
– Apotoma cefálico oscurecido solo en la mitad de la base (Fig. 63 D) o en 1/4 basal
y continuando suavemente sobre el área medio longitudinal ................................... 8
7(6) Hendidura postgenal en forma de domo (Fig. 63 C); anillo anal con 88-100 hileras
de ganchos ............................................................................................... Simulium pertinax
– Hendidura postgenal subtriangular; anillo anal con 76-86 hileras de ganchos .......
..........................................................................................................Simulium spinibranchium
8(6) Apotoma cefálico oscurecido en el 1/4 basal y suavemente continuado sobre el
área mediana longitudinal; manchas positivas medio longitudinales y submedianas
.................................................................................................................. Simulium laneportoi
– Apotoma cefálico oscurecido solo en la mitad basal; manchas negativas medio
longitudinales y submedianas (Fig. 63 D) ..................................................................... 9
9(8) Anillo anal con 85-88 hileras de ganchos; papilas rectales con 15-18 divertículos
en cada lóbulo ..........................................................................................Simulium serranus
– Anillo anal con 105-122 hileras de ganchos; papilas rectales con 20-24 divertículos
en cada lóbulo ..................................................................................... Simulium friedlanderi
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 247

PERTINAX species group

Simulium (Chirostilbia) pertinax KOLLAR (Figs. 57 A-E; 58 A-C; 59 A-C; 63 A-C.)


Simulium pertinax KOLLAR, 1832: 117; LUTZ, 1910: 228-230, 266; LUTZ et al., 1918: 107-109;
D’ANDRETTA & D’ANDRETTA, 1950: 193-213; SHELLEY et al., 1984: 149.
Simulium (Chirostilbia) pertinax COSCARÓN, 1987: 26; 1991: 215-217; GIL ACEVEDO et al., 2004: 433-437.
Simulium pertinax PINTO, 1931: 678-680, 722-724, 725, 727, 742, 743, 760; LANE & PORTO, 1939: 169-
171; LANE & VULCANO, 1943: 432.
Trichodagmia pertinax ENDERLEIN, 1936: 128.
Simulium infuscatum LUTZ, 1910: 26-29.
Simulium venustum (nec SAY, 1922): LUTZ, 1909: 132, 136-138.
Chirostilbia flavifemur ENDERLEIN, 1921b: 79.
Trichodagmia lutziana ENDERLEIN, 1934b: 291; 1935: 195.
Simulium distinctum (nec LUTZ, 1910): PINTO, 1931: part: 670, 669-701, part: 724.
Simulium inexorabile SCHROTTKY, 1909: 63; WYGODZINSKY, 1953a: 302.
Simulium septentrionale CERQUEIRA & BARBOSA DE ALMEIDA, 1970: 1-11.
Simulium cerqueirai BARBOSA DE ALMEIDA, 1979: 789.
Chirostilbia pertinax PY-DANIEL & MOREIRA SAMPAIO, 1995: 118; STRIEDER, 2002: 133-135; 2004: 126-127.
Female: Wing length 2.2-2.8 mm; general coloration blackish with soft gray
pollinosity and gray to golden pile; with some frontal illumination, black with median wide
stripe; changing light to latero-posterior position shows grayish with 1+1 black submedian
stripes nearly reaching blackish posterior border, similar as shown for laneportoi but wider
and without contrast with gray area; in specimens preserved for long time, coloration is
dark grayish brown homogeneous and without evident vittae; scutellum dark brown gray;
metanotum velvety black with gray pollinosity; pleurae brown to violaceous with silvery
pruinosity; legs brown blackish on third distal area of tibia and basitarsus of hind leg and
tarsomeres of all legs; abdomen black with shiny tergal plates V-IX; cibarium with median
area light and smooth (Fig. 57 A); fronto-ocular triangle deep (Fig. 57 B); basal sector of
R with hair arranged in 2-3 rows, and Sc with 18 setae; claw with well developed subbasal
tooth; eighth sternite well sclerotized, with about 20 setae on each side (Fig. 57 C)
Gonapophysis subtriangular, divergent with rounded apex and slightly sclerotized inter-
nally; paraproct about 2.5 times longer than wide at base and without basal lobe (Fig. 57
D); anterior branch of genital fork with strong curvature (Fig. 57 E).
Male: Wing length 2.2-2.5 mm; scutum black with golden pile and gray pollinosity
showing with some light, a wide and lighter lateral stripe joined anteriorly with 1+1
fronto-lateral spots and posteriorly with gray prescutellar area, similar as shown for
Simulium serranus (Fig. 58 E); frontolateral spots with antero-lateral illumination are
continuous anteriorly as in Simulium laneportoi (Fig. 58 D); scutellum and metanotum black;
abdomen black with 1+1 silvery pruinose spots on tergites II, V-VII and in some
specimens on IX; gonostylus about 3/4 of gonocoxite length with curved apex (Figs. 58
A, B); ventral plate about as long as wide at base apically slightly acuminate and curved
dorsally (Figs. 58 A, C); median sclerite with slight median cleft and endoparameres with
strong hooks (Fig. 58 A).
248 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

M H

O
F G J
B A

E I
C

D
R T

L
V P

K
Figure 57. Female. A-E: S. (Chirostilbia) pertinax; A: cibarium, B: fronto-ocular triangle; C: genitalia
ventral view showing eighth sternite, gonapophysis, cercus, paraproct, genital fork and
spermatheca; D: cercus and paraproct mounted on slide; E: genital fork. F-G: S. (C.) laneportoi; F:
scutum with frontal illumination; G: scutum with posterior illumination. H-I: S. (C.) serranus; H:
frons; I: cercus and paraproct. J-L: S. (C.) spinibranchium; J: scutum with frontal illumination; K:
eighth sternite and gonapophysis; L: cercus and paraproct. M-N: S. (C.) distinctum; M: claw; N:
cercus and paraproct. O-V: S. (C.) obesum; O: scutum with frontal illumination, P: sensory vesicle of
maxillary palp and palpomere; Q-S: anterior, median and hind leg, respectively; T: eighth sternite
and gonapophysis; U: cercus and paraproct; V: genital fork.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 249

Pupa: Cocoon slipper shaped, weave resistant with visible threads and reinforced
anterior border (Fig. 59 A), length at base 4.0-4.5 mm, at dorsum 3.0-3.5 mm, gills 1.5-2.0
mm; frontoclypeus and exposed portion of thorax with numerous tubercles (Fig. 59 B),
cephalic trichomes with 1-3 branches, thorax with 4-6 branches; gill with 8 filamentous
branches, with short primary and secondary branches (Fig. 59 C).
Larva: Maximum length 6.5-7.0 mm; body without hair or spines; cephalic apotome
with dark subtriangular basal spot reaching half of length (Fig. 63 A); ratio of proximal,
medial and distal antennal segments = 1:1.2:1.4 (Fig. 63 B); cephalic fan with about 40
rays; postgenal cleft dome shaped (Fig. 63 C); postgenal bridge a little longer than half of
hypostomium length; anal ring with about 88-100 rows of 15-21 hooks each; rectal
papillae with 15 diverticula on central lobe and 12 on each lateral lobe.
Distribution: Argentina: Corrientes, Entre Rios, Misiones; Brazil: Bahia, Mato
Grosso, Mato Grosso do Sul, Minas Gerais, Paraná, Rio de Janeiro, Rio Grande do Sul,
Rondônia, Santa Catarina, São Paulo; Paraguay.
Bionomics: Simulium pertinax breeds in creeks with fast current and clear water.
Larvae attach to aquatic plants, trees, submerged branches and leaves. Females are very
anthropophilic and the bite is uncomfortable. This species is the most dangerous in
southern Brazil and northeastern Argentina and Paraguay.
Discussion: Simulium pertinax is a very peculiar species and easy to identify. The closest
species is S. laneportoi with a very similar pupa, having gill branches only a little thinner. The
female of S. laneportoi has a scutum with narrower black submedian stripes, and when seen
with posterior light, the 1+1 grayish submedian vittae are more evident than in S. pertinax.

Simulium (Chirostilbia) laneportoi VARGAS (Figs. 57 F, G; 58 D.)


Simulium laneportoi VARGAS, 1941a: 118 (subst. name).
Simulium pilosum LANE & PORTO, 1940: 190 (preocup).
Eusimulium laneportoi LANE & VULCANO, 1943: 437.
Simulium (Chirostilbia) laneportoi PY-DANIEL, 1987: 336-341; COSCARÓN, 1987: 26, 1991: 217-219.
Simulium (Chirostilbia) dekeyseri SHELLEY & PY-DANIEL, 1981: 23-32.
Chirostilbia laneportoi PY-DANIEL & MOREIRA SAMPAIO, 1995: 118.
Female: Wing length 2.2-2.9 mm; general coloration black; scutum black with gray
pollinosity, pile relatively scarce with golden tones; with frontal light, median and 1+1
sublateral grayish areas appear on black scutum (Fig. 57 F); with posterior illumination,
coloration reverses (Fig. 57 G). Metanotum and abdomen black; abdominal tergite II with
1+1 lateral silvery spots and tergites V-VIII shiny; basal sector of R with hair and Sc with
10-20 setae; claw with large subbasal tooth; eighth sternite with 15-17 setae; gonapophysis
and paraprocts as in Simulium spinibranchium; genital fork as in S. pertinax.
Male: Wing length 2.1-2.7 mm; scutum velvety black with anterior lateral and
posterior border gray pollinose similar to Simulium serranus; with frontal light, anterior
border gray, wide and continuous (Fig. 58 D); gonocoxite, gonostylus and ventral plate as
in S. spinibranchium.
Pupa: Cocoon shape, numerous tubercles on frontoclypeus and thorax, and gill with
8 branches practically indistinguishable from those of Simulium pertinax, with only small
250 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

A
E

B H
C

K O

Figure 58. Male. A-C: S. (Chirostilbia) pertinax; A: genitalia ventral view showing gonocoxite,
gonostylus, ventral plate, endoparameres and median sclerite; B: gonocoxite and gonostylus in
ventral view; C: ventral plate. D: S. (C.) laneportoi, scutum with frontal illumination. E-F: S. (C.)
serranus; E: scutum with lateral view; F: gonocoxite and gonostylus. G-H: S. (C.) spinibranchium; G:
gonocoxite and gonostylus, dorsal view; H: ventral plate. I: S. (C.) empascae, gonocoxite and
gonostylus. J-K: S. (C.) distinctum; J: gonocoxite and gonostylus, K: ventral plate. L-N: S. (C.)
riograndense; L: gonocoxite and gonostylus; M: ventral plate; N: endoparamere and aedeagal
membrane. O-P: S. (C.) obesum; O: gonocoxite and gonostylus, ventral view; P: ventral plate.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 251

thin gill branches; size slightly smaller with cocoon length on base 2.4-3.9 mm, on dorsum
2.8-3.8 mm and gills 1.5-2.4 mm.
Larva: Maximum length 5.6-7.5 mm; cephalic apotome darkened on mid longitudi-
nal and 1/4 basal area, with discontinuous positive spots on median line and 1+1 on
submedian and subbasal areas; postgenal cleft subtriangular with blunt apex; postgenal
bridge a little shorter than hypostomium length; ratio of proximal, medial and distal
antennal segments = 1:1-1.3:1.3-1.7; cephalic fan with 38-45 rays; anal ring with 90-115
rows with 15-18 hooks each; rectal papillae with 19-30 diverticula on each lobe.
Distribution: Brazil: Goiás, Mato Grosso, Minas Gerais, Rio de Janeiro, São Paulo.
Bionomics: Simulium laneportoi breeds in fast-flowing creeks, where the larvae and
pupae attach themselves to stones and aquatic vegetation. Females bite humans.
Discussion: Simulium laneportoi’s similarity to S. pertinax was explained in the description.
Simulium spinibranchium is very similar and the adults can be differentiated only by the more
abundant pilosity on the scutum. The pupa with its shoe-shaped cocoon and thicker gill
branches allow it to be separated easily. Simulium serranus is also very similar and can be
separated only by its small size and lower number of head and thoracic tubercles. Comparative
cytological or molecular studies would be convenient to assure the identity of these species.

Simulium (Chirostilbia) serranus COSCARÓN (Figs. 57 H, I; 58 E, F; 59 D-F; 63 D-G.)


Simulium serranus COSCARÓN, 1981a: 158-160.
Simulium (Chirostilbia) serranus COSCARÓN, 1987: 26; 1991: 219-220.
Chirostilbia serranus PY-DANIEL & MOREIRA SAMPAIO, 1995: 118.
Female: Wing length 2.0-2.2 mm; scutum black with gray pollinosity, with ornamentation
as in Simulium laneportoi; legs darkened on tibial apex and tarsi of anterior leg, distally on tarsi,
medially and apically on tibia and terminal portion of tarsi of hind leg; scutellum light brown;
metanotum blackish; pleurae and abdomen brown grayish; frons with parallel sides (Fig. 57 H);
Sc without hair; eighth sternite with about 30 setae, gonapophysis and genital fork as in
Simulium pertinax; paraproct subtriangular, two times longer than wide at base (Fig. 57 I).
Male: Wing length 2.0-2.1 mm; scutum black bordered by grayish pollinose 1+1
anterior spots joining lateral stripe to posterior gray area (Fig. 58 E); changing light
position shows continuous grayish anterior band as in Simulium laneportoi; scutellum brown;
metanotum blackish; length/width ratio of hind basitarsus = 5.4; gonostylus subtrapezoi-
dal, distally truncate with acute apex (Fig. 58 F); ventral plate as in Simulium pertinax.
Pupa: Cocoon slipper shaped, weakly projected basally (Fig. 59 D); weave compact,
sometimes with threads surpassing anterior border and covering basal portion of gills;
length at base 2.9-3.2 mm, at dorsum 2.8-3.0 mm, gills 2.0-2.5 mm; frontoclypeus and
exposed portion of thorax with tubercles relatively abundant (Fig. 59 E) but more scarce
than in Simulium pertinax; cephalic trichomes with 1-3 branches and thoracic with 3-8
branches; gill with 8 branches, with primary ventral branch longer than primary dorsal;
secondary median branch two times longer than secondary dorsal branch (Fig. 59 F).
Larva: Maximum length 6.0 mm; cephalic apotome scarcely pigmented (Fig. 63 D);
antenna as long as cephalic fan stem; ratio of proximal, medial and distal antennal
segments = 1:1-1.3:1.6-2; cephalic fan with 45-50 rays. Mandible with second marginal
252 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

A D J

G O

P
B
E

C N
H

F
K

Q I R

Figure 59. Pupa. A-C: S. (Chirostilbia) pertinax; A: lateral view; B: frontoclypeus; C: portion of
thorax and gill. D-F: S. (C.) serranus; D: lateral view; E: frontoclypeus; F: portion of thorax and gill.
G-I: S. (C.) spinibranchium; G: lateral view; H: gill; I: thoracic trichome. J-K: S. (C.) empascae; J: lateral
view; K: gill. L-N: S. (C.) distinctum; L: lateral view; M: frontoclypeus; N: portion of thorax and gill.
O-P: S. (C.) riograndense; O: lateral view; P: frontoclypeus, Q: gill. R: S. (C.) friedlanderi, gill, lateral
view (extracted from original description). S-U: S. (C.) obesum; S: lateral view; T: frontoclypeus; U:
portion of thorax with trichomes and gill.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 253

tooth very short (Fig. 63 E); postgenal cleft subtriangular with acute apex; postgenal
bridge a little longer than half hypostomium length (Fig. 63 F); anal sclerite with single
hairs (Fig. 63 G); anal ring with 85-88 rows with 13-15 hooks each; lateral sclerite of
proleg with about 30 teeth; rectal papillae with about 15-18 diverticula on each lobe.
Distribution: Brazil: Mato Grosso, Minas Gerais.
Bionomics: Simulium serranus larvae live in small creeks with moderate current and
crystalline water, flowing over aquatic vegetation. The females are haematophagous and
very aggressive in Chapada dos Guimarães (MT), Brazil.
Discussion: Simulium serranus is very close to S. laneportoi. The differences were
explained in the description of that species. Simulium spinibranchium is similar but has more
abundant pilosity on the scutum and the pupal gill is thicker and has 8 or 9 branches. The
pupal cocoon is shoe shaped. Simulium pertinax can be distinguished by the absence of
submedian silvery vittae on the scutum.

Simulium (Chirostilbia) spinibranchium LUTZ (Figs. 57 J-L; 58 G, H; 59 G-I; 63 H.)


Simulium spinibranchium LUTZ, 1910: 248, 261, 263-264; PINTO, 1931: 712-713, 726-727, 756-762; PY-
DANIEL & SHELLEY, 1980: 213-223.
Simulium (Chirostilbia) spinibranchium COSCARÓN, 1987: 26; 1991: 220-223; SHELLEY et al., 2000: 175-176.
Chirostillbia spinibranchia PY-DANIEL & MOREIRA SAMPAIO, 1995: 118; STRIEDER, 2004: 128.
Female: Wing length 2.7-3.0 mm; scutum blackish gray pollinose with abundant gray
pile; with frontal light, 1+1 submedian gray pollinose stripes wider anteriorly and posteriorly,
last portion joining with grayish, prescutellar area (Fig. 57 J); with another light position this
submedian vittae changes to black with median and sublateral gray lead coloration;
scutellum brown grayish, metanotum velvety brown grayish; legs brown yellowish with
darker apical portion of segments blackish; abdomen brown blackish with tergite I and 1+1
lateral spots on tergite II light grayish brown; tergites V-VIII shiny; fronto-ocular triangle
absent or very reduced; eighth sternite well sclerotized with about 30 setae on each side,
gonapophysis with internal border well sclerotized (Fig. 57 K); paraproct about two times
longer than wide at base (Fig. 57 L); genital fork with anterior branch well sclerotized.
Male: Wing length 2.4-2.7 mm; scutum black with pilosity gray, and bordered
anteriorly, laterally and posteriorly by silvery pruinose stripe; with frontal light, grayish
anterior border very wide as in Simulium laneportoi; scutellum and metanotum as in female;
abdomen black with 1+1 grayish pollinose spots on tergites II, VI-VII, decreasing in size
from anterior to posterior segments; gonostylus apically acuminate (Fig. 58 G); ventral
plate body wider than high (Fig. 58 H).
Pupa: Cocoon shoe-shaped, with reinforced anterior border and protecting gill base
(Fig. 59 G); weave thick, with visible threads; frontoclypeus and exposed portion of
thorax with numerous tubercles, similar to Simulium pertinax; cephalic and thoracic
trichomes with 6-8 branches (Fig. 59 I); gills each with 8-9 branches emerging from 4
short primary branches from short basal trunk; branches thick at base and slightly
thinning to blunt apex (Fig. 59 H). Microstructure of gill branches with thin porosity.
Larva: Maximum length 6.0-6.7 mm; cephalic apotome darkened basally as in
Simulium pertinax, antennae a little shorter than cephalic fan stem; ratio of proximal,
254 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

medial and basal segments = 1:1-1.1:1.7-1.9. Mandible with latero-mandibular process;


postgenal cleft subtriangular with blunt apex; postgenal bridge about 2/3 of hypostomi-
um height; cephalic fan with 30-37 rays; lateral sclerite of proleg with 20 teeth arranged in
9-10 groups (Fig. 63 H); anal sclerite with hair and some scales like spines; anal ring with
76-86 rows with 12-14 hooks each; rectal papillae with 11-13 diverticula on each lobe;
ninth segment with 1+1 very small ventral tubercles; body cuticle without hair.
Distribution: Brazil: Bahia, Goiás, Mato Grosso, Minas Gerais, Rio de Janeiro, Rio
Grande do Sul, São Paulo; Venezuela.
Bionomics: Simulium spinibranchium larvae live in small creeks with slight discharge,
deep, clear water and regular current velocity, from 400 to 1200 m in altitude. The feeding
habits of the females are unknown.
Discussion: The closest species to S. spinibranchium is S. laneportoi, whose adults are
practically inseparable when the scutum pilosity is absent; the peculiar cocoon and gill
shape permit separation from similar species.

Simulium (Chirostilbia) empascae PY-DANIEL & MOREIRA (Figs. 58 I ; 59 J, K; 63 I-J.)


Simulium (Chirostilbia) empascae PY-DANIEL & MOREIRA, 1988: 77-86; COSCARÓN, 1991: 222-223.
Chrostillbia empascae PY-DANIEL & MOREIRA SAMPAIO, 1995: 118; STRIEDER, 2004: 126.
(Based on original description).
Female: (Pharate) General coloration black to dark brown; Sc and basal sector of R
with hair; genitalia similar to those of Simulium spinibranchium and S. obesum.
Male: (Pharate) Gonostylus wide basally, blunt apically, with very small subapical
spine (Fig. 58 I); ventral plate as in Simulium distinctum.
Pupa: Cocoon shoe shaped, protecting gills (Fig. 59 J); frontoclypeus without
tubercles; exposed portion of thorax with short and acute tubercles on posterior area;
cephalic trichomes single and short, thoracic single but more enlarged; gills with 8
branches emerging from short basal trunk and with primary and secondary branches also
very short (Fig. 59 K).
Larva: Maximum length 8.1-11.8 mm; cephalic apotome similar to that of Simulium
pertinax; antenna relatively thick (Fig. 63 I); ratio of proximal, medial and distal segments
= 1:1.3-1.6:1-1.3; cephalic fan with 53-54 rays. Mandible with latero-mandibular process
(Fig. 63 J); postgenal cleft subtriangular with acute apex; postgenal bridge about half of
hypostomium length; lateral sclerite of proleg with about 28 teeth; anal ring with 201-204
rows with 19-26 hooks each; rectal papillae with 4-6 diverticula on each lobe.
Distribution: Brazil: Santa Catarina, Rio Grande do Sul.
Bionomics: Simulium empascae breeds in fast-flowing creeks in areas with direct
sunlight. Feeding habits of the females are unknown.
Discussion: The peculiar cocoon shape covering all the gills and the gill branching
allow the differentiation of this species from similar species in the subgenus. The
gonostylus with a blunt apex and with a spine and the abundant rows of hooks on the anal
ring also help to separate this species from others.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 255

Simulium (Chirostilbia) distinctum LUTZ (Figs. 57 M, N; 58 J, K; 59 L-N; 63 K, L.)


Simulium distinctum LUTZ, 1910: 241-243; PINTO, 1931: 699-701; D’ANDRETTA & D’ANDRETTA, 1950:
197-198; EDWARDS, 1933: 210; D’ANDRETTA, 1954: 63-66; SHELLEY et al., 1984: 145.
Simulium (Simulium) distinctum ORFILA, 1939: 1533.
Simulium (Chirostilbia) distinctum COSCARÓN, 1987: 26; 1991: 223-225; PY-DANIEL et al., 1988: 43-48.
?Trichodagmia distincta ENDERLEIN, 1934a: 194; 1936: 128.
Trichodagmia spadicidorsum ENDERLEIN, 1934a: 194.
Simulium pertinax LANE & PORTO, 1929: 170; VARGAS, 1945a: 182.
Simulium prumirimense COSCARÓN, 1981a: 160-164.
Chirostilbia distincta PY-DANIEL & MOREIRA SAMPAIO, 1995: 118; STRIEDER, 2004: 133-135; 2004: 125-126.
Female: Wing length 2.5-3.4 mm; general coloration dark grayish brown; scutum
brown grayish with decumbent golden to silvery pilosity, with posterior border darker
pollinose, giving dark tones, one median and 1+1 sublateral stripes gray pollinose not
reaching anterior and posterior borders; scutellum brown; metanotum reddish brown with
grayish pollinosity; pleura dark grayish brown with gray pollinosity; legs grayish brown
with tarsi, distal portion of femur, tibia and basitarsus of hind leg blackish; abdomen dark
brown grayish with tergite I, center of II-IV brown with dark pile; VI-IX tergites shiny;
tergite II with 1+1 silvery pruinose lateral spots; ventrally light brown grayish; fronto-
ocular triangle higher than wide; basal sector of R hairy and Sc with 10 setae; claw with
subbasal tooth (Fig. 57 M); eighth sternite with about 60 setae; gonapophysis with internal
margin more sclerotized and without microtrichia; paraprocts acuminate distally (Fig. 57
N) and genital fork with anterior branch strongly sclerotized.
Male: Wing length 2.4-3.3 mm; scutum blackish with yellow greenish pile; abdomen
blackish with 1+1 silvery spots on tergites II, V-VI and smaller spots on VII-VIII and IX;
length/width ratio of hind basitarsus = 6; gonostylus subquadrate, with smooth external
distal process (Fig. 58 J); ventral plate elongated with blunt apex (Fig. 58 K).
Pupa: Cocoon shoe shaped with thick weave dark brown leaving holes anteriorly and
protecting gill branches laterally (Fig. 59 L); length at base 3.0 mm; at dorsum 2.8, gills 1.0
mm; frontoclypeus and thorax with moderate quantity of tubercles (Fig. 59 M); cephalic
trichomes with 2-3 branches, and thoracic with 3-6 branches; gills each with 10 short
branches, with 3 short primary branches emerging from short basal trunk; gill filaments
relatively thick, curved and sclerotized distally, terminal portion frequently lost (Fig. 59 N).
Larva: Maximum length 8.5-9.0 mm; general aspect as in figure 63 K; cephalic
apotome brown with median basal area darkened but not like triangle as in Simulium
pertinax; slightly darkened antero-lateral and posterior spots; antennae as long as or
surpassing cephalic fan stem; ratio of proximal, medial and distal segments = 1:1.5-
1.6:1.2-1.3; cephalic fan with 45-50 rays. Mandibles with second marginal tooth nearly as
long as first (Fig. 63 L); postgenal bridge longer than half of hypostomial length; lateral
sclerite of proleg with 32-38 teeth arranged in 10-13 groups; anal ring with 170-190 rows
of 18-20 hooks each; body integument with scarce, short hair; rectal papillae with 7-9
diverticula on each lobe.
Distribution: Brazil: Minas Gerais, Paraná, Rio de Janeiro, Rio Grande do Sul, São
Paulo.
256 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Bionomics: Simulium distinctum breeds in creeks of fast-flowing clear water, attached


to stones. Feeding behavior of the females is unknown.
Discussion: The closest species to S. distinctum is S. riograndense, but it can be
separated by the light brown yellowish scutum, and very sparse pupal tubercles reduced
on the frontoclypeus to 1+1 groups on the base at the sides of the facial trichomes and on
the posterior area of the thorax.

Simulium (Chirostilbia) riograndense PY-DANIEL, SOUZA & CALLAS (Figs. 58 L, M; 59 O-Q.)


Simulium (Chirostilbia) riograndense PY-DANIEL et al., 1998: 38-43; CROSSKEY & HOWARD, 2004: 20.
Chirostilbia riograndense PY-DANIEL & MOREIRA SAMPAIO, 1995: 118; STRIEDER, 2002: 133-135; 2004: 127.
Female: Wing length 3.2-3.6 mm; scutum light brown-yellowish with light grayish to
yellowish pile; with frontal light, 1+1 sublateral grayish pruinose stripes, becoming a little
darker with posterior light and darker than prescutellar area; scutellum brown and metanotum
and pleurae dark brown with gray pruinosity; legs yellowish with tibiae and tarsi of anterior leg,
apices of basitarsi and tarsi of middle and posterior legs and apex of hind tibiae blackish;
abdomen brown with 1+1 grayish spots on tergite II; eighth sternite with about 26-38 setae on
each side, gonapophysis, paraproct and genital fork as in Simulium distinctum.
Male: Wing length 3.3-3.5 mm; scutum brown-reddish to yellowish, more darkened
than in female, with silvery to yellowish pile; with postero-lateral light, silvery pollinose
band bordering posterior and sides; scutellum and metanotum as in female; abdomen
blackish with 1+1 silvery pollinose spots on tergites II-VIII, more developed on tergite II;
genitalia very similar to those of Simulium distinctum but gonostylus more sclerotized with
well-developed process on external distal side, and microscopic subapical spur (Fig. 58 L);
ventral plate not elongated and more rounded apically (Fig. 58 M).
Pupa: Cocoon shoe shaped, weave resistant with abundant holes and protecting
branches laterally (Fig. 59 O); length at base 4.2-4.7 mm, at dorsum 4.6-4.7 mm; gills 1.7-
2.0 mm; frontoclypeus and exposed portion of thorax with scarce tubercles reduced to
1+1 small areas at frontoclypeal base (Fig. 59 P) and to posterior dorsal area of thorax,
those mostly acuminate; cephalic trichomes short, single or bifid; thoracic trichomes with
1-4 branches; gills each with 10 branches emerging from short basal trunk and very short
primary branches (Fig. 59 Q).
Larva: Maximum length 9.5-10.5 mm; cephalic apotome with median basal area
brownish; spots positive, median longitudinal and 1+1 submedian subbasally positioned;
antennae scarcely surpassing cephalic fan stem; ratio of proximal, medial and distal
segments = 1:1.8-2.1:0.9-1.3; cephalic fan with 43-50 rays; mandible very similar to that of
Simulium distinctum; postgenal cleft subtriangular, blunt apically; postgenal bridge more
than half hypostomium length; anal ring with 145-152 rows with 19-25 hooks each; rectal
papillae with 9-11 diverticula on each lobe.
Distribution: Brazil: Paraná, Rio Grande do Sul.
Bionomics: Simulium riograndense breeds in slow, acidic water in temperatures of 19-
20 °C, attached to stones. Feeding behavior of females is unknown.
Discussion: The closest species to S. riograndense is S. distinctum in whose species
description the differences are mentioned.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 257

Simulium (Chirostilbia) bifenestratum HAMADA & PEPINELLI


Simulium (Chirostilbia) bifenestratum HAMADA & PEPINELLI, 2004: 45-52.
(Based on original description).
Female: Wing length 3.3 mm; scutum dark gray with 1+1 submedian small subrectan-
gular silver spots, pilosity pale golden; scutellum light brown, metanotum brown; pleurae
dark brown; legs light brown with darkened black portion; abdomen dark brown with
membranous areas light brown; tergite II with 1+1 silver pruinose spots, VI-IX shiny;
cibarium smooth with well sclerotized external cornuae branches; basal sector of R and Sc
with hair; claw with subbasal tooth; genitalia very similar to those of Simulium distinctum.
Male: Wing length 3.1 mm; scutum black with dense golden pile; scutellum dark
brown, metanotum black; legs coloration as in female; genitalia as in Simulium distinctum.
Pupa: Cocoon slipper shaped, with strong and thick weave, showing 1+1 anterior
sublateral subovoidal holes, and protecting most gill branches; length at base 1.8-2.2 mm; at
dorsum 3.9-4.5 mm; frontoclypeus with rounded tubercles on thoracic middle area larger,
with protuberances and sometimes acuminate, frontoclypeus with marginal area elevated, in
special cases lateral border with a crown; cephalic trichomes with 1-3 branches, thoracic with
3-9 branches; gill with 10 branches, thicker at base and surrounding head.
Larva: Maximum length 8.4-9.0 mm; cephalic apotome darkened on mid to
posterior area, and postgenal cleft shaped as in Simulium pertinax; antenna as long as
cephalic fan stem; ratio of proximal, medial and distal segments = 1:1.8-1.9:0.8-.9;
cephalic fan with 45-48 rays; anal ring with 152-158 rows with 15-19 hooks each; rectal
papillae with 20-26 elongated diverticula in total; body cuticle with single-branched hairs.
Distribution: Brazil: São Paulo.
Bionomics: Simulium bifenestratum breeds in creeks with rocky bottoms, about 10 m
wide, at an altitude of 1500 m. Feeding habits of the females are unknown.
Discussion: Similar to species of the PERITNAX group, in particular to S. distinctum,
S. bifenestratum can be separated by the presence on the female scutum of small anterior
submedian silvery spots and 1+1 anterior sublateral apertures in the thick cocoon.

Simulium (Chirostilbia) friedlanderi PY-DANIEL (Fig. 59 R.)


Simulium (Chirostilbia) friedlanderi PY-DANIEL, 1987: 331-336; COSCARÓN, 1991: 227-228.
(Based on original description).
Female: Unknown.
Male: Scutum dark brown with yellowish pile; lateral illumination shows whitish
pollinosity bordering scutum, frontal light shows 1+1 lateral and antero-dorsal pruinose
spots; genitalia similar to those of Simulium obesum.
Pupa: Cocoon shoe shaped very similar to Simulium riograndense and S. obesum, and
very elevated anteriorly; weave with abundant holes and protecting all gill filaments
laterally; length at base 3.1-3.9 mm; at dorsum 3.4-5.0 mm, gills 1.2 mm; frontoclypeus
and exposed portion of thorax with numerous small tubercles; cephalic and thoracic
trichomes with 3-7 branches; gill with 11 very thick filaments, thinning to acute apices
with strong curvature (Fig. 59 R).
258 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Larva: Maximum length 6.2-6.4 mm; cephalic apotome darkened on mid basal area
and with median longitudinal and median sublateral positive spots; antennae surpassing
cephalic fan stem; ratio of proximal, medial and distal segments length = 1:1-1.2:1.3-1.5;
cephalic fan with 43-47 rays; postgenal cleft subtriangular; with blunt apex; postgenal
bridge a little longer than half hypostomium length. Mandible with second marginal tooth
surpassing half of anterior length; latero-mandibular process present; lateral sclerite of
proleg with about 23 teeth in 7-8 groups; anal ring with 105-122 rows of 16-20 hooks
each; rectal papillae with 20-24 diverticula on each lobe.
Distribution: Brazil: Mato Grosso.
Bionomics: Larval stages of S. friedlanderi live in flows with rocky bottoms. Feeding
behavior of the females is unknown.
Discussion: The known stages are very close to those of S. distinctum, S. riograndense
and S. obesum, but the peculiar thick gill of S. friedlanderi permits separation.

Simulium (Chirostilbia) obesum VULCANO (Figs. 57 O-V; 58 O, P; 59 S-U.)


Simulium obesum VULCANO, 1959: 351-360; LUTZ, 1922: 89-92 (nomen nudum); PINTO, 1931: 753, 762
(nomen nudum).
Simulium (Chirostilbia) obesum COSCARÓN, 1987: 26; 1991: 228-229.
Chirostilbia obesa PY-DANIEL & MOREIRA SAMPAIO, 1995: 118.
Female: Wing length 3.0-3.6 mm; scutum dark brown reddish to blackish with soft
gray pollinosity and golden decumbent pilosity, lateral and posterior edges with silvery
pollinose band; frontal illumination shows 1+1 silvery submedian and sublateral vittae
separated by one median and 1+1 sublateral black stripes that becomes wider with lateral
light (Fig. 57 O); scutellum dark brown reddish; metanotum velvety black and pleural
areas blackish with gray pollinosity; legs light brown with darkened areas dark brown to
black, as in figures 57 Q-S; abdomen blackish with tergal posterior border dark gray;
tergite II with 1+1 lateral silvery spots; tergites I-V velvety; VI-IX shiny. Maxillary palpal
sensory organ small, shorter than half of palpomere length (Fig. 57 P); claw with subbasal
tooth well developed; Sc with about 30 setae; eighth sternite with about 40 setae on each
side; gonapophysis with rounded apex and without microtrichia (Fig. 57 T); paraprocts
two times longer than width at base (Fig. 57 U); genital fork with strong anterior branch
(Fig. 57 V); spermatheca subovoidal without sclerotized neck and light circular areas
between spiculae.
Male: Wing length 3.2-3.5 mm; general coloration as in female; scutum velvety black,
bordered laterally and posteriorly by silvery band; frontal light shows 1+1 anterior
sublateral silver spots; legs as in female but more darkened especially on anterior legs;
abdomen velvety black, tergites II and VI with 1+1 silvery spots, bigger on II; gonostylus
about half gonocoxite length (Fig. 58 O); ventral plate subtriangular, a little wider than
high (Fig. 58 P).
Pupa: Cocoon with threads evident, leaving large apertures, anteriorly elevated,
projected anteriorly and protecting all gill filaments (Fig. 59 S); length at base 3.1-4.0 mm;
at dorsum 4.0 mm; gills 1.3 mm; frontoclypeus and exposed portion of thorax with
abundant tubercles (Fig. 59 T); cephalic trichomes with 2-5 branches; thorax dorsally
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 259

covered with abundant trichomes with 1-3 branches; gills filiform, arborescent, each with
about 120 terminal branches; primary branches about 7 in number, relatively short,
emerging form base without evident basal trunk (Fig. 59 U); abdomen with two strong
terminal spines.
Larva: Undescribed.
Distribution: Brazil: Rio de Janeiro, Santa Catarina, São Paulo.
Bionomics: Simulium obesum breeds in torrential creeks with crystalline, relatively
cold water. The female feeding habits are unknown.
Discussion: Coloration of the adults of S. obesum is similar to that of S. distinctum,
but is lighter. The peculiar pupa with its many gill branches and abundant trichomes on
the thorax allows separation of S. obesum from other species.

SUBPALLIDUM species group

Simulium (Chirostilbia) subpallidum LUTZ (Figs. 60 A-C; 61 H-K; 62 A, B; 63 M-P.)


Simulium subpallidum LUTZ, 1910: 247-248; PINTO, 1931: 714, 742, 755; COSCARÓN, 1982: 66, 67, 73-76.
Simulium (Chirostilbia) subpallidum COSCARÓN, 1987: 26; 1991: 230-232; SHELLEY et al., 2000: 176-177.
Simulium subviride LUTZ & MACHADO, 1915: 46 (nomem nudum).
Simulium guarani COSCARÓN & WYGODZINSKY, 1972a: 210-216.
Chirostilbia subpallida PY-DANIEL & MOREIRA SAMPAIO, 1995: 118; STRIEDER, 2004: 128.
Female: Wing length 2.0-2.8 mm; scutum and scutellum light brown yellowish with
gray yellowish pile, some specimens with 1+1 small dark gray postero-lateral spots;
posterior illumination shows submedian light gray stripes joined posteriorly with lyre
pattern; pleurae gray yellowish with basal area silvery gray. Metanotum velvety brown
blackish; legs yellowish with dark portion brown, nearly black on anterior tarsi; abdomen
with tergites I-II yellowish, III-V yellow grayish with tergal plate brown dark gray not
reaching posterior border and in III-IV another 1+1 lateral dark spot; VI-IX with cereous
sheen on tergal plates and yellowish laterally; frons slightly narrowed (Fig. 60 A); fronto-
ocular triangle as high as wide (Fig. 60 B); basal portion of cibarium smooth with cornuae
well sclerotized; basal portion of R with hair in 1-2 rows and Sc generally without hair
(Fig. 60 C); claws with or without small tooth; eighth sternite, gonapophyses and genital
fork as in Simulium acarayense; spermatheca with abundant microtrichia arranged in rows;
paraproct about two times longer than wide at base and with small basal lobe.
Male: Wing length 1.8-2.5 mm; scutum and scutellum brown yellowish to reddish,
with 1+1 blackish spots postero-laterally; laterally and posteriorly bordered by silvery gray
band; with changing light position, lateral band can increase in size, covering one quarter
of antero-lateral area; scutum pilosity golden; pleurae gray, silvery and shiny basally; legs
brown yellowish with dark areas larger than in female; abdomen black with 1+1 silvery
spots on tergites II, VI-VIII; gonostylus as long as gonocoxite, longer than wide, slightly
curved and with lateral crest (Figs. 61 H, I); ventral plate wider than high, with thin hair
(Figs. 61 J, K). Median sclerite and endoparameres as in Simulium papaveroi.
Pupa: Cocoon slipper shaped, with straight anterior border and slightly projected
anteriorly (Fig. 62 A); compact weave with evident threads; length at base 3.6-4.0 mm, at
260 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

A B

N
I
J
D

M
E
P

G F

Figure 60. Female. A-C: S. (Chirostilbia) subpallidum; A: frons; B: fronto-ocular triangle; C: veins of
anterior portion of wing. D-K: S. (C.) acarayense; D: basal portion of cibarium; E: abdomen; F:
eighth sternite and gonapophysis; G: genital fork; H: cercus and paraproct; I-K: fore, mid and hind
leg of male, respectively. L-P: S. (C.) papaveroi; L-M: scutum with lateral and frontal illumination; N:
sensory vesicle of maxillary palp and palpomere; O: claw; P: cercus and paraproct.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 261

dorsum 2.6-3.5 mm, gills 1.5-2.3 mm; frontoclypeus and thorax without tubercles or these
restricted to basal area on frontoclypeus, area of gill and postero-dorsally on thorax with
conical shape; cephalic trichomes large with 3-5 branches and thoracic trichomes with 4-
6 branches; gills with 8 branches emerging from short basal trunk and with short primary
branches (Fig. 62 B).
Larva: Maximum length 5.0 mm; color light green grayish; cephalic apotome light
brown with positive spots (Fig. 63 M); antennae as long as or shorter than stem of
cephalic fan; ratio of proximal, medial and distal segments = 1:1.2-1.5 (Fig. 63 N).
Mandible with second marginal tooth half length of first tooth (Fig. 63 O); cephalic fan
with 32-39 rays; postgenal cleft subovoidal; postgenal bridge half of hypostomium length;
anal sclerite with single hairs, and scales with 3-5 branches (Fig. 63 P); anal ring with 70-80
rows with 12-14 hooks each; body cuticle without hairs; segment nine with 2 small ventral
tubercles; rectal papillae with 4-7 diverticula on each lobe.
Distribution: Argentina: Misiones; Brazil: Bahia, Espírito Santo, Goiás, Minas
Gerais, Pará, Paraná, Rio Grande do Sul, Santa Catarina, São Paulo.
Bionomics: Simulium subpallidum breeds in small creeks, with moderate current,
attached to aquatic vegetation. The feeding habits of the females are unknown.
Discussion: The closest species to S. subpallidum is S. acarayense. Females can be
separated by the gray-lead scutum, and males by the blackish scutum and shorter
gonostylus. Simulium papaveroi also is similar but can be separated in the female by the dark
grayish brown scutum, and more elongated paraproct; in the male by the blackish scutum;
in the pupa by the more grouped branches and cephalic trichomes with 1-2 branches.
Karyological or molecular studies are needed to assure the species identity, particularly of
S. subpallidum and S. acarayense, which are very close.

Simulium (Chirostilbia) acarayense COSCARÓN & WYGODZINSKY (Figs. 60 D-K; 61 A-E; 62 C-F.)
Simulium acarayense COSCARÓN & WYGODZINSKY, 1972a: 216-223; COSCARÓN, 1982: 67-68.
Simulium (Chirostilbia) acarayense COSCARÓN, 1987: 26; 1991: 232-233.
Chirostilbia acarayense PY-DANIEL & MOREIRA SAMPAIO, 1995: 118; STRIEDER, 2004: 125.
Female: Wing length 2.0-2.2 mm; scutum gray-lead with dark background and gray
pollinosity, pile yellowish; scutellum light brown with gray pollinosity; metanotum velvety
dark brown; pleurae brown grayish; legs light yellow with dark areas blackish (Figs. 60 I-K);
abdomen gray greenish with tergal plates of segments III-VI dark brown and more dilute
laterally on I-VI (Fig. 60 E); fronto-ocular triangle higher than wide; cibarium with median
portion light (Fig. 60 D); basal sector of R with hair; eighth sternite scarcely sclerotized with
about 20 setae; gonapophysis longer than wide (Fig. 60 F); genital fork with anterior branch
well sclerotized (Fig. 60 G); paraproct about 3 times longer than wide at base (Fig. 60 H).
Male: Wing length 2.2 mm; scutum blackish with silvery band laterally and
posteriorly and enlarged antero-medially (Figs. 61 A, B); scutellum, metanotum and
pleurae as in female; legs yellowish with distribution of pigmentation and relative
proportion of size similar to female; abdomen gray greenish, with large light areas on
tergites II, and smaller ones on III-VI (Fig. 61 C); gonocoxite, gonostylus and ventral plate
as shown in figures 61 D, E.
262 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

B
C

I
H

F G

E K

Figure 61. Male. A-E: S. (Chirostilbia) acarayense; A-B: scutum in dorsal and lateral views; C:
abdomen; D: gonocoxite and gonostylus; E: ventral plate. F-G: S. (C.) papaveroi; F: gonocoxite and
gonostylus; G: median sclerite and endoparameres. H-K: S. (C.) subpallidum; H-I: gonocoxite and
gonostylus in dorsal and ventral view; J-K: ventral plate in different position.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 263

Pupa: Cocoon slipper shaped with straight edge and slightly projected anteriorly
(Figs. 62 C, D); shorter than in Simulium subpallidum; weave compact with threads obvious;
length at base 2.6-2.7 mm; at dorsum 2.2-2.3 mm; gills 1.1-1.2 mm; frontoclypeus and
thorax with scarce tubercles (Fig. 62 E); cephalic and thoracic trichomes with 3-5
branches; gills with 8 branches, open from base (Fig. 62 F).
Larva: Maximum length 5.3-5.8 mm; cephalic apotome with more pigmentation at
center and base; ratio of proximal, medial and distal antennal segments = 1:1.1-1.2:1.1-
1.6; cephalic fan with 44-50 rays; postgenal cleft subovoidal, postgenal bridge a little
shorter than half of hypostomium length; anal ring with 50-70 rows with 14-15 hooks
each; rectal papillae with 8-10 diverticula on each lobe.
Distribution: Argentina: Entre Ríos, Misiones; Brazil: Ceará, Mato Grosso, Minas
Gerais, Rio Grande do Sul, Santa Catarina, São Paulo; Paraguay; Uruguay.
Bionomics: Simulium acarayense breeds in creeks with much discharge and fast-
flowing, crystalline water, attached to leaves and branches. The female feeding habits are
unknown.
Discussion: Similarities of S. acarayense with S. subpallidum were treated in the species
description.

Simulium (Chirostilbia) papaveroi COSCARÓN (Figs. 60 L-P; 61 F, G; 62 G-I; 63 Q.)


Simulium papaveroi COSCARÓN, 1982: 67-73.
Simulium (Chirostilbia) papaveroi COSCARÓN, 1987: 26; 1991: 233-235.
Chirostilbia papaveroi PY-DANIEL & MOREIRA SAMPAIO, 1995: 118.
Female: Wing length 1.8-2.0 mm; general coloration dark brown grayish; scutum
brown with gray pollinosity and pilosity silver to golden; laterally and posteriorly bordered
by silvery band; with lateral light, scutum shows 1+1 submedian silvery vittae, joined
posteriorly to grayish posterior area (Fig. 60 L); with frontal illumination, vittae and border
change to black (Fig. 60 M); scutellum light brown; metanotum velvety blackish with scarce
darker portion; abdomen brown grayish with tergite I light brown grayish; tergite II with
1+1 silvery lateral spots; tergites III-VI black with silvery narrow posterior margin; tergites
VII-IX green grayish; fronto-ocular triangle short; sensory vesicle of maxillary palp about
half of palpomere length and with abundant tuberosity (Fig. 60 N); basal sector of R and Sc
without hair; claw without basal tooth (Fig. 60 O); eighth sternite with about 28 setae;
gonapophysis subtriangular, without microtrichia; paraproct nearly two times longer than
wide at base (Fig. 60 P); genital fork with anterior branch well sclerotized.
Male: Wing length 1.7-1.8 mm; general coloration blackish; scutum black, bordered
laterally and posteriorly by silvery pruinose band; with lateral illumination, silvery area
enlarged antero-laterally as in Simulium acarayense; scutellum, metanotum and pleurae as in
female; abdomen blackish with 1+1 silvery spots on tergites II, V-IX; length/wide ratio of
hind basitarsus = 6; gonostylus relatively thin, about two times longer than wide at base,
with median diagonal crest (Fig. 61 F); ventral plate very similar to that of Simulium
subpallidum; median sclerite with deep incision; endoparameres with big teeth (Fig. 61 G).
Pupa: Cocoon slipper shaped, weave thick, with evident threads, frequently with thin
weave covering thorax dorsally and gills (Fig. 62 G); length at base 2.5-2.8 mm, at dorsum
264 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

H
D

Figure 62. Pupa. A-B: S. (Chirostilbia) subpallidum; A: lateral view; B: anterior portion of thorax
and gill. C-F: S. (C.) acarayense; C-D: lateral and dorsal views; E: frontoclypeus; F: anterior portion
of thorax and gill. G-I: S. (C.) papaveroi; G: lateral view; H: frontoclypeus; I: anterior portion of
thorax and gill.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 265

K M

A
N H
B I

E L

J O

Q
F

G P

Figure 63. Larva. A-C: S. (Chirostilbia) pertinax; A: cephalic apotome; B: antenna, C: postgenal cleft
and hypostomium. D-G: S. (C.) serranus; D: head, dorsal view; E: teeth of mandible; F: postgenal
cleft and hypostomium; G: anal sclerite. H: S. (C.) spinibranchium; lateral sclerite of proleg. I-J: S. (C.)
empascae; I: antenna; J: teeth of mandible. K-L: S. (C.) distinctum; K: lateral view; L: teeth of
mandible. M-P: S. (C.) subpallidum; M: head, dorsal view; N: antenna; O: teeth of mandible; P: anal
sclerite. Q: S. (C.) papaveroi, postgenal cleft and hypostomium.
266 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

2.0-2.3 mm, gills 1.5-1.7 mm; frontoclypeus (Fig. 62 H) and exposed portion of thorax
without tubercles, or only very scarce on thorax; cephalic and thoracic trichomes single or
bifid and very short; gills each with 8 branches with short basal trunk, primary and
secondary branches (Fig. 62 I); abdomen with short terminal spines.
Larva: Maximum length 5.5-6.0 mm; head brownish, body gray green-bluish;
cephalic apotome scarcely pigmented and light on anterior area and lateral borders; ratio
of proximal, medial and distal antennal segments = 1:1.2:1.5-2; postgenal cleft rounded
anteriorly with postgenal bridge about half of hypostomium length (Fig. 63 Q); second
marginal tooth about 1/4 of first; latero-mandibular process present; cephalic fan with
about 40-46 rays; anal sclerite with single hairs; anal ring with 70-81 rows with 12-14
hooks each; lateral sclerite of thoracic proleg with about 25 teeth arranged in 8 groups;
body cuticle without hair.
Distribution: Brazil: Bahia, Minas Gerais.
Bionomics: Simulium papaveroi breeds in torrential creeks with large discharge and
crystalline water. The female feeding habits are unknown.
Discussion: Imagoes of the PERTINAX species group have coloration similar to
that of S. papaveroi, in particular S. serranus. Differences in gonostylus morphology,
presence of frontoclypeal and thoracic tubercles and disposition of the gill branches allow
separation of S. papaveroi from other species. Simulium acarayense is also very similar but has
a relatively shorter gonostylus and very open pupal gill branches.

Species inquirenda

Simulium striginotum (ENDERLEIN)


Thyrsopelma striginotum ENDERLEIN, 1934b: 284.
Female type material from the Dresden Museum is very similar to S. laneportoi. Lack
of knowledge of the genitalia morphology and other stages, do not permit its comparison
with other species; we, therefore, maintain S. striginotum as a species inquirenda.

Simulium (Inaequalium) COSCARÓN & WYGODZINSKY (Figs. 64-70; 141.)


Type-species: Eusimulium inaequale PATERSON & SHANNON, 1927.
Female: Species medium sized and blackish; frons and clypeus gray pollinose;
without bright opalescence; scutum black with grayish pollinosity, generally with gray
band around lateral and posterior edges and in some species with 1+1 submedian anterior
light gray stripes; scutum with decumbent homogeneous silvery to yellowish pilosity;
scutellum, metanotum and pleurae black with gray pilosity; abdomen black with 1+1
silvery spots on tergite II; tergites I-IV velvety and V-VIII with cereous sheen; frons
slightly convergent; fronto-ocular triangle deep; cibarium with basal portion projected
medially and covered with robust and well-sclerotized spines; legs with petaloid hair;
calcipala well developed, longer than wider; basal sector of R with hair; Sc with about 20
hairs; claws generally with well-developed subbasal tooth; gonapophysis divergent,
subtriangular, with sclerotized internal edge; eighth sternite with about 30-40 hairs on
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 267

each side; cerci rounded apically; paraprocts generally shorter than wide, apically rounded
or truncated; genital fork elongated, about 2 times length of each anterior branch.
Male: Scutum velvety black, grayish silvery on lateral and posterior borders; with
some illumination, 1+1 submedian gray pollinose spots evident on anterior third; scutum
pilosity yellowish to golden; abdomen blackish with tergites II, VI, VII and 1+1 on VIII
with small anterior portion whitish-silvery; basal sector of R with hair arranged in one
row; Sc with 3-4 hairs; calcipala as in female; gonocoxite subtrapezoidal, about as long as
wide; gonostylus slightly shorter than gonocoxite, subtriangular to subtrapezoidal, with
large apical or subapical spur; ventral plate subovoidal to subtriangular, wider than long;
endoparameres with robust spines.
Pupa: Cocoon slipper shaped with anterior edge generally straight and reinforced;
weave strong with threads obvious; gill generally with 6 filiform branches or with robust
2-3 irregular shaped branches; cephalothorax with obvious tubercles; frontoclypeal
trichomes with 3-5 branches, and thoracic trichomes with 3-8 branches; abdominal spines,
hooks and terminal spines well developed.
Larva: Body without hair or scales; cephalic apotome generally with darkened median
posterior and basal area, margin with light spots, or with more diffuse ornamentation;
postgenal cleft generally deep with postgenal bridge smaller or about as long as hypostomi-
um. Median hypostomium tooth lower than lateral teeth; hypostomial setae about 6 on each
side; antennae surpassing stem of cephalic fan; terminal segment longer than each preceding
segment. Mandible with 2 marginal teeth, anterior tooth over two times length of second;
with robust latero-mandibular tooth; cephalic fan with 35-40 rays; anal sclerite with single
hairs; anal ring with 65-90 rows with 12-15 hooks in each; ninth segment with two small
ventral tubercles, sometimes not evident; rectal papillae with 2-13 diverticula on each lobe.
Distribution: Simulium (Inaequalium) is a peculiarly South American taxon (although
reaching Panama) from Venezuela to northern Argentina, present in all the countries,
except Chile; the greatest diversity is in the southeastern Brazil mountain areas (Fig. 141).
Discussion: Simulium (Inaequalium) has 14 known species; adults are very similar, but
pupae have good characters to differentiate the species, especially by gill morphology;.
Two species groups are evident, as shown through cladistic analysis (COSCARÓN &
COSCARÓN ARIAS (1997) and STRIEDER (1997):
· The INAEQUALE species group is characterized as follows: basal portion of
cibarium pyramidal, acuminate apically; paraprocts longer than high, not acumi-
nate distally; gonostylus shorter than gonocoxite; ventral plate wider than long;
pupal gill composed of 6 filamentous branches and postgenal bridge shorter than
hypostomium length. Simulium inaequale, S. clarki, S. subnigrum, S. nogueirai, S.
leopoldense, S. beaupertuyi, S. hauseri, S. clavibranchium, S. subclavibranchium, S. nahimi, S.
mariavulcanoae, S. diversibranchium, S. rappae, S. travassosi..
· The BOTULIBRACHIUM species group is characterized as follows: basal
portion of cibarium with very low pyramidal median process to relatively
depressed medially; paraprocts higher than long and acuminate distally; gonosty-
lus generally as long as gonocoxite; ventral plate as long as wide; pupa gill
composed of 2-6 thick irregular filaments; larval postgenal bridge as long as
hypostomium length. Simulium botulibranchium, S. souzalopesi, S. petropoliense.
268 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

KEYS TO SIMULIUM (INAEQUALIUM) SPECIES

Females
Simulium hauseri and S. leopoldense females are unknown.

1 Paraprocts longer than high (Figs. 64 G, L, P) ............................................................ 2


– Paraprocts higher than long, acuminate distally (Figs. 69 D, Q) ............................. 6
2(1) Scutum without 1+1 submedian anterior band in certain lighting ...........................
............................................................ Simulium mariavulcanoae, Simulium diversibranchium
– Scutum with 1+1 submedian anterior band in certain light (Fig. 64 A) ................ 3
3(2) Coxae I blackish; femora and tibiae with strong blackish coloration; paraprocts
proportionally low (Fig. 64 L) ................................................... Simulium clavibranchium
– Coxae I light grayish-brown, femora and tibiae with slight blackish coloration (Fig.
64 C); paraprocts proportionally higher (Figs. 64 G, P) ............................................ 4
4(3) Paraproct relatively high with apical ovoidal edge (Fig. 64 P); genital fork with
stem relatively longer and posterior branches proportionately thicker (Fig. 64 O)
...................................................................................................................Simulium travassosi
– Paraproct relatively low with apical rounded edge (Fig. 64 G); genital fork with
stem relatively shorter and posterior branches proportionately thinner (Fig. 64 N)
................................................................................................................................................ 5
5(4) Wing length 1.8-2.0 mm ................ Simulium rappae, Simulium clarki, Simulium nahimi
– Wing length 2.3-2.8 mm .....................................................................................................
....... Simulium inaequale, Simulium nogueirai, Simulium subclavibranchium, Simulium subnigrum
6(1) Gonapophyses with parallel inner margins (Fig. 69 T) ............ Simulium petropoliense
– Gonapophyses with divergent inner margins (Fig. 69 P) .......................................... 7
7(6) Basal portion of cibarium with subconical medial elevation (Fig. 69 C) .................
........................................................................................................ Simulium botulibranchium
– Basal portion of cibarium slightly elevated, with small median depression (Fig. 69 K)
................................................................................................................ Simulium souzalopesi

Males
Simulium leopoldense, S. hauseri and S. petropoliense males are unknown.

1 Gonostylus short, about 1/2 as long as gonocoxite (Figs. 65 C, G, M), and not
more than 2 times longer than wide at base (Figs. 65 C, D, F, G, H, J, K, L, N, P);
ventral plate wider than long (Figs. 65 C, E, I, O) ..................................................... 2
– Gonostylus elongated, about same length as gonocoxite (Fig. 69 R), and about 3
times longer than wide at base (Fig. 69 I); ventral plate about as long as wide (Figs.
69 H, S) ................................................................................................................................ 6
2(1) Gonostylus with 3-8 spurs on dorsal external edge to apex (Fig. 65 K) .................
.......................................................................................................................Simulium nahimi
– Gonostylus with 1 spur at apex (Figs. 65 D, F, H, J, L, N, P) .................................. 3
3(2) Gonostylus not acuminate apically and with subapical spur (Figs. 65 J, L) ............
............................................................ Simulium mariavulcanoae, Simulium diversibranchium
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 269

– Gonostylus acuminate and with apical spur (Figs. 65 D, F, G, M, N) .................... 4


4(3) Ventral plate rounded apically (Fig. 65 I) ........................... Simulium subclavibranchium
– Ventral plate acuminate apically (Figs. 65 E, O) ......................................................... 5
5(4) Gonostylus thinning from distal half of its length (Figs. 65 D, P) ...........................
..................................................................................... Simulium inaequale, Simulium rappae
– Gonostylus thinning from distal third of its length (Figs. 65 F, G, H, N) ..............
........ Simulium clavibranchium, Simulium nogueirai, Simulium subnigrum, Simulium travassosi
6(1) Ventral plate well expanded basally (Fig. 69 H) .................. Simulium botulibranchium
– Ventral plate not well expanded basally (Fig. 69 S) .................... Simulium souzalopesi

Pupae
1 Gill with 6 branches generally filiform (Figs. 66 A B, D, G; 67 A-L) .................... 2
– Gill with 2-3 thick branches, frequently asymmetrical and with diverticula (Figs.
66 N; 70 A-C, G-I, P-R) ................................................................................................. 12
2(1) Gill branches thick and about half of cocoon length; secondary branches about as
long as primaries (Fig. 66 I) .................................................................... Simulium rappae
– Gill branches filiform, as long as or longer than cocoon length; secondary
branches longer than primary branches (Figs. 66 A, B, D, G; 67 A-L) .................. 3
3(2) Gill filaments thicker distally (Figs. 67 G, H, I) .......................................................... 4
– Gill filaments not thicker distally (Figs. 67 A-C, F, J, K) ........................................... 5
4(3) Gill filaments greatly thickened distally and thinning abruptly (Figs. 67 G, H);
secondary dorsal and median branches bifurcating far from primary ventral
bifurcation; gill length 3.5-4.2 mm ........................................... Simulium clavibranchium
– Gill filaments slightly thickened distally and thinning gradually (Fig. 67 I);
secondary dorsal and median branches bifurcated close to primary ventral branch;
gill length 4.8-5.5 mm ............................................................ Simulium subclavibranchium
5(3) Gill filaments branching very close to base and at about same distance, with very
short primary and secondary branches (Fig. 67 A) .................................................... 6
– Gill filaments branching at variable and different distances from base, with
variable lengths for primary and secondary branches (Figs. 67 B-F, J, K) ............ 7
6(5) Frontoclypeus and exposed portion of thorax with abundant tubercles ................
....................................................................................... Simulium inaequale, Simulium clarki
– Frontoclypeus and exposed portion of thorax with scarce tubercles ......................
...................................................................................................................... Simulium hauseri
7(5) Gill filaments with dorsal branch thicker than others (Figs. 67 K, L) .................... 8
– Gill filaments of about same diameter ......................................................................... 9
8(7) Primary and secondary dorsal branches much thicker and longer than secondary
ventral branches; secondary ventral branches directed downward and posteriorly
(Fig. 67 L) ................................................................................... Simulium diversibranchium
– Primary and secondary dorsal branches only slightly thicker than and about same
length as secondary ventral branches (Fig. 67 K); secondary ventral branches
directed anteriorly (Fig. 67 K) .................................................. Simulium mariavulcanoae
9(7) Gill longer than cocoon .................................................................................................. 10
– Gill about same length as cocoon ................................................................................ 11
270 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

10(9) Primary ventral branch relatively short; secondary dorsal and median branches
bifurcated at about same level (Fig. 67 B) ..................................... Simulium subnigrum
– Primary ventral branch relatively long, reaching at least to 1/3 of secondary
ventral branch; secondary dorsal and median branches bifurcated at different
levels ....................................................................................................... Simulium leopoldense
11(9) Secondary ventral branch bifurcated at median distance between ventral primary
and secondary dorsal bifurcation (according to D’ANDRETTA & GONZALEZ, Fig. 67
C), or sometimes at same level (PY-DANIEL & MOREIRA, 1989; & STRIEDER, 1998,
Fig. 67 D) .................................................................................................Simulium nogueirai
– Secondary dorsal and ventral branches bifurcated at greater distance than primary
ventral bifurcation and at about same distance between them (Fig. 67 J) ...............
.......................................................................................................................Simulium nahimi
12(1) Gill with 3 branches of about uniform diameter, symmetrical, without diverticula
or branches (Figs. 66 L-N) ..................................................................Simulium travassosi
– Gill with 2 primary branches of variable diameter, asymmetrical, with diverticula
or terminal branches ........................................................................................................ 13
13(12) Gill integument with abundant and large microtrichiae; dorsal branch with 4
relatively thin branches (Figs. 70 P-R); very abundant tubercles on head and
thorax (Fig. 70 S) .............................................................................. Simulium petropoliense
– Gill integument with sparse spiculae and tubercles; dorsal branch with different
apical branches; head and thorax with few tubercles ............................................... 14
14(13) Dorsal branch with 4 terminal diverticula with blunt apices (Fig. 70 C) .................
........................................................................................................ Simulium botulibranchium
– Dorsal branch with 6 apically acuminate branches (Fig. 70 I) ...................................
................................................................................................................ Simulium souzalopesi

CLAVES DE ESPECIES DE SIMULIUM (INAEQUALIUM)


Hembras
Simulium hauseri y S. leopoldense con hembras desconocidas.

1 Paraproctos aproximadamente más largos que altos (Figs. 64 G, L, P) ................. 2


– Paraproctos más altos que largos, aguzados distalmente (Figs. 69 D, Q) .............. 6
2(1) Escudo sin 1+1 banda submediana anterior observada con cierta luz ....................
............................................................ Simulium mariavulcanoae, Simulium diversibranchium
– Escudo con 1+1 banda submediana anterior observada con cierta luz (Fig. 64 A)
................................................................................................................................................ 3
3(2) Coxa I negruzca; fémur y tibia con fuerte coloración negruzca; paraproctos
proporcionalmente bajos (Fig. 64 L) ........................................ Simulium clavibranchium
– Coxa I castaño grisáceo claro; fémures y tibias con escasa coloración negruzca
(Fig. 64 C); paraproctos proporcionalmente más altos (Figs. 64 G, P) .................. 4
4(3) Paraprocto relativamente alto y de borde apical ovoidal (Fig. 64 P); horquilla
genital con el tallo relativamente largo y ramas posteriores proporcionalmente
gruesas (Fig. 64 O) ................................................................................Simulium travassosi
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 271

– Paraprocto relativamente bajo y de borde apical redondeado (Fig. 64 G); horquilla


genital con el tallo relativamente corto, y ramas posteriores proporcionalmente
más finas (Fig. 64 N) ......................................................................................................... 5
5(4) Longitud alas 1,8-2,0 mm ............... Simulium rappae, Simulium clark, Simulium nahimi
– Longitud alas 2,3-2,8 mm ...................................................................................................
....... Simulium inaequale, Simulium nogueirai, Simulium subclavibranchium, Simulium subnigrum
6(1) Gonapófisis con borde interno paralelo (Fig. 69 T) ................. Simulium petropoliense
– Gonapófisis con bode interno divergente (Fig. 69 P) ................................................ 7
7(6) Porción basal del cibario medianamente elevado subcónico (Fig. 69 C) .................
........................................................................................................ Simulium botulibranchium
– Porción basal del cibario escasamente elevada, con pequeña depresión mediana
(Fig. 69 K) ............................................................................................ Simulium souzalopesi

Machos
Simulium hauseri, S. leopoldense y S. petropoliense con machos desconocidos.

1 Gonostilo corto, aproximadamente la mitad del largo del gonocoxito (Figs. 65 C,


G, M) y no más de 2 veces más largo que ancho en la base (Figs. 65 C, D, F, G, H,
J, K, L, N, P); placa ventral más ancha que larga (Figs. 65 C, E, I, O) .................. 2
– Gonostilo alargado, aproximadamente del mismo largo que el gonocoxito (Fig. 69
R), y unas 3 veces más largo que ancho en la base (Fig. 69 I); placa ventral
aproximadamente tan larga como ancha (Figs. 69 H, S) ........................................... 6
2(1) Gonostilo con 3-8 espolones en el borde dorsal externo hasta el ápice (Fig. 65 K)
.......................................................................................................................Simulium nahimi
– Gonostilo con 1 espolón sobre el ápice (Figs. 65 D, F, H, J, N, L, P) ................... 3
3(2) Gonostilo romo apicalmente con espolón subapical (Figs. 65 J, L) .........................
............................................................ Simulium mariavulcanoae, Simulium diversibranchium
– Gonostilo aguzado y con espolón apical (Figs. 65 D, F, G, M, N) ......................... 4
4(3) Placa ventral redondeada apicalmente (Fig. 65 I) ............ Simulium subclavibranchium
– Placa ventral aguzada apicalmente (Figs. 65 E, O) ..................................................... 5
5(4) Gonostilo afinándose desde la mitad distal de su largo (Figs. 65 D, P) ...................
..................................................................................... Simulium inaequale, Simulium rappae
– Gonostilo afinándose desde el tercio distal de su largo (Figs. 65 F, G, H, N) .......
........... Simulium clavibranchium, Simulium nogueirai, Simulium subnigrum, Simulium travassosi
6(1) Placa ventral bien expandida basalmente (Fig. 69 H) ........ Simulium botulibranchium
– Placa ventral no expandida basalmente (Fig. 69 S) ..................... Simulium souzalopesi

Pupas
1 Branquias con 6 ramas generalmente filiforme (Figs. 66 A, B, D, G; 67 A-L) ..... 2
– Branquias con 2-3 ramas gruesas, frecuentemente asimétricas y con divertículos
(Figs. 66 N; 70 A-C, G-I, P-R) ...................................................................................... 12
2(1) Branquias de ramas gruesas, aproximadamente la mitad del largo del capullo;
ramas secundarias aproximadamente tan largas como las primarias (Fig. 66 I) .....
....................................................................................................................... Simulium rappae
272 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

– Branquias de ramas filiformes, tan largas o más que el largo del capullo; ramas
secundarias más largas que las primarias (Figs. 66 A, B, D, G; 67 A-L) ................ 3
3(2) Filamentos de las branquias más gruesos distalmente (Figs. 67 G-I) ..................... 4
– Filamentos de las branquias no más gruesos distalmente (Figs. 67 A-C, F, J, K)
................................................................................................................................................ 5
4(3) Filamentos de las branquias muy engrosados distalmente y afinándose
abruptamente (Figs. 67 G-H); ramas secundaria dorsal y mediana bifurcadas lejos
de la bifurcación primaria ventral; longitud de la branquia 3,5-4,2 mm ..................
.......................................................................................................... Simulium clavibranchium
– Filamentos de la branquia moderadamente engrosados distalmente y afinándose
suavemente (Fig. 67 I); ramas secundaria dorsal y mediana bifurcadas cerca de la
rama primaria ventral; longitud de las branquias 4,8-5,5 mm ....................................
..................................................................................................... Simulium subclavibranchium
5(3) Filamentos de las branquias ramificados muy cerca de la base y aproximadamente a
la misma distancia, mostrando ramas primarias y secundarias muy cortas (Fig. 67 A)
................................................................................................................................................ 6
– Filamentos de las branquias ramificados a diferente distancia de la base,
mostrando ramas primarias y secundarias de variable longitud (Figs. 67 B-F, J, K)
................................................................................................................................................ 7
6(5) Frontoclípeo y porción expuesta del tórax con abundantes tubérculos ..................
....................................................................................... Simulium inaequale, Simulium clarki
– Frontoclípeo y porción expuesta del tórax con escasos tubérculos ..........................
...................................................................................................................... Simulium hauseri
7(5) Filamentos de las branquias con rama dorsal más gruesa que las otras (Figs. 67 K,
L) ........................................................................................................................................... 8
– Filamentos de las branquias aproximadamente del mismo diámetro ..................... 9
8(7) Ramas primaria dorsal y secundaria dorsal notoriamente gruesas y más gruesas
que la secundaria ventral; ramas ventrales secundarias dirigidas hacia abajo y atrás
(Fig. 67 L) ................................................................................... Simulium diversibranchium
– Ramas primaria y secundaria dorsales solamente un poco más gruesas y
aproximadamente de la misma longitud que la secundaria ventral (Fig. 67 K);
ramas ventrales secundarias dirigidas en dirección frontal (Fig. 67 K) ....................
......................................................................................................... Simulium mariavulcanoae
9(7) Branquia más larga que el largo del capullo ............................................................... 10
– Branquia aproximadamente del mismo largo que el capullo .................................. 11
10(9) Rama primaria ventral relativamente corta; ramas secundarias dorsal y media
bifurcadas aproximadamente al mismo nivel (Fig. 67 B) ........... Simulium subnigrum
– Rama primaria ventral relativamente larga, llegando por lo menos a 1/3 de la
secundaria ventral; ramas dorsales y medias bifurcadas a diferentes niveles ..........
................................................................................................................. Simulium leopoldense
11(9) Rama secundaria ventral bifurcada a media distancia entre la bifurcación de la
primaria ventral y la bifurcación de la secundaria dorsal (según D’ANDRETTA &
GONZÁLEZ, 1964), Fig. 67 C), o a veces al mismo nivel (PY-DANIEL & MOREIRA,
1989; STRIEDER, 1998; Fig. 67 D) ........................................................Simulium nogueirai
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 273

– Rama secundaria dorsal y ventral bifurcada a mayor distancia de la bifurcación de


la primaria ventral y al mismo nivel entre ellas (Fig. 67 J) ...............Simulium nahimi
12(1) Branquias con 3 ramas de aproximadamente diámetro uniforme, simétricas, sin
divertículos o ramas (Figs. 66 L-N) ................................................... Simulium travassosi
– Branquias con 2 ramas primarias de diámetro variable, asimétricas, con
divertículos o ramas terminales ..................................................................................... 13
13(12) Tegumento de la branquia con largas y abundantes microtriquias; rama dorsal con
4 ramas relativamente finas (Figs. 70 P-R); cabeza y tórax con abundantes
tubérculos (Fig. 70 S) ....................................................................... Simulium petropoliense
– Tegumento de la branquia con espículas y escasos tubérculos; rama dorsal
presenta diferentes ramas apicales; cabeza y tórax sin o con escasos tubérculos
.............................................................................................................................................. 14
14(13) Rama dorsal con 4 divertículos terminales con ápice romo (Fig. 70 C) ..................
........................................................................................................ Simulium botulibranchium
– Rama dorsal con 6 ramas terminales con ápice aguzado (Fig. 70 I) .........................
................................................................................................................ Simulium souzalopesi

INAEQUALE species group

Simulium (Inaequalium) inaequale (PATERSON & SHANNON)


(Figs. 64 A-G; 65 A-E; 66 A, 67 A; 68 B, C.)
Eusimulium inaequalis PATERSON & SHANNON, 1927: 738-740; PINTO, 1931: 677; VULCANO, 1967: 13.
Simulium inaequalis WYGODZINSKY, 1949: 305; 1953a: 304.
Simulium inaequale VARGAS, 1945a: 147; COSCARÓN & WYGODZINSKY, 1973b: 142.
Simulium (Inaequalium) inaequale COSCARÓN & WYGODZINSKY, 1984: 80-84; COSCARÓN, 1987: 21; 1991: 157-
158; COSCARÓN & COSCARÓN ARIAS, 1997: 111; COSCARÓN & MIRANDA ESQUIVEL, 1998b: 337.
Inaequalium inaequale PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 1998: 44; 2004: 118-119;
STRIEDER & PY-DANIEL, 1999: 51; 2000: 34; 2002: 66.
Trichodagmia manicata ENDERLEIN, 1934b: 288.
Simulium jundiaense D’ANDRETTA & GONZÁLEZ, 1964: 108-110.
Simulium baiensis PINTO, 1931: 685.
Thyrsopelma argentata ENDERLEIN, 1936: 125.
Female: Wing length 2.4-2.7 mm; frons and clypeus grayish pollinose; scutum black
with grayish pollinosity and pilosity gray to tin colored; with some light directions, 1+1
submedian gray pollinose band (Fig. 64 A) that can be thinner and extended to posterior
edge; (these bands can be easier to see if there is no hair); cibarium with basal portion well
elevated medially; sensory vesicle of maxillary palp about 1/3 of palpomere length (Fig. 64
B); legs gray yellowish, with darkened areas dark brown to blackish as shown in figures 64 C,
D, E; calcipala well developed, about as long as wide (Fig. 64 F); genitalia as shown in figure
64 G; paraproct longer than high with abundant microtrichiae and rounded distal edge.
Male: Wing 2.3-2.6 mm; scutum black with golden pilosity; with some illumination.
1+1 submedian light gray pollinose subtriangular spots (Fig. 65 A) and band of gray
pollinosity laterally and posteriorly (Fig. 65 B); legs with coloration as in female but more
274 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

J I

C D
F E

L
N

P
K

Figure 64. Female. A-G: S. (Inaequalium) inaequale; A: scutum, dorsal view; B: third palpomere and
sensory vesicle; C-E: fore, mid and hind leg, respectively; F: calcipala and pedisulcus; G: genitalia in
ventral view: eighth sternite, gonapophyses, spermatheca, genital fork, paraproct and cercus (the
latter in two positions). H: S. (I.) subnigrum, chaetotaxy of wing, anterior border. I-L: S. (I.)
clavibranchium; I: frons; J: fronto-ocular triangle; K: claw; L: paraproct and cercus. M-N: S. (I.)
subclavibranchium; M: basal portion of cibarium; N: genital fork. O-P: S. (I.) travassosi; O: genital fork;
P: paraproct and cercus.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 275

darkened; genitalia as shown in figure 65 C; gonostylus elongated, subtriangular and


robust spur at apex (Fig. 65 D); ventral plate subtriangular (Fig. 65 E); endoparameres
with large teeth.
Pupa: Cocoon length on base 3.3-3.5 mm, on dorsum 3.0-3.2 mm, gill length 4.0-4.2
mm; aspect of pupa in accordance with figure 66 A; gill with 6 similar filamentous
branches emerging very close to base from 3 short primary branches (Fig. 67 A);
frontoclypeus and exposed portion of thorax with abundant and obvious tubercles; head
with 2+2 frontal and 1+1 facial trichomes with 2-4 branches; thoracic trichomes with 4-6
branches; abdomen chaetotaxy as in Simulium subnigrum.
Larva: Maximum length 5.5-7.0 mm; coloration light yellowish-gray, darkened and
greenish gray dorso-laterally; head brown, with light median subbasal window on cephalic
apotome (Fig. 68 B); antennae longer than stem of cephalic fan; ratio of proximal, medial
and distal antennal segments = 1:1.1-1.3:1.2-1.3. Mandible with teeth as explained for
subgenus; anal sclerite with abundant single hairs (Fig. 68 C); anal ring with 62-68 rows of
10-12 hooks each; rectal papillae with 11-16 diverticula on each lobe.
Distribution: Argentina: Jujuy, Misiones, Salta, Tucumán; Bolivia: La Paz; Brazil:
Amapá, Bahia, Espírito Santo, Mato Grosso, Minas Gerais, Paraná, Rio Grande do Sul,
Santa Catarina, São Paulo; Paraguay: from the Cordillera and Alto Paraná.
Bionomics: Immature stages of S. inaequale live in small shallow creeks, mostly on
aquatic vegetation in fast-flowing areas. Females can be anthropophilic and sometimes
pestiferous, as in Santo Inácio (Bahia, Brazil).
Discussion: Arrangement of the gill branches helps differentiate the most similar
species, such as S. subnigrum, with branching at further distances and in different positions. The
most similar species to S. inaequale is S. clarki, with similar pupal gill branching, and adult
ornamentation and morphology, but it is smaller in size. Simulium inaequale could represent
subspecific variation in Panama and Venezuela. Simulium hauseri STRIEDER has very similar pupal
gill branching, and small morphological differences in the number of frontoclypeal and
thoracic tubercles and abdominal chaetotaxy. As the adults and larva are unknown, we do not
have enough information to consider it a variation of S. inaequale or S. subnigrum. Simulium
nogueirai, in accordance with the original illustrations, is also very close to S. inaequale.

Simulium (Inaequalium) clarki FAIRCHILD


Simulium clarki FAIRCHILD, 1940: 703; VARGAS & DÍAZ NÁJERA, 1948a: 75; RAMÍREZ PÉREZ, 1971: 286;
1983: 42; RAMÍREZ PÉREZ et al., 1982: 31.
Simulium (Inaequalium) clarki COSCARÓN, 1983: 21; 1991: 159; SHELLEY et al., 2004: 449, 453, 460.
Inaequalium clarki PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 1998: 29; STRIEDER & PY-
DANIEL, 2000: 26.
This species is very close to S. inaequale; females and males of the type material are
morphologically and ornamentally similar to S. inaequale, only a little smaller (wing length
1.8-1.9 mm) and with lighter legs, more yellowish, and lighter distal blackish areas on the
hind femur, tibia and tarsi. The pupa illustrated by FAIRCHILD of S. clarki shows gill
branching similar to that of S. inaequale. Material from Venezuela determined by RAMÍREZ
PÉREZ (1971-1973) as S. clarki, is similar to the type material, but is larger.
276 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

A
B

H
C F
D

I
M
O

N L K P

Figure 65. Male. A-E: S. (Inaequalium) inaequale; A: scutum, dorsal view; B: scutum, lateral view; C:
genitalia in ventral view: gonocoxite, gonostylus, ventral plate, endoparameres and median sclerite;
D: gonostylus; E: ventral plate in another position than C. F: S. (I.) subnigrum, gonostylus. G-H: S.
(I.) clavibranchium; G: gonocoxite and gonostylus; H: gonostylus in different position. I: S. (I.)
subclavibranchium, ventral plate. J: S. (I.) mariavulcanoae, gonostylus. K: S. (I.) nahimi, gonostylus (from
PY-DANIEL, 1984). L: S. (I.) diversibranchium, gonostylus. M-O: S. (I.) travassosi; M: gonocoxite and
gonostylus; N: gonostylus in different position; O: ventral plate P: S. (I.) rappae, gonostylus; N:
Gonostylus in different position; O: Ventral plate.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 277

B G

D C

M I

E
K

F
J H

Figure 66. Pupa. A: S. (Inaequalium) inaequale, lateral view. B-C: S. (I.) clavibranchium; B: lateral view;
C: frontoclypeus. D-E: S. (I.) subclavibranchium; D: lateral view; E: frontoclypeus. F: S. (I.)
mariavulcanoae, thoracic trichomes. G-H: S. (I.) diversibranchium; G: lateral view; H: thoracic
trichomes. I-K: S. (I.) rappae; I: lateral view; J: frontoclypeus; K: cephalic trichomes. L-N: S. (I.)
travassosi; L-M: dorsal and lateral view, respectively; N: gill.
278 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Female: Wing length 2.3 mm.


Male: Wing length 2.2 mm.
Pupa: Cocoon length on base 3.0 mm, dorsum 2.8 mm, gill filaments 3.3 mm.
Distribution: Panama: Coclé, Darien; Venezuela: Amazonas, Aragua, Bolivar,
Barinas, Carabobo, Cojedes, Guárico, Lara, Miranda, Trujillo.
Bionomics: Breeds in small creeks in temperate to warm water from 150 to 1900 m
altitude. Females bite humans.
Discussion: It will be necessary to study the immature stages at the type locality to
decide if S. clarki is a synonym of S. inaequale or a subspecies of S. inaequale from Central
America, as suggested by WYGODZINSKY (1953), with variation characteristic of a species
with a large distribution.

Simulium (Inaequalium) subnigrum Lutz (Figs. 64 H; 65 F; 67 B, 68 D-F.)


Simulium subnigrum LUTZ, 1910: 239-241; LUTZ & NUNEZ TOVAR, 1928: 43-44; LANE & VULCANO,
1943: 138; VULCANO, 1967: 21.
Simulium (Inaequalium) subnigrum COSCARÓN & WYGODZINSKY, 1984: 90; COSCARÓN, 1987: 21; 1991:
162-163; STRIEDER et al., 1992: 60; COSCARÓN & COSCARÓN ARIAS, 1997: 111.
Inaequalium subnigrum PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 1998: 90; 2004: 135;
2004: 120-121; STRIEDER & PY-DANIEL, 1999: 58; 2000: 66; 2002: 66.
Simulium diversifurcatum LUTZ, 1910: 258-263; LUTZ & MACHADO, 1915: 46.
Simulium mbarigui COSCARÓN & WYGODZINSKY, 1973a: 142-149.
Female: Wing length 2.3-2.8 mm; color similar to that of Simulium inaequale but with
scutum pilosity silky grayish and femur of anterior and middle legs lighter without
blackish apex; cibarium elevated medially with strong teeth covering median area, similar
to Simulium subclavibranchium (Fig. 64 M); sensory vesicle of maxillary palp shorter than
half length of palpomere; wing with one row of hair on Sc and more abundant on basal
sector of R (Fig. 64 H); claw with or without subbasal tooth; genitalia as in Simulium
inaequale, but paraprocts with distal edge shorter as in Simulium clavibranchium (Fig. 64 L).
Male: Wing length 2.2-2.7 mm; coloration and morphology as for Simulium inaequale;
gonostylus thicker and wider subapically than in S. inaequale (Fig. 65 F), and ventral plate
subovoidal with apex well curved , similar to Simulium subclavibranchium.
Pupa: Cocoon length at base 3.3-4.5 mm, dorsally 2.8-3.8 mm, body o f pupa 2.8-3.6
mm, gill filaments 3.8-5.0 mm; cocoon weave thick, with obvious threads, with anterior
reinforcement and without dorsal projection; gill branching at different distances from base:
dorsal and median secondary branches bifurcated at same distance or a little far than ventral
branch (Fig. 67 B) with relative variability of distances in different specimens; frontoclypeus
and exposed portion of thorax with abundant tubercles of medium size; cephalic trichomes
with 2-3 and thoracic with 3-5 branches; abdominal chaetotaxy as shown in Fig. 68 A.
Larva: Maximum length 5.3-6.0 mm; cephalic apotome darkened in basal and
median areas; ratio of proximal, medial and distal antennal segments = 1:1.1:1.3-1.5 (Fig.
68 D). Mandible with one long latero-mandibular process surpassing inferior mandible
rim (Fig. 68 E); cephalic fan with 35-40 rays; rectal papillae with 10-15 diverticula on each
lobe (Fig. 68 F); anal ring with 67-75 rows with 12-14 hooks each.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 279

H
F

K
A J

G
B

I C

Figure 67. Pupa. Gill. A: S. (Inaequalium) inaequale. B: S. (I.) subnigrum. C-E: S. (I.) nogueirai; C:
extracted from D’ANDRETTA & GONZÁLEZ, 1964, D: from STRIEDER, 1998; E: from PY-DANIEL &
MOREIRA, 1989. F: S. (I.) leopoldense. G-H: S. (I.) clavibranchium (H: gill, terminal portion with higher
magnification). I: S. (I.) subclavibranchium. J: S. (I.) nahimi ( PY-DANIEL, 1984). K: S. (I.) mariavulcanoae.
L: S. (I.) diversibranchium.
280 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

G
D E

K
B

M H L

Figure 68. A: Pupa; S. (Inaequalium) subnigrum, chaetotaxy of abdomen. B-M: Larva; B-C: S. (I.)
inaequale; B: head, dorsal view showing cephalic apotome; C: anal sclerite and trichomes; D-F: S. (I.)
subnigrum; D: antenna; E: teeth of mandible; F: rectal papillae; G-J: S. (I.) diversibranchium; G: general
aspect, lateral view; H: head, dorsal view; I: hypostomium and postgenal area; J: gill histoblast; K-L:
S. (I.) rappae; K: general aspect; L: antenna; M: S. (I.) travassosi, head dorsal view.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 281

Distribution: Argentina: Corrientes, Misiones; Brazil: Bahia, Mato Grosso, Minas


Gerais, Paraná, Rio de Janeiro, Rio Grande do Sul, Santa Catarina, São Paulo; Colombia:
Meta; Paraguay: Caaguzu; Trinidad; Venezuela: Bolivar, Distrito Federal, Miranda;
Bionomics: Simulium subnigrum breeds in creeks with clear water and fast current in
open areas. Female feeding habits are unknown.
Discussion: A close species to S. subnigrum is S. inaequale, which occurs in the same
habitats. It shows small differences in adult genitalia but the best character is the branching
of the pupal gill; that of S. inaequale is very close to the base and the emergence of filaments
is at the same distance. Simulium leopoldense has a very similar pattern of gill branching but the
primary ventral branch reaches about the basal third of the secondary ventral branch, and
the secondary dorsal and median branches are bifurcated at different distances. In S.
subnigrum the primary ventral branch is shorter and is bifurcated near the base of the
secondary branches, and the secondary dorsal and median branches are bifurcated at the
same distance. Simulium hauseri also has the gill bifurcation closer to the base and sparser
tubercles on the frontoclypeus and thorax than does S. subnigrum. Simulium nogueirai also is
very similar, but the female scutum pilosity is golden and the pupal gill is shorter with a
length about that of the venter of the cocoon; that of S. subnigrum is longer. The pupa of S.
beaupertuyi has the primary ventral branch longer than the primary dorsal branch, the
bifurcation of the secondary, median and dorsal branches proportionately lower than in S.
subnigrum, and the thoracic tubercles larger in diameter than in S. subnigrum.

Simulium (Inaequalium) nogueirai D’ANDRETTA & GONZÁLEZ (Figs. 67 C-E.)


Simulium nogueirai D’ANDRETTA & GONZÁLEZ, 1964: 103-106; MOREIRA et al., 1994: 525.
Simulium (Inaqualium) nogueirai COSCARÓN, 1987: 21; PY-DANIEL & MOREIRA, 1989: 55; STRIEDER et al.,
1992: 55.
Inaqualium nogueirai PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 1998: 65; 2004: 133-135;
2004: 119-120; STRIEDER & PY-DANIEL, 1999: 54; 2000: 46; 2002: 66.
Adults: Morphology and ornamentation as for INAEQUALE species group, without
peculiar characters that permit separation; based on detailed redescription by PY-DANIEL &
MOREIRA (1989) of Santa Catarina material (type material from São Paulo is lost), female
genitalia as in Simulium inaequale, male gonostylus as in S. diversibranchium and S. clavibranchium,
ventral plate as in S. travassosi; material from Santa Catarina studied by PY-DANIEL &
MOREIRA does not show differences in adults described by D’ANDRETTA & GONZÁLEZ.
Pupa: Cocoon slipper shaped, similar to Simulium subnigrum, but smaller, and
proportionally with shorter gill in relation to cocoon basal length. PY-DANIEL & MOREIRA
(1989), STRIEDER (1998) and STRIEDER & PY-DANIEL (2000) redescribed S. nogueirai from
material of Rio Grande do Sul, Santa Catarina and São Paulo; it shows different levels of
bifurcation of secondary dorsal and median branches, (Fig. 67 D, STRIEDER, 1998 or Fig.
67 E, PY-DANIEL & MOREIRA, 1989) in which median branch is shorter than dorsal; this is
not in accordance with original descriptions, as in figure 67 C, showing approximately
same level of bifurcation.
Larva: Maximum length a little shorter than that of Simulium subnigrum; antennal
segment ratio with medial segment shorter than proximal and distal; anal sclerite with
282 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

single hairs and scales; rectal papillae with 3-6 diverticula on each lobe, anal ring with 77-
83 rows with 11-15 hooks each.
Distribution: Brazil: Paraná, Rio de Janeiro, Rio Grande do Sul, Santa Catarina, São Paulo.
Bionomics: According to PY-DANIEL & MOREIRA (1989), immature stages were
collected in small creeks with or without direct sunshine at 100-300 m altitude. Females
feed on cows, horses and pigs (MOREIRA et al., 1994: 525, STRIEDER et al., 1992: 55).
Discussion: Simulium nogueirai is very close to some other species of the
INAEQUALE species group, especially to S. subnigrum, but can be separated by
characters of the pupal gill and larva. It will be useful to study more material and with
morphological and other methods to determine species validity, because the size of the
pupa and the gill branching can be interpreted as variation in S. subnigrum, which
frequently shows variation of gill bifurcation even in the same specimen.

Simulium (Inaequalium) leopoldense STRIEDER & PY-DANIEL (Fig. 67 F.)


Inaequalium leopoldense STRIEDER & PY-DANIEL, 2000: 14-19; STRIEDER, 2004: 119.
Adults: Unknown
Pupa: Simulium leopoldense is very close to Simulium subnigrum but can be differentiated
by the small size of Simulium leopoldense (cocoon length dorsally 2.4-3.0 mm, basally 2.9-3.2
mm), and gill branching; primary ventral branch three times longer than primary dorsal
branch (Fig. 67 F) (about two times longer in Simulium subnigrum) and primary ventral
branch reaching 1/3 of secondary ventral branch length (in S. subnigrum, about 1/5).
Larva: Small differences also permit Simulium leopoldense to be differentiated from
Simulium subnigrum; in S. leopoldense antenna surpasses length of cephalic fan stem, rectal
papillae with 10-14 diverticula (in S. subnigrum 15-16) and body coloration with darker
dorsal band relative to S. subnigrum.
Distribution: Brazil: Rio Grande do Sul.
Bionomics: Simulium leopoldense lives on aquatic vegetation in small, shallow creeks in
open areas.
Discussion: Simulium leopoldense shows few differences from S. subnigrum and they are
difficult to separate. Simulium nogueirai is also similar, and studies to separate S. nogueirai
from S. subnigrum could be used to separate it from S. leopoldense.

Simulium (Inaequalium) beaupertuyi RAMÍREZ PÉREZ, Rassi & RAMÍREZ


Simulium (Simulium) beaupertuyi RAMÍREZ PÉREZ et al., 1977: 165-177; RAMÍREZ PÉREZ et al., 1982: 29-31.
Simulium beaupertuyi RAMÍREZ PÉREZ, 1983: 29.
Simulium (Inaequalium) beaupertuyi COSCARÓN & WYGODZINSKY, 1984: 84; COSCARÓN, 1987: 21; 1991: 163-164.
Inaequalium beaupertuyi PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 1998: 20; STRIEDER &
PY-DANIEL, 1999: 47; 2000: 19; 2002: 66.
Female: Very similar to S. subnigrum, but a little smaller (wing length 2.2 mm) and
with golden pilosity on scutum.
Male and Pupa: Very similar to Simulium subnigrum, having secondary central and
dorsal gill branches bifurcated at about same distance.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 283

Larva: Very similar to Simulium subnigrum, but with proportionally (a little) shorter
medial antennal segment.
Distribution: Brazil: Roraima; Venezuela: Amazonas.
Bionomics: Simulium beaupertuyi breeds in fast-flowing areas of small creeks with
clear water.
Discussion: As previously explained, S. beaupertuyi is very similar to S. subnigrum and
it is very difficult to separate the two, similar to the case of S. inaequale and S. clarki, having
few morphological differences and living in different regions from the other two well-
known southern species. A comparative study with cytological or molecular methods
could help to resolve the identity of this species.

Simulium (Inaequalium) hauseri (STRIEDER)


Inaequalium hauseri STRIEDER, 1998: 41.
Known only from pupa.
Pupa: Cocoon with gill branching and number of filaments as in Simulium inaequale;
cocoon length ventrally 4.1 mm, on dorsum 3.7 mm, gills 4.2 mm; frontoclypeus and
exposed portion of thorax with scarce rounded tubercles, on head mostly present on basal
area around facial trichomes; on thorax irregularly arranged; cephalic trichomes with 2-4
branches, thoracic trichomes with 4-6 branches.
Distribution: Brazil: Rio Grande do Sul.
Bionomics: This species has been collected in a small creek.
Discussion: Simulium hauseri is very similar to S. inaequale but can be differentiated
by the large number of tubercles on the head and thorax and more basal gill branching.

Simulium (Inaequalium) clavibranchium LUTZ (Figs. 64 I-L; 65 G, H; 66 B, C; 67 G, H.)


Simulium clavibranchium LUTZ, 1910: 257-258; PINTO, 1931: 699; VARGAS, 1945a: 127; VULCANO,
1967: 9.
Simulium (Inaequalium) clavibranchium COSCARÓN & WYGODZINSKY, 1984: 84-87; COSCARÓN, 1987: 21;
1991: 159-160; COSCARÓN & COSCARÓN ARIAS, 1997: 111.
Inaequalium clavibranchium PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 1998: 31; 2004: 133-
135; 2004:117-118; STRIEDER & PY-DANIEL, 1999: 49; 2000: 27; 2002: 66.
Female: Wing length 2.3-2.8 mm; head blackish, palp and proboscis brown blackish,
antenna base light brown; scutum and scutellum black with grayish pollinosity and hair
yellowish-gray with copper tones, homogeneously arranged and not abundant. Metano-
tum velvet black; legs light brown with darkened areas as in Simulium inaequale; frons
slightly convergent (Fig. 64 I); basal portion of cibarium as in Simulium subclavibranchium;
fronto-ocular triangle wider than high (Fig. 64 J); Sc with about 10 hairs; claws with
obvious subbasal tooth (Fig. 64 K); eighth sternite with about 80 setae and gonapophyses
divergent; paraproct short with slight curvature distally (Fig. 64 L).
Male: Wing length 2.3 mm; coloration as in Simulium inaequale but with abdominal
silvery spots reaching segment IX; legs as in S. inaequale but more darkened with fore coxa
blackish and also more darkened femora and tibiae; gonocoxite subtrapezoidal (Fig. 65
284 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

G), gonostylus (Fig. 65 H) relatively shorter than in Simulium inaequale and S. subnigrum;
ventral plate acuminate distally.
Pupa: Cocoon with anterior edge reinforced and perpendicular to base (Fig. 66 B); weave
with open threads obvious; length at base 3.2-3.5 mm, on dorsum 2.8-3.0 mm, gills 3.5-4.2 mm;
gill with primary ventral branch bifurcated at half length of secondary central branch, and
these latter bifurcated at half of secondary dorsal branch (Fig. 67 G ); all filaments enlarged
apically (Fig. 67 H); cephalothoracic trichomes with 3-5 branches; frontoclypeus and exposed
portion of thorax with tubercles in regular quantity and well darkened (Fig. 66 C).
Larva (based on STRIEDER, 1998): Color yellowish-gray; cephalic apotome with
negative spots; antenna longer than stem of cephalic fan; ratio of proximal, medial and
distal antennal segments = 1:0.8-0.9:1.3; cephalic fan with about 35 rays; postgenal cleft
subtriangular; latero-mandibular process approximately straight, not surpassing inferior
rim of mandible; anal ring with 67-76 rows with 11-14 hooks each; rectal papillae with 10-
12 diverticula on each lobe.
Distribution: Brazil: Minas Gerais, Rio de Janeiro, Rio Grande do Sul, Rondônia,
Santa Catarina, São Paulo.
Bionomics: Simulium clavibranchium breeds in swift creeks of crystalline water
flowing over aquatic vegetation or submerged tree branches. Feeding behavior of the
females is unknown.
Discussion: Adults of S. clavibranchium are similar to those of S. inaequale, S.
subclavibranchium and S. subnigrum, although S. clavibranchium has the femora, tibiae and
coxae of the anterior legs darker, the paraproct with a curved distal edge and the
gonostylus with a more flattened distal edge. The subapically inflated gill filaments permit
differentiation from the closest species, S. subclavibranchium, which also has enlarged gill
filaments distally, but with smaller diameter and longer filaments. Another difference
from this species is the proportional length of the secondary dorsal gill branch
bifurcation; that in S. clavibranchium, in relation to the primary ventral branch, is more than
four times the length; the gills also are more sclerotized.

Simulium (Inaequalium) subclavibranchium LUTZ (Figs. 64 M, N; 65 I; 66 D, E; 67 I.)


Simulium subclavibranchium LUTZ, 1910: 260-261; PINTO, 1931: 713; VULCANO, 1967: 21.
Simulium (Inaequalium) subclavibranchium COSCARÓN & WYGODZINSKY, 1984: 87-90; COSCARÓN, 1987:
21; 1991: 160-162; COSCARÓN & COSCARÓN ARIAS, 1997: 111.
Inaequalium subclavibranchium PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 1998: 85; 2002:
133-135; 2004:120; STRIEDER & PY-DANIEL, 1999: 57; 2000: 61; 2002: 66.
Female: Wing length 2.4-2.5 mm; coloration similar to that of Simulium inaequale
with scutum pilosity from grayish to tin color; basal portion of cibarium as in figure 64 M;
fronto-ocular triangle as in Simulium clavibranchium; cercus short, paraproct shorter than
high with rounded edge; genital fork as in figure 64 N; eighth sternite with about 70 setae;
gonapophyses scarce sclerotized on internal edge.
Male: Wing length 2.5 mm; coloration as in Simulium inaequale with variation in dark
color of legs; gonocoxite and gonostylus as in S. clavibranchium; ventral plate not
acuminate apically (Fig. 65 I).
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 285

Pupa: Cocoon similar to Simulium clavibranchium but with thicker weave and threads
more evident; aspect as in figure 66 D, with gill much longer than body of pupa and cocoon;
cocoon length on base 3.2-3.5 mm; on dorsum 2.8-3.2 mm; gill 4.8-5.5 mm; gill with lighter
base than in S. clavibranchium, with secondary dorsal and ventral branches bifurcating nearer
to base than in S. clavibranchium (Fig. 67 I); gill filaments slightly widened distally but
gradually thinning toward apex; frontoclypeal and thoracic trichomes with 3-6 branches;
tubercles abundant, about same as in S. clavibranchium, but a little lighter (Fig. 66 E).
Larva: Maximum length 6.0-6.5 mm; coloration light yellowish-gray, darkened
dorsolaterally in greenish-gray bands; head dark-brown; cephalic apotome darkened on
basal and median area but more darkened than in Simulium inaequale; ratio of proximal,
medial and distal antennal segments = 1:1.1-1.3:1.4-1.5; mandible as in Simulium
diversibranchium; cephalic fan with about 40 rays; anal sclerite as in Simulium inaequale; rectal
papillae with 6-9 diverticula on each lobe; anal ring with 74-86 rows of 13-15 hooks each.
Distribution: Brazil: Minas Gerais, Rio de Janeiro, Rio Grande do Sul, Rondônia,
Santa Catarina, São Paulo.
Bionomics: Simulium subclavibranchium breeds in small creeks, with clear water, on
aquatic vegetation and stones. Feeding behavior of the females is unknown.
Discussion: Imagoes of S. subclavibranchium are similar to those of S. inaequale, S.
subnigrum and S. clavibranchium. The first two species have more pilosity and more grayish
coloration on the scutum; the latter has darkened legs especially on coxa I, more elongated
paraprocts and a shorter gonostylus. A great difference is shown in the pupal gill: Simulium
inaequale and S. subnigrum do not have the distal expansion, and in S. clavibranchium the
distal enlargement is bigger, the gills are proportionally shorter, and the bifurcation of the
secondary dorsal and ventral branches are at a greater distance from the base. Also, the
frontoclypeal and thoracic tubercles of S. clavibranchium are more darkened and abundant.
It is difficult to separate S. subnigrum and S. subclavibranchium pupae when the former has
very long gills that are slightly enlarged distally.

Simulium (Inaequalium) nahimi PY-DANIEL (Figs. 65 K; 67 J.)


Simulium nahimi PY-DANIEL, 1984: 125-137.
Simulium (Inaequalium) nahimi COSCARÓN, 1987: 21; 1991: 164; COSCARÓN & COSCARÓN ARIAS, 1997: 111.
Inaequalium nahimi PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 1998: 60; STRIEDER & PY-
DANIEL, 1999: 53; 2000: 42; 2002: 66.
(Diagnosis based on PY-DANIEL, 1984 and STRIEDER, 1998).
Female: Wing length 1.9-2.0 mm; coloration and morphology similar to that of
Simulium inaequale and S. subnigrum, although scutellum in Simulium nahimi is brown yellowish.
Male: General coloration and morphology similar to that of Simulium inaequale and S.
subnigrum, but Simulium nahimi with gonostylus bearing 3-8 spurs positioned on distal edge
(Fig. 65 K from PY-DANIEL, 1984).
Pupa: Cocoon length on base 3.2-4.0 mm, on dorsum 2.1-3.2 mm; maximum length
of gills 3.6-3.7 mm; primary dorsal branch shorter than primary ventral branch; secondary
dorsal and median branches bifurcated at about same distance, at greater distance than
bifurcation of dorsal primary branch (Fig. 67 J, from PY-DANIEL, 1984).
286 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Larva: Maximum length 5.0; antenna reaching apex of cephalic fan stem; ratio of
proximal, medial and distal antennal segments = 1:0.9-1.03:1.2-1.5. Mandible with latero-
mandibular process; rectal papillae with 15-19 diverticula on each lobe; anal sclerite with
small spiniform setae or filiform hair.
Distribution: Brazil: Mato Grosso.
Bionomics: Breeds in temporary swift springs or creeks in open areas, flowing over
aquatic vegetation or rocks. Female feeding habits are unknown.
Discussion: It is difficult to separate S. nahimi from similar species. The small size
and gonostylus with a greater number of spurs than only one other species allows it to be
readily identified. The number of diverticula on the rectal papillae is the highest for the
species group and is another character that can help distinguish this species.

Simulium (Inaequalium) mariavulcanoae COSCARÓN & WYGODZINSKY (Figs. 65 J; 66 F; 67 K.)


Simulium (Inaequalium) mariavulcanoae COSCARÓN & WYGODZINSKY, 1984: 96; COSCARÓN, 1987: 21;
1991: 166; COSCARÓN & COSCARÓN ARIAS, 1997: 111.
Inaequalium mariavulcanoae PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 1998: 55; STRIEDER
& PY- DANIEL, 1999: 52; 2000: 40; 2002: 66.
Female: Wing length 2.4-2.6 mm; coloration and morphology similar to that of
Simulium diversibranchium; paraproct slightly shorter is the only appreciable morphological
difference.
Male: Wing length 2.3-2.6 mm; similar to Simulium diversibranchium; gonostylus wider
(Fig. 65 J) is the only difference.
Pupa: Cocoon shape and weave structure as in Simulium diversibranchium; length at
base 3.2-3.5 mm, on dorsum 3.0-3.2 mm, gills 3.2-3.6 mm; gills with short basal trunk;
ventral primary branch thin and short, dorsal primary branch also is short but little
thicker; secondary dorsal branch always a little thicker (Fig. 67 K); frontoclypeus and
thorax with abundant and obvious tubercles; trichomes with 2-7 branches with basal
portion curved (Fig. 66 F).
Larva: Unknown.
Distribution: Brazil: São Paulo.
Bionomics: Simulium mariavulcanoae breeding sites and female feeding habits are
unknown.
Discussion: Adults of S. mariavulcanoae are similar to those of S. diversibranchium, but
the more elongated paraproct and narrower gonostylus of this species help differentiate
it. The bifurcation of the pupal gill secondary branches at a greater distance, the thicker
dorsal branch and the more elongated tertiary dorsal branch, as well as the first ventral
secondary branch directed downward and backward, help identify this species.

Simulium (Inaequalium) diversibranchium LUTZ (Figs. 65 L; 66 G, H; 67 L; 68 G-J.)


Simulium diversibranchium LUTZ, 1910: 238-239, 263; PINTO, 1931: 701; VULCANO, 1967: 10.
Simulium (Inaequalium) diversibranchium COSCARÓN & WYGODZINSKY, 1984: 93-96; COSCARÓN, 1987: 21;
1991: 164-166; COSCARÓN & COSCARÓN ARIAS, 1997: 111.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 287

Inaequalium diversibranchium PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 1998: 36; 2004:
118; STRIEDER & PY-DANIEL, 1999: 50; 2000: 31; 2002: 66.
Simulium missionum COSCARÓN, 1976b: 151-154.
Female: Wing length 2.3-2.7 mm; coloration blackish with ornamentation similar to
that in Simulium inaequale species group; frons as in Simulium clavibranchium; fronto-ocular
triangle about as wide as high; scutum pilosity golden to tin colored; metanotum black,
velvety; legs yellowish, with darkened areas blackish and pilosity yellowish to black in
accordance with position of lighting; abdomen brownish-gray with blackish tergal plates on
segments II-IV; paraprocts relatively shortened, with truncate apical edge; eighth sternite
with about 72 setae; gonapophyses slightly sclerotized internally and scarcely divergent.
Male: Wing length 2.3-2.6 mm; color generally as in female; scutum similar to that of
Simulium inaequale; abdominal segments I-II brownish, subsequent segments blackish, with
1+1 silvery spots on tergites II, V, VI; sternites grayish pollinose; gonostylus subtrapezoi-
dal, with strong spur near inner corner (Fig. 65 L).
Pupa: Cocoon with thin weave, with few evident threads, reinforced edge and slowly
projected ventrally (Fig. 66 G); cocoon length on base 3.2-4.0 mm, on dorsum 2.6-3.5 mm,
gills 2.8-3.5 mm; gills with 5-6 filaments emerging from two short primary branches: ventral
branch thinner and bifurcated at short distance; secondary ventral branch curved downward
and posteriorly; primary dorsal branch bifurcated at short distance, with secondary dorsal
branch thicker; secondary dorsal and median branches bifurcated at same distance (Fig. 67
L); frontoclypeus and thorax with abundant tubercles; trichomes of frontoclypeus with 2-6
branches, thoracic trichomes with 5-8 branches and with basal trunk curved (Fig. 66 H).
Larva: Maximum length 6.0-6.8 mm; aspect as in figure 68 G; coloration yellowish gray;
head yellowish brown; cephalic apotome darkened in median basal area (Fig. 68 H); antenna as
long as or slightly shorter than stem of cephalic fan; ratio of proximal, medial and distal
segments = 1:1:1.4; mandible teeth with latero-mandibular process; hypostomium with 4-6
setae on each side; postgenal cleft deep (Fig. 68 I); pupal gill histoblast as in figure 68 J; anal ring
with about 84 rows of 12-13 hooks each; rectal papillae with about 20 diverticula.
Distribution: Brazil: Minas Gerais, Rio de Janeiro, Rio Grande do Sul, Santa
Catarina, São Paulo.
Bionomics: Simulium diversibranchium breeds in small, clear creeks flowing over
submerged aquatic vegetation or trees branches. Female feeding habits are unknown.
Discussion: Adults are difficult to separate from those of the other INAEQUALE
species group. Simulium diversibranchium shows only small differences such as the paraproct
being shorter and gonostylus subtrapezoidal. The asymmetrical gill of S. diversibranchium
permits it to be identified.

Simulium (Inaequalium) rappae PY-DANIEL & COSCARÓN (Figs. 66 I-K; 68 K, L.)


Simulium rappae PY-DANIEL & COSCARÓN, 1982: 155-162.
Simulium (Inaequalium) rappae COSCARÓN, 1987: 21; 1991: 167-168; COSCARÓN & COSCARÓN ARIAS,
1997: 111.
Inaequalium rappae PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 1998: 77; STRIEDER &
PY-DANIEL, 1999: 55; 2000: 55; 2002: 66.
288 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Female: Wing length 1.8-1.9 mm; scutum blackish with pilosity yellowish; with
anterior light, showing median and 1+1 submedian grayish pollinose bands; with posterior
illumination, grayish spots become blackish and black area becomes grayish; Sc with 9-10
hairs; eighth sternite with about 40 setae on each side; gonapophyses sclerotized on
internal edge and divergent; paraproct short as in Simulium clavibranchium and genital fork
as in Simulium travassosi.
Male: Wing length 1.8 mm; scutum black with yellowish pilosity; with frontal
illumination, median third of scutum dark; Sc without setae; length/width ratio of hind
basitarsus = 5.7; gonostylus subtriangular; ventral plate similar to that of Simulium inaequale.
Pupa: Cocoon with thick weave, and reinforced anterior edge (Fig. 66 I); length at
base 2.8-3.6 mm, on dorsum 2.4-3.0 mm, gills 1.6-2.1 mm; gills light brown, with 6 thick
branches emerging from 3 long primary branches bifurcated at about same distance;
frontoclypeus and thorax with scarce tubercles (Fig. 66 J); cephalic trichomes of 2-3 long
branches and thoracic trichomes with 3-5 branches (Fig. 66 K).
Larva: Maximum length 4.2-5.0 mm; general aspect as in figure 68 K; postgenal cleft
deep, with postgenal bridge shorter than hypostomium length; antenna as in figure 68 L,
ratio of proximal, medial and distal antennal segments = 1:0.7-1:1.2-1.7; anal ring with 64
rows with 12-15 hooks each; rectal papillae with 13-17 diverticula on each lobe.
Distribution: Brazil: Goiás, Minas Gerais, Rio de Janeiro, São Paulo.
Bionomics: Simulium rappae breeds in clear creeks, on aquatic vegetation in fast
current and in open areas. Feeding habits of the females are unknown.
Discussion: Simulium rappae adults are very similar to those of the INAEQUALE
species group, with very few differences seen with different directions of illumination.
The pupal gill of S. rappae is unique and allows the species to be identified.

Simulium (Inaequalium) travassosi D’ANDRETTA & D’ANDRETTA


(Figs. 64 O, P; 65 M, O; 66 L-N; 68 M.)
Simulium travassosi D’ANDRETTA & D’ANDRETTA, 1947: 161-179; VULCANO, 1967: 22.
Simulium (Inaequalium) travassosi COSCARÓN & WYGODZINSKY, 1984: 96-98; COSCARÓN, 1987: 22; 1991:
168-169; STRIEDER et al., 1992: 59; COSCARÓN & COSCARÓN ARIAS, 1997: 111.
Inaequalium travassosi PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 1998: 97; 2004: 121;
STRIEDER & PY-DANIEL, 1999: 59; 2000: 71; 2002: 66.
Simulium botulibranchium LUTZ, 1910: 256; PINTO, 1931: 675.
Female: Wing length 2.2 mm; scutum blackish with golden dispersed hair; abdomen
blackish, tergite I bright silvery with yellowish setae on posterior edge; tergite II blackish
brown, bright silvery laterally; tergites III and V black matte, remaining tergites with
cereous sheen; legs brown with darkened area blackish; fronto-ocular triangle deep;
genital fork with elongate stem (Fig. 64 O); paraproct slightly higher than long with
convex edge (Fig. 64 P); gonapophysis as in Simulium inaequale.
Male: Wing length 2.1 mm; coloration as in Simulium inaequale; Sc with 3-4 hairs;
gonocoxite about two times longer than gonostylus (Fig. 65 M); gonostylus short,
subtrapezoidal, distally elongated, with strong apical spur (Fig. 65 N); ventral plate wider
than long, with distal edge subacute (Fig. 65 O).
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 289

Pupa: Cocoon with thin weave, threads evident, with straight reinforced anterior
edge (Figs. 66 L, M); length at base 3.2-3.6 mm, on dorsum 2.9-3.2 mm, gills 1.5-1.8 mm;
gills each of three branches with microscopical granulosity; dorsal branch largest, showing
shape variation, with 1 or 2 small apical processes (Fig. 66 N); frontoclypeus and exposed
portion of thorax with abundant tubercles; cephalic and thoracic trichomes with 2-4
branches.
Larva: Maximum length 5.0-5.2 mm; color yellowish brown with tones greenish-gray
dorso-laterally on abdomen; cephalic apotome with diffuse ornamentation (Fig. 68 M);
antennal apex surpassing cephalic fan stem; ratio of proximal, medial and distal antennal
segments = 1:0.9:1.6; mandible with latero-mandibular process surpassing inferior edge;
cephalic fan with about 35 rays; hypostomium with 4 setae on each side; anal ring with
about 78 rows with 15 hooks each; anal sclerite dorsally with 3-4 rows of scales with 2.5
spines each; rectal papillae with 3-7 diverticula on each lobe.
Distribution: Argentina: Misiones; Brazil: Goiás, Minas Gerais, Rio de Janeiro, Rio
Grande do Sul, Santa Catarina, São Paulo.
Bionomics: Simulium travassosi breeds on aquatic vegetation in small fast-flowing
creeks. Female feeding habits are unknown.
Discussion: Simulium travassosi adults are very difficult to separate from those of
similar species such as S. inaequale, S. subnigrum and S. subclavibranchium, showing
differences only in the genital fork and gonostylus. However, the characteristic pupal gill
morphology allows it to be identified.

BOTULIBRANCHIUM species group

Simulium (Inaequalium) botulibranchium LUTZ (Figs. 69 A-I; 70 A-F.)


Simulium (Inaequalium) botulibranchium LUTZ, 1910: 256-257, 263; D’ANDRETTA & D’ANDRETTA, 1947:
145; VULCANO, 1967: 7; 19; COSCARÓN, 1980: 293-295.
Simulium (Inaequalium) botulibranchium COSCARÓN, 1987: 22; 1991: 170-171; COSCARÓN & COSCARÓN
ARIAS, 1997: 111.
Simulium tulibranchium LUTZ, 1922: 92 (error).
Inaequalium botulibranchium PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 1998: 24; 2004:
117; STRIEDER & PY-DANIEL, 1999: 48; 2000: 22; 2002: 66.
Female: Wing length 2.1-2.6 mm; scutum blackish with abundant pilosity tin-colored
to golden; scutellum dark brown; metanotum and pleurae blackish with nacreous pollinosity;
abdomen black, tergite I laterally with grayish pollinosity; tergites II-III black matte, showing
1+1 postero-lateral whitish spots on tergite II; remaining tergites with cereous sheen; frons
convergent and relatively narrow (Fig. 69 A); fronto-ocular triangle deep (Fig. 69 B);
cibarium with basal portion elevated and with thick spines (Fig. 69 C); Sc with about 20
setae; paraprocts subtriangular, about two times higher than wide and with basal posterior
lobe (Fig. 69 D); gonapophysis divergent, with edge slightly sclerotized; genital fork with
elongated stem, longer than two posterior branches combined (Fig. 69 E).
Male: Wing length 2.1-2.4 mm; general coloration as in female, but more darkened,
especially legs, with more abundant black hair; scutum black, with some directions of
290 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

B
A J D

C K G

F P

L
N M

O S
H

Figure 69. A-I: S. (Inaequalium) botulibranchium; A-E: Female; A: frons; B: fronto-ocular triangle; C:
basal portion of cibarium; D: cercus and paraproct; E: genital fork; F-I: Male; F: scutum and
metanotum; G: abdomen, lateral view; H: ventral plate; I: gonostylus. J-S: S. (I.) souzalopesi; J-Q:
Female; J: frons; K: basal portion of cibarium; L-N: fore, mid and hind leg, respectively; O: claw; P:
eighth sternite and gonapophyses; Q: cercus and paraproct; R-S: Male; R: gonocoxite and
gonostylus; S: ventral plate. T: S. (I.) petropoliense, eighth sternite and gonapophyses.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 291

illumination, silvery spot of variable size on antero-lateral area (Fig. 69 F); abdomen
blackish with light silvery spots as shown in figure 69 G; ventral plate about as long as
wide, slightly acuminate distally (Fig. 69 H).Gonostylus elongated about as long as
gonocoxite, with external lateral curvature and apical spur (Fig. 69 I).
Pupa: Cocoon of thin weave, sparse threads, with straight and reinforced edge (Figs.
70 A, B); length at base 2.6-3.0 mm, on dorsum 2.1-2.6 mm, gill 1.3-1.5 mm; gills very
peculiar, each of 2 thick branches: one antero-lateral and one ventral; the first has 2 short
basal branches and 4 on distal half, the ventral has several curvatures at different levels
and no secondary branches (Fig. 70 C); gill integument with small spiculae; frontoclypeus
and thorax with scarce or no tubercles; cephalic trichomes bifid, thoracic trichomes with
2-6 branches; abdominal chaetotaxy as Simulium inaequale.
Larva: Maximum length 6.0-6.5 mm; color grayish-brown with greenish-gray dorsolat-
erally on abdominal segments, aspect as in figure 70 D; antennae surpassing apex of cephalic
fan stem; ratio of proximal, medial and distal antennal segments = 1:0.8-0.9:1.2-1.6; latero-
mandibular process sinuous and surpassing inferior dental edge; cephalic apotome scarcely
ornamented, with negative spots; cephalic fan with 40-46 rays; hypostomium with 5-7 lateral
setae (Fig. 70 E); postgenal cleft subtriangular, postgenal bridge about as long as
hypostomium length (Fig. 70 F); anal sclerite with single hairs; anal ring with 80-92 rows of
14-15 hooks each; rectal papillae with 2-9 diverticula on each lobe.
Distribution: Argentina: Misiones; Brazil: Rio de Janeiro, Rio Grande do Sul, Santa
Catarina, São Paulo.
Bionomics: Simulium botulibranchium breeds in the fast flow of small, clear creeks
with aquatic vegetation. Feeding habits of females are unknown.
Discussion: Adults of Simulium (Inaequalium) are very similar externally. Small
differences in the shape of the paraproct, gonostylus and ventral plate help differentiate
them. The peculiar pupal gill is different from that of any other known Neotropical
simuliid.

Simulium (Inaequalium) souzalopesi COSCARÓN (Figs. 69 J-S; 70 G-O.)


Simulium souzalopesi COSCARÓN, 1980: 296-298.
Simulium (Inaequalium) souzalopesi COSCARÓN, 1987: 23; 1991: 171-172; COSCARÓN & COSCARÓN ARIAS,
1997: 111.
Inaequalium souzalopesi PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 1998: 81; STRIEDER &
PY- DANIEL, 1999: 56; 2000: 58; 2002: 66.
Female: Wing length 2.3-2.5 mm; coloration and morphology similar to that of
Simulium botulibranchium; frons a little wider than in Simulium botulibranchium (Fig. 69 J); Sc
with about 10 hairs; cibarium with abundant spines, with small median depression at apex
(Fig. 69 K); aspect of fore mid and hind legs as in figures 69 L, M, N; claw with well-
developed subbasal tooth (Fig. 69 O); eighth sternite with about 60 setae; gonapophyses
divergent, slightly sclerotized on internal margin (Fig. 69 P); paraprocts subtriangular,
about as high as wide and with small protuberance at base (Fig. 69 Q).
Male: Wing length 2.2 mm; coloration and morphology, including genitalia, similar
to that of Simulium botulibranchium; gonocoxite subtrapezoidal with same length as
292 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

C
A

B
I

H
R

J
P

Q M

E S

O
D

L F
N

Figure 70. A-F: S. (Inaequalium) botulibranchium; A-C: Pupa; A: lateral view; B: dorsal view; C: gill; D-
F: Larva; D: lateral view; E: hypostomium; F: hypostomium and postgenal area. G-O: S. (I.)
souzalopesi; G-J: Pupa; G: lateral view; H: dorsal view; I: gill; J: frontoclypeus; K-O: Larva; K:
antenna; L: teeth of mandible; M: head, dorsal view; N: hypostomium and postgenal area; O: anal
sclerite. P-S: S. (I.) petropoliensye; P-S: Pupa; P: lateral view; Q: dorsal view; R: gill; S: frontoclypeus.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 293

gonostylus; gonostylus elongated, with ventral subapical curvature externo-laterally


positioned (Fig. 69 R); ventral plate relatively narrow basally (Fig. 69 S).
Pupa: Cocoon with thin weave and evident threads, slipper shaped with dorsal projection
(Figs. 70 G-H); length on dorsum 3.0-3.4 mm; on base 2.8-3.0 mm; gills 2.0-2.1 mm; gills
asymmetrical, relatively thickened, each with 6 terminal branches; two primary branches
emerging from short basal trunk; dorsal primary branch bifurcated at short distance dorso-
laterally; another dorsal secondary branch with strong curvature and bifurcated (Fig. 70 I); gill
integument process with small-spaced tubercles; integument of head and exposed portion of
thorax with scarce tubercles (Fig. 70 J); frontoclypeal trichomes simple, thoracic trichomes
single or bifid; abdominal chaetotaxy as in Simulium subnigrum.
Larva: Maximum length 5.5-6.2 mm; coloration light grayish-brown with greenish-
brown on thorax dorsally and abdomen dorso-laterally; head light brown, cephalic
apotome with diffuse to positive spots, darkened on median and basal area (Fig. 70 M);
cervical sclerites well developed; hypostomium with 5-6 setae on each side; postgenal cleft
not deep, with length about half of hypostomium (Fig. 70 N); ratio of proximal, medial
and distal antennal segments = 1:0.8:1.3 (Fig, 70 K); teeth of mandible as figure 70 L;
cephalic fan with about 35 rays; anal ring with about 75 rows with 13 hooks each; anal
sclerite with abundant simple hairs, without scales (Fig. 70 O).
Distribution: Brazil: Rio de Janeiro.
Bionomics: Simulium souzalopesi was collected on aquatic vegetation and stones in
small currents about 0.30-0.50 m wide and 0.05-0.10 m deep, with crystalline water, living
with S. botulibranchium and S. petropoliense. Feeding habits of the females are unknown.
Discussion: Simulium souzalopesi adults are similar to those of S. botulibranchium,
showing small differences such as a narrow frons, the fronto-ocular triangle more acute
and the cibarium elevated medially. Differences are obvious in the pupal gill, which has a
unique morphology not seen in other Neotropical species.

Simulium (Inaequalium) petropoliense COSCARÓN (Figs. 69 T; 70 P-S.)


Simulium petropoliense COSCARÓN, 1980: 298-301.
Simulium (Inaequalium) petropoliense COSCARÓN, 1987: 23; 1991: 172-173; COSCARÓN & COSCARÓN
ARIAS, 1997: 111.
Inaequalium petropoliense PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 1998: 72; STRIEDER &
PY- DANIEL, 1999: 55; 2000: 52; 2002: 66.
Female: Wing length 2.6 mm; coloration and external morphology as in Simulium
botulibranchium; Sc with about 16 hairs; eighth sternite with about 25 setae on each side;
gonapophyses with internal borders separated and parallel (Fig. 69 T); paraprocts as in
Simulium souzalopesi.
Male: Unknown.
Pupa: Cocoon with compact weave and very close threads, with reinforced anterior
edge and without dorsal anterior projection (Figs.70 P, Q); length on dorsum 2.7 mm; on
base 3.1 mm; gills 1.1 mm; gill asymmetrical with 2 wide primary branches; dorsal branch
with 4 short thin terminal branches and ventral branch with several curves, undivided
distally (Fig. 70 R); gill integument with abundant thin microscopical projections like hair;
294 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

head and exposed portion of thorax with abundant tubercles (Fig. 70 S); frontoclypeal
trichomes with 2-4 branches, thoracic trichomes with 4-6 branches; thorax with dark
brown tones not uniformly distributed (Fig. 70 R).
Larva: Unknown.
Distribution: Brazil: Rio de Janeiro.
Bionomics: Simulium petropoliense is similar to S. souzalopesi, which was collected in the
same place.
Discussion: The peculiar pupal gill allows S. petropoliense to be differentiated from
the other Neotropical species.

Species inquirendae

Simulium parimaensis RAMÍREZ PÉREZ, YARZÁBAL, TAKAOKA, TADA & RAMÍREZ, 1986: 61.
Simulium (Inaequalium) parimaensis COSCARÓN, 1991: 155.
Inaequalium parimaensis STRIEDER, 1998: 71; STRIEDER & PY-DANIEL, 1999: 46; 2000: 51.
Simulium (Psaroniocompsa) parimaensis CROSSKEY & HOWARD, 1997: 58; 2004: 52.
The female description of S. parimaensis corresponds to any Simulium (Inaequalium)
species. (As explained before, it is difficult to establish the species identity in this subgenus
with only adult material). The pupal gill has 18-20 branches, and is different from known
Simulium (Inaequalium) species and closer to Simulium (Ectemnaspis) species. Based on the
pupae, it is similar to S. parimaensis from Roraima (Brazil), which is close to the type
locality in the Federal Territory of Amazonia (Venezuela). STRIEDER (1998) and STRIEDER
& PY-DANIEL (2000) verify that females obtained by dissection correspond to the genus
Ectemnaspis. The authors believe a mistake was made when the holotype was mounted on
the slide. Based on this assertion, and pending new material reared from the type locality,
we maintain S. parimaensis as species inquirenda.
Distribution: Venezuela: Amazonas.

Simulium luribayae SMART


Simulium luribayae SMART, 1944: 132 n. nom; for Trichodagmia angustifrons ENDERLEIN, 1934b: 288.
Simulium (Inaequalium) luribayae COSCARÓN & WYGODZINSKY, 1984: 22; 1991: 173.
The type deposited at the Berlin Museum is externally similar to S. subnigrum but with
more greenish scutum pilosity. Without knowledge of the pupa, it is impossible to define
this species.

Simulium pseudoexiguum NUNES DE MELLO & DE ALMEIDA


Simulium pseudoexiguum NUNES DE MELLO & DE ALMEIDA, 1974: 65-68.
Simulium pseudoexiguum RAMIREZ PÉREZ et al., 1986: 63.
Simulium (Inaequalium) pseudoexiguum COSCARÓN, 1987: 22.
As this species was described only from a female, it is impossible to separate it from
other species of the subgenus. We consider S. pseudoexiguum as species inquirenda.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 295

Simulium (Psaroniocompsa) ENDERLEIN (Figs. 71-75; 142.)


Type species: Psaroniocompsa opalinifrons ENDERLEIN, 1934a: 192 (= S. incrustatum LUTZ, 1910: 243).
Syn. Pliodasina ENDERLEIN, 1936: 124 (Type species Pliodasina guttata ENDERLEIN, 1936: 124).
Female: Medium-sized to small species. General coloration black; frons and clypeus
grayish silvery pollinose or with nacreous sheen with greenish-blue tones. Scutum velvety
black with silvery lateral and posterior margins, and generally with 1+1 silvery to blackish,
according to direction of light, short cuneiform anterior submedian spots that can be
continuous posteriorly as thin silvery stripe, sometimes reaching posterior border. Scutum
with homogeneous pilosity, gray to bronze; sometimes hairs grouped, simulating scales
with golden to violaceous greenish sheen. Metanotum black with velvety gray pollinosity.
Abdomen black with tergites I-V with velvety tones, sometimes showing 1+1 small
sublateral silvery spots on tergite II; tergites VI-IX with cereous sheen. Frons relatively
wide; fronto-ocular triangle scarcely developed. Basal portion of cibarium with sclero-
tized rim and teeth arranged in several rows reaching cornuae base at middle, glabrous and
depressed. Basal portion of R generally without hairs; Sc with or without hairs. Legs with
single, spatulate hair. Calcipala well developed. Claws generally with small subbasal tooth.
Fore tarsus relatively wide. Genital fork with elongated stem; gonapophyses with
subparallel internal border, scarcely sclerotized; eighth sternite with about 40 setae;
paraproct short with rounded border.
Male: Scutum velvety black with hairs as in female. Pronotal areas grayish dark
with silvery lateral and posterior margins; with anterior light source, 1+1 silvery
submedian small cuneiform spots. Abdomen black with 1+1 silvery spots on tergites II,
VI and VII and another small spot on lateral border of VIII. Gonocoxite subtrapezoi-
dal, about as long as wide; gonostylus shorter than gonocoxite, subquadrate or
subtrapezoidal to subtriangular, with great submedian to apical spur. Ventral plate
subtriangular, wider than long.
Pupa: Cocoon with thick or thin weave, with anterior border from straight to
dorsally projected. Gill generally with 6 filaments, exceptionally with 4 or 2 thicker ones.
Frontoclypeus and exposed portion of thorax with platelets from abundant to reduced
or absent. Head trichomes with one frontal pair and 2 facial pairs; thorax with 5 pairs,
generally bifid but can have 5-12 branches in some species. Abdomen with spine combs
on distal tergites.
Larva: Body without hairs or scales or these very short to well developed on
abdomen. Cephalic apotome generally with positive spots, discontinuous, arranged in
median and 1+1 sublateral groups. Cephalic capsule with short trichomes, generally bifid.
Postgenal cleft deep, with postgenal bridge shorter than half of hypostomial length.
Antenna surpassing apex of cephalic fan stem, with median segment generally shorter
than fore and hind segments. Mandible with 2 marginal teeth and generally with 1-2
latero-mandibular teeth surpassing level of tooth apex. Hypostomium median tooth
about as high as lateral teeth. Cephalic fan with about 35 rays. Anal sclerite generally with
scarce simple hairs. Anal ring with about 65 rows with about 12 hooks each. Ninth
segment with 1 pair of ventral tubercles. Rectal papillae with 3-8 diverticula on each lobe.
Distribution: From Argentina to Panama (Fig. 142).
296 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

The subgenus S. (Psaroniocompsa) includes 18 species that can be arranged in 2 species


groups:
• INCRUSTATUM: with S. anamariae, S. angrense, S. auripellitum, S. auristriatum, S.
bonaerense, S. brevifurcatum, S. fuliginis, S. incrustatum, S. jujuyense, S. limbatum, S.
minuanum, S. schmidtmummi and S. stellatum.
• SIOLII: with S. damascene, S. guaporense, S. lourencoi, S. siolii, S. tergospinosum.
The first group has species with the scutum scarcely silvery ornamented, from absent
or reduced to 1+1 anterior submedian spots that can be continued posteriorly by thin
stripes not joining with the posterior grayish area; and the male gonostylus smooth
without large hairs or scales.
The second group is differentiated by having a scutum with wider 1+1 submedian
silvery stripes; male gonostylus subtriangular with a dorsal longitudinal carina, and larvae
with dorsal protuberances on the abdominal tergites and abundant and large hairs and
scales on the integument.
Species arrangement in the text was done considering those with more generalized
morphology first, showing scutal hairs homogeneously distributed, scarce silvery spots or
longitudinal stripes and pollinose frons without shiny iridescent pigmentation.
CROSSKEY & HOWARD (1997) include in this subgenus, the species in the AMAZONI-
CUM, AURISTRIATUM, INCRUSTATUM, QUADRIFIDUM, and SIOLII groups. The
INCRUSTATUM, AURISTRIATUM and SIOLII groups treated here correspond to the
subgenus S. (Psaroniocompsa), AMAZONICUM to S. (Cerqueirellum) PY-DANIEL and
QUADRIFIDUM to S. (Coscaroniellum) PY-DANIEL.

KEYS TO SIMULIUM (PSARONIOCOMPSA) SPECIES

Key to larvae not included.

Females and males


1 Scutum hairs grouped, scale like (Fig. 71 K) frequently arranged in longitudinal
rows, at least on median and submedian rows (Figs. 71 L, O, Q); frons and clypeus
silvery with greenish-blue to violaceous sheen ............................................................ 2
– Scutum hairs homogeneously distributed, not grouped or scale like; frons and
clypeus with grayish pollinosity, with velvety sheen ................................................... 7
2(1) Small species, wing length about 1.8 mm; scutum without silvery submedian
cuneiform spots or longitudinal bands ............................................... Simulium fuliginis
– Medium-sized species, wing length >2.0 mm; scutum with or without silvery
spots, if wing length <2.0 mm, scutum with 1+1 submedian longitudinal bands
................................................................................................................................................ 3
3(2) Scutum without silvery anterior submedian cuneiform spots; hairs grouped like
isolated scales arranged in 1 median and 2 submedian rows (Fig. 71 Q) .............. 4
– Scutum with 1+1 silvery anterior submedian cuneiform spots; sometimes
prolonged posteriorly, reaching grayish posterior area (Figs. 71 L, O); hairs
grouped, appearing like scales sometimes not isolated nor arranged in well-
defined rows ........................................................................................................................ 6
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 297

4(3) Scutum hair bright golden ................................ Simulium anamariae, Simulium stellatum
– Scutum hair shiny greenish violet or golden-violet .................................................... 5
5(4) Female: frons wider than opposite option with parallel sides; fronto-ocular
triangle scarcely deep (Fig. 71 R). Male: gonostylus spur very close to inner angle
(Fig. 72 H) .......................................................................................... Simulium auristriatum
– Female: frons relatively narrow with convergent sides (Fig. 71 S); fronto-ocular
triangle relatively deep (Fig. 71 T). Male: gonostylus spur almost halfway to edge
............................................................................................................ Simulium brevifurcatum
6(3) Scutum with 1+1 silvery anterior cuneiform spots not prolonged posteriorly as stripes
(Fig. 71 L), if prolonged, they do not surpass anterior 1/4 of scutum; scutum hairs
golden-bronze or golden-green ............. Simulium incrustatum, Simulium schmidtmummi
– Scutum with 1+1 silvery cuneiform spots prolonged posteriorly as grayish
pollinose stripe that reaches posterior grayish area (Fig. 71 O); scutum hairs pale
greenish gray ..........................................................................................Simulium limbatum
7(1) Scutum without 1+1 silvery submedian longitudinal stripes; pilosity very
abundant with silky appearance. Male gonostylus subtrapezoidal (Figs. 72 A, F)
................................................................................................................................................ 9
– Scutum with 1+1 wide silvery submedian longitudinal bands that extend to
posterior grayish area in females or can be shorter and not surpass half length of
scutum in males (Figs. 72 J, K); pilosity scarce, without silky appearance; male
gonostylus subtriangular with longitudinal carina (Figs. 72 L, M).
Female ................................ Simulium siolii, Simulium tergospinosum, Simulium damascenoi
Male ....................................................................................................................................... 8
8(7) Scutum with 1+1 silvery submedian, cuneiform spots (Fig. 72 K) ..........................
..................................................................................Simulium siolii, Simulium tergospinosum
– Scutum with 1+1 silvery submedian, subtrapezoidal spots (Fig. 72 J) .....................
................................................................................................................ Simulium damascenoi
9(8) Large species, wing length: female 2.5-2.8 mm, male 2.1-2.6 mm; scutum pilosity
very abundant hiding anterior silvery cuneiform spots; scutum hairs from pale
greenish yellow to bronze .................................................................... Simulium angrense
– Medium-sized species, wing length: female 1.7-2.5 mm, male 1.7-2.4 mm ......... 10
10(9) Claws without subbasal tooth (Fig. 71 I) ....................................... Simulium minuanum
– Claws with subbasal tooth .............................................................................................. 11
11(10) Scutum pilosity predominantly bronze to yellowish gray-green; wing length:
female 1.7-2.1 mm, male 1.6-2.0 mm ........................................... Simulium auripellitum
– Scutum pilosity predominant grayish; wing length: female 2.1-2.5 mm, male 1.9-
2.4 mm ...................................................................Simulium jujuyense, Simulium bonaerense

Pupae
1 Gill with 2 thick subcylindrical branches (Fig. 73 J) ............... Simulium auristriatum
– Gill with 4-6 thin filamentous branches ........................................................................ 2
2(1) Gill with 4 branches (Figs. 73 K, 74 H) ........................................................................ 3
– Gill with 6 branches (Figs. 74 B-G) ............................................................................... 8
3(2) Cocoon with well-developed dorsal anterior projection (Fig. 73 K) ...................... 4
298 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

– Cocoon without dorsal anterior projection (Fig. 73 N) or very short (Figs. 73 L, M)


................................................................................................................................................ 5
4(3) Cephalic and thoracic trichomes with 2 branches ..........................Simulium stellatum
– Cephalic trichomes with 1-4 branches, thoracic with 4-12 branches .......................
.......................................................................................................... Simulium schmidtmummi
5(3) Gill shorter than cocoon length (Figs. 73 M, N); primary branches bifurcating
close to base (Fig. 74 I), frontoclypeus without tubercles ........................................ 6
– Gill longer than cocoon length (Fig. 73 L); primary branches bifurcating relatively
far from base, frontoclypeus with tubercles ................................................................. 7
6(5) Cocoon without dorsal anterior projection and with very short ventral projection
(Fig. 73 M) ........................................................................................... Simulium damascenoi
– Cocoon with short dorsal and relatively long ventral projection (Fig. 73 N) .........
............................................................................................................ Simulium tergospinosum
7(5) With sclerotized areas on abdominal membrane .................................. Simulium siolii
– Without sclerotized areas on abdominal membrane ...................... Simulium lourencoi
8(2) Cocoon with dorsal anterior projection (Figs. 73 H, J) ............................................. 9
– Cocoon without dorsal anterior projection (Figs. 73 A-G) .................................... 10
9(8) Cocoon and gills expanded laterally; anterior dorsal projection relatively short
(Fig. 73 I) ......................................................................................... Simulium brevifurcatum
– Cocoon and gills not expanded laterally; anterior dorsal projection very elongated
and bifurcated, protecting gills laterally (Fig. 73 H) .................... Simulium anamariae
10(8) Cocoon with open weave, with threads very evident; frontoclypeal and thoracic
trichomes with 3-4 branches ............................................................. Simulium guaporense
– Cocoon with compact weave, with threads not very evident; frontoclypeal and
thoracic trichomes with 2 branches (Fig. 74 J) .......................................................... 11
11(10) Gill bifurcation of primary ventral and secondary branches very close to base
(Fig. 74 C) ............................................................................................. Simulium bonaerense
– Gill bifurcation of primary ventral and secondary branches not very close to base
(Figs. 74 B, D, E-G) ......................................................................................................... 12
12(11) Pupa large; gill length 3.1-4.5 mm; frontoclypeus and thorax with abundant (Fig.
73 P) to moderately abundant tubercles (Fig. 73 Q) ................................................ 13
– Pupa medium sized; gill length 2.1-4.0 mm, frontoclypeus and thorax with sparse
tubercles (Fig. 73 O) ........................................................................................................ 14
13(12) Frontoclypeus and thorax with abundant tubercles and trichomes moderately long
(Fig. 73 P) ................................................................................................ Simulium angrense
– Frontoclypeus and thorax with relatively abundant tubercles and trichomes
proportionately longer (Fig. 73 Q) ................................................ Simulium incrustatum
14(12) Primary ventral and secondary gill branches bifurcated relatively close to base
(Fig. 74 F) ...............................................................................................Simulium limbatum
– Primary ventral and secondary gill branches bifurcated farther from base (near
two times than the opposite option ) (Figs. 74 D, E, G) ......................................... 15
15(14) Gill in profile approximately straight (Fig. 73 C) ............................ Simulium jujuyense
– Gill in profile with slight median ventral curvature (Figs. 73 A, B) ...................... 16
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 299

16(15) Secondary gill branches with bifurcation surpassing 1/3 of gill length (Fig. 73 A)
...............................................................................................................Simulium auripellitum
– Secondary gill branches with bifurcation surpassing 1/4 of gill length (Fig. 73 B)
................................................................................................................. Simulium minuanum

CLAVES DE ESPECIES DE SIMULIUM (PSARONIOCOMPSA)


Clave para larvas no incluida.

Hembras y machos
1 Pelos del escudo agrupados con aspecto de escamas (Fig. 71 K) frecuentemente
ordenados en hileras longitudinales, por lo menos en las hileras medianas y
submedianas (Figs. 71 L, O, Q); frente y clípeo plateado con brillo verdoso
azulado a violáceo .............................................................................................................. 2
– Pelos del escudo distribuidos homogeneamente no agrupados con aspecto de
escamas, frente y clípeo con polinosidad grisácea, con brillo aterciopelado ........ 7
2(1) Especies pequeñas, longitud ala 1,8 mm; escudo sin manchas cuneiformes o
bandas longitudinales plateadas ........................................................... Simulium fuliginis
– Especies medianamente grandes; longitud del ala >2,0 mm; escudo con o sin
manchas plateadas, si la longitud de la ala <2,0 mm, muestran 1+1 bandas
medianas longitudinales .................................................................................................... 3
3(2) Escudo sin las manchas plateadas anteriores submedianas cuneiformes; pelos
agrupados como escamas aisladas dispuestas en una hilera mediana y dos
submedianas (Fig. 71 Q) ................................................................................................... 4
– Escudo con 1+1 manchas plateadas cuneiformes anteriores submedianas, a veces
prolongadas posteriormente y llegando al área grisácea posterior (Figs. 71 L, O);
pelos agrupados aparentando escamas a veces no bien aisladas y no bien
ordenadas en hileras .......................................................................................................... 6
4(3) Pelos del escudo dorado brillante ................... Simulium anamariae, Simulium stellatum
– Pelos del escudo con brillo verde violáceo o dorado violáceo ................................ 5
5(4) Hembra: frente de lados paralelos; triángulo frontoocular escasamente profundo
(Fig. 71 R). Macho: espolón del gonostilo muy próximo al ángulo interno (Fig. 72
H) ......................................................................................................... Simulium auristriatum
– Hembra: frente de lados convergentes (Fig. 71 S); triángulo frontoocular
relativamente profundo (Fig. 71 T). Macho: espolón del gonostilo próximo a la
distancia mediana del borde ........................................................ Simulium brevifurcatum
6(3) Escudo con 1+1 manchas plateadas cuneiformes no prolongadas posteiormente
(Fig. 71 L), si estuvieran prolongadas no sobrepasan el 1/4 anterior del escudo;
pelos del escudo con coloración dorado bronceado a dorado verdoso ...................
.................................................................... Simulium incrustatum, Simulium schmidtmummi
– Escudo con 1+1 manchas plateadas prolongadas posteriormente por una banda
grisácea polinosa que llega al área grisácea posterior (Fig. 71 O); pelos del escudo
de coloración gris verdoso pálido ......................................................Simulium limbatum
300 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

7(1) Escudo sin 1+1 bandas plateadas longitudinales submedianas; pilosidad muy
abundante con apariencia sedosa. Macho: gonostilo subtrapezoidal (Figs. 72 A, F)
................................................................................................................................................ 9
– Escudo con 1+1 bandas plateadas longitudinales submedianas anchas que llegan al
borde posterior grisáceo en hembras o pueden ser mas cortas y no sobrepasar la
mitad del largo del escudo en machos (Figs. 72 J, K); pilosidad escasa y sin
apariencia sedosa. Macho: con gonostilo subtriangular, con una carena longitudinal
(Figs. 72 L, M).
Hembra .............................. Simulium siolii, Simulium tergospinosum, Simulium damascenoi
Macho ................................................................................................................................... 8
8(7) Escudo con 1+1 manchas plateadas submedianas cuneiformes (Fig. 72 K) ..........
..................................................................................Simulium siolii, Simulium tergospinosum
– Escudo con 1+1 manchas plateadas submedianas subtrapezoidales (Fig. 72 J) ....
................................................................................................................ Simulium damascenoi
9(7) Especies grandes, longitud ala: hembra 2,5-2,8 mm; macho 2,1-2,6 mm; pilosidad
del escudo muy abundante, impidiendo ver las manchas plateadas cuneiformes
anteriores; pelos del escudo desde amarillo verdoso pálido a bronceado ...............
.................................................................................................................... Simulium angrense
– Especies medianas, longitud ala: hembra 1,7-2,5 mm; macho: 1,7-2,4 mm ........ 10
10(9) Uñas sin diente subbasal (Fig. 71 I) ................................................ Simulium minuanum
– Uñas con diente subbasal ............................................................................................... 11
11(10) Pilosidad del escudo predominantemente bronceado a amarillento gris-verdoso;
longitud ala: hembra 1,7-2,1 mm. Macho 1,6-2,0 mm .............. Simulium auripellitum
– Pilosidad del escudo predominantemente grisácea; longitud ala: hembra 2,1-2,5
mm; macho 1,9-2,4 mm ..................................... Simulium jujuyense, Simulium bonaerense

Pupas
1 Branquia con 2 ramas gruesas, subcilíndricas (Fig. 73 J) ......... Simulium auristriatum
– Branquia con 4-6 ramas finas, filamentosas ................................................................. 2
2(1) Branquias con 4 ramas (Figs. 73 K, 74 H) ................................................................... 3
– Branquias con 6 ramas (Figs. 74 B-G) ........................................................................... 8
3(2) Capullo con bien desarrollada prolongación dorsal anterior (Fig. 73 K) .............. 4
– Capullo sin prolongación dorsal anterior (Fig. 73 N) o muy corta (Figs. 73 L, M)
................................................................................................................................................ 5
4(3) Tricomas cefálicos y torácicos con 2 ramas .....................................Simulium stellatum
– Tricomas cefálicos con 1-4 ramas, torácicos con 4-12 tricomas .....................................
.......................................................................................................... Simulium schmidtmummi
5(3) Branquias más cortas que la longitud del capullo (Figs. 73 M, N); bifurcación de
ramas primarias próxima a la base (Fig. 74 I); frontoclípeo sin tubérculos .......... 6
– Branquia más larga que el largo del capullo (Fig. 73 L); bifurcación de ramas
primarias relativamente lejos de la base, frontoclípeo con tubérculos ................... 7
6(5) Capullo sin prolongación dorsal anterior y con muy corta prolongación ventral
(Fig. 73 M) ........................................................................................... Simulium damascenoi
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 301

– Capullo con corta prolongación dorsal anterior y relativamente larga la


prolongación ventral (Fig. 73) ..................................................... Simulium tergospinosum
7(5) Con áreas esclerotizadas sobre las membranas del abdomen ............ Simulium siolii
– Sin áreas esclerotizadas sobre las membranas del abdomen ........ Simulium lourencoi
8(2) Capullo con prolongación anterior dorsal (Figs. 73 H, J) ......................................... 9
– Capullo sin prolongación anterior dorsal (Figs. 73 A-G) ........................................ 10
9(8) Capullo y branquias expandidas lateralmente; prolongación anterior dorsal
relativamente corta (Fig. 73 I) ..................................................... Simulium brevifurcatum
– Capullo y branquias no expandidas lateralmente; prolongación anterior dorsal
muy alargada y bifurcada protegiendo las branquias lateralmente (Fig. 73 H) .......
................................................................................................................. Simulium anamariae
10(8) Capullo con tejido abierto, con las fibras bien evidentes; tricomas del frontoclípeo
y tórax con 3-4 ramas ......................................................................... Simulium guaporense
– Capullo con tejido compacto con fibras no muy evidentes; tricomas del
frontoclípeo y tórax con 2 ramas (Fig. 74 J) .............................................................. 11
11(10) Bifurcación de la rama primaria ventral y ramas secundarias de la branquia muy
próximas a la base (Fig. 74 C) .......................................................... Simulium bonaerense
– Bifurcación de la rama primaria ventral y ramas secundarias de la branquia no muy
próximas a la branquia (Figs. 74 B, D, E, G) ............................................................. 12
12(11) Pupa grande; longitud branquia 3,1-4,5 mm; frontoclípeo y tórax con abundante
(Fig. 73 P) a medianamente abundantes tubérculos (Fig. 73 Q) ............................ 13
– Pupa medianamente grande; longitud branquia 2,1-4,0 mm; frontoclípeo y tórax
con escasos tubérculos (Fig. 73 O) .............................................................................. 14
13(12) Frontoclípeo y tórax con abundantes tubérculos y tricomas moderadamente
largos (Fig. 73 P) .................................................................................... Simulium angrense
– Frontoclípeo y tórax con abundantes tubérculos y tricomas proporcionalmente
más largos (Fig. 73 Q) ...................................................................... Simulium incrustatum
14(12) Rama primaria ventral y secundarias de la branquia bifurcadas relativamente
próximas a la base (Fig. 74 F) .............................................................Simulium limbatum
– Rama primaria ventral y secundarias de las branquias bifurcadas aproximadamente
al doble de distancia de la base que en la opción opuesta (Figs. 74 D, E, G) ....... 15
15(14) Branquia vista de perfil aproximadamente recta (Fig. 73 C) ........ Simulium jujuyense
– Branquia vista de perfil con una suave curvatura mediana (Figs. 73 A-B) .......... 16
16(15) Bifurcación de ramas secundarias sobrepasando 1/3 de la longitud de la branquia
(Fig. 73 A) ...........................................................................................Simulium auripellitum
– Bifurcación de ramas secundarias sobrepasando 1/4 de la longitud de la branquia
(Fig. 73 B) ..............................................................................................Simulium minianum
302 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

INCRUSTATUM species group

Simulium (Psaroniocompsa) auripellitum ENDERLEIN (Figs. 71 A-C; 72 A; 73 A, O; 74 J.)


Simulium auripellitum ENDERLEIN, 1934b: 286; VULCANO, 1967: 7.
Simulium (Psaroniocompsa) auripellitum COSCARÓN & WYGODZINSKY, 1984: 54-58; COSCARÓN, 1987: 23;
1991: 180-181; 2000: 6; COSCARÓN & COSCARÓN ARIAS, 1997: 115; CROSSKEY & HOWARD, 1997:
57; 2004: 51; SHELLEY et al., 2000: 178-180.
Psaroniocompsa auripellita: PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 2004: 122-123.
Female: Wing length 2.1 mm. General coloration as described for subgenus. Frons
and clypeus grayish with nacreous sheen and reflecting bluish silvery to velvety
appearance. Scutum with homogeneous and abundant pilosity, bronze to yellowish gray-
green coloration with silky appearance, allowing 1+1 gray pollinose spots to be seen. Leg
coloration as in S. bonaerense. Abdomen blackish with 1+1 gray pollinose spots on tergite
II. Frons relatively wide (Fig. 71 A); fronto-ocular triangle reduced. Basal portion of
cibarium as in S. limbatum. Sensory vesicle of maxillary palp shorter than half length of
palpomere. Sc generally without hair. Claws with short subbasal tooth. Paraprocts
relatively short and curved distally (Fig. 71 B); genital fork with lateral branches scarcely
sclerotized (Fig. 71 C); eighth sternite with 12-15 setae and gonapophyses with internal
border subparallel and slightly sclerotized.
Male: Wing length 1.6-2.0 mm. Area around antenna insertion silvery. Scutum
velvety black with hair as in female, male basitarsus ratio 6.1-7.5. Abdomen as described
for subgenus. Genitalia as in S. bonaerense. Gonostylus subquadrate with strong spur
positioned near midsection of distal border (Fig. 72 A).
Pupa: Cocoon with small dorsal projection, and reinforced anterior border (Fig. 73
A); threads evident. Cocoon base length 2.5-3.0 mm; on dorsum 2.0-2.7 mm; gills: 1.9-2.7
mm. Gill filaments with 6 terminal relatively joined branches, with short ventral curvature
at half total length; secondary bifurcations at relatively long distance from base;
bifurcation of dorsal secondaries about 1/3 or more of total gill length (Fig. 73 A).
Platelets relatively scarce (Fig. 73 O) and trichomes of frontoclypeus and exposed portion
of thorax bifid (Fig. 74 J).
Larvae: Color, size and morphology very close to S. incrustatum. Ratio of proximal,
medial and distal antennal segments = 1:1.0-1.1:1.0-1.2. Anal sclerite with scarce hairs. Anal
ring with 62-65 rows of 11-13 hooks each. Rectal papillae with 6-11 diverticula on each lobe.
Distribution: Argentina: Corrientes, Misiones, Santa Fe; Brazil: Goias, Mato
Grosso, Mato Grosso do Sul, Minas Gerais, Paraná, Rio Grande do Sul, Santa Catarina,
São Paulo, Tocantins; Paraguay: Alto Paraná, Asunción, La Cordillera; Uruguay.
Bionomics: This species is very abundant in small creeks with moderate flow and
clean temperate water. Larval stages live on submerged aquatic vegetation or on grasses at
the border of the streams. Females will bite humans and cattle.
Discussion: This species is difficult to separate from the other species with
generalized characters. Specimens of S. jujuyense in austral and western populations are of
smaller size, with gill branches bifurcating nearer to the base and without a ventral
concavity, but these differences are not evident in the eastern populations, living in the
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 303

L O
Q
U R

P K
S
A

M
G
F

D E

J
N
C H B
I
Figure 71. Female. A-C: S. (Psaroniocompsa) auripellitum; A: frons; B: cercus and paraproct; C: genital fork.
D-H: S. (P.) bonaerense; D-F: fore, mid and hind leg, respectively; G: genitalia in ventral view showing
eighth sternite, gonapophyses, genital fork and spermatheca; H: cercus and paraproct. I-J: S. (P.)
minuanum; I: claw; J: cercus and paraproct. K-N: S. (P.) incrustatum; K: grouped hair of scutum; L: scutum;
M: anterior veins of wing; N: claw. O-P: S. (P.) limbatum; O: scutum; P: basal portion of cibarium. Q-R:
S. (P.) auristriatum; Q: scutum; R: fronto-ocular triangle. S-T: S. (P.) brevifurcatum; S: frons; T: fronto-ocular
triangle. U: S. (P.) damascenoi, head, dorsal view and scutum (from HAMADA et al., 2003).
304 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

C
E

A
D

H I M

Figure 72. Male. A: S. (Psaroniocompsa) auripellitum, gonocoxite and gonostylus. B-D: S. (P.) bonaerense;
B: hairs of legs; C: distal portion of hind basitarsus with calcipala and portion of second tarsite; D:
genitalia in ventral view, showing gonocoxite, gonostylus, ventral plate and endoparameres. E-F: S. (P.)
limbatum; E: scutum; F: gonostylus. G: S. (P.) anamariae, ventral plate. H: S. (P.) auristriatum, gonocoxite
and gonostylus. I: S. (P.) schmidtmummi, gonocoxite and gonostylus (from WYGODZINSKI, 1973). J: S. (P.)
damascenoi, scutum and portion of head (from HAMADA et al., 2003). K-M: S. (P.) tergospinosum (from
HAMADA, 2000); K: scutum; L: gonocoxite and gonostylus; M: gonostylus dorsal view showing
longitudinal carina.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 305

same areas with S. auripellitum. Simulium minuanum is differentiated by having scutum hairs
of silvery with greenish to bronze tones, and claws without a subbasal tooth. Simulium
angrense shows hairs on the Sc, and the more abundant scutum hairs do not permit the
anterior 1+1 subtriangular silvery spots to be seen. The pupal gill filaments are more
expanded and the frontoclypeal and thoracic tubercles are more abundant. Simulium
bonaerense has scutum hairs more grayish in coloration and the pupal gill bifurcation is
closer to the base. Simulium incrustatum is similar, but the grouped scutum hairs, more
abundant platelets on the pupal frontoclypeus and thorax, and gill bifurcation of the
secondary branches at a greater distance from the base, allow it to be separated.

Simulium (Psaroniocompsa) minuanum STRIEDER & COSCARÓN (Figs. 71 I, J.)


Simulium (Psaroniocompsa) minuanum STRIEDER & COSCARÓN, 2000: 105-111; COSCARÓN, 2003: 5;
CROSSKEY & HOWARD, 2004: 51.
Psaroniocompsa minuana: STRIEDER, 2004: 124.
Female: Very close to some others species of subgenus; it is characterized by wing
length 2.0-2.2 mm, scutum hair homogeneously distributed; frons relatively narrow and
frontal angle lower than in closest species; sensory vesicle of maxillary palp surpassing
half length of palpomere; claws without subbasal tooth (Fig. 71 I); paraproct longer than
high and with rounded border (Fig. 71 J); gonapophysis with internal border slightly
sclerotized and distal border with nearly straight margin.
Male: Wing length 1.8-2.0 mm, no special differences from other species of the group.
Pupa: Cocoon without anterior projection. Primary ventral branch elongated, with
bifurcation surpassing 3 times primary dorsal branch bifurcation length (Fig. 73 B).
Frontoclypeus and exposed portion of thorax with scarce platelets, fewer than in S.
auripellitum; trichomes of thorax with 1-3 branches.
Larva: Maximum length 3.4-3.6 mm. Body integument with abundant short hairs.
Cephalic apotome slightly positive. Antenna surpassing apex of cephalic fan stem; ratio of
antennal segments: proximal, medial and distal = 1:0.6-0.7:1.1-1.2. Hypostomium with 4
setae on each side. Mandible with one latero-mandibular process. Postgenal cleft deep
with rounded margin. Anal ring with 60-66 rows with 8-13 hooks each. Rectal papillae
with 8-10 diverticula on each lobe.
Distribution: Argentina: Corrientes, Entre Ríos, Misiones; Brazil: Rio Grande do Sul.
Bionomics: Larval stages live in creeks of fast, clear water, with soft milky tones,
flowing over aquatic vegetation or submerged tree branches. Female feeding habits are
unknown.
Discussion: As it is difficult to differentiate S. minuanum from other closely related
species, we have added a table with differentiating characteristics:

Simulium Simulium Simulium Simulium Simulium


angrense auripellitum minuanum jujuyense bonaerense
‡ wing length (mm) 2.5-3.0 2.1-3.0 2.0-2.2 2.1 -2.4 2.2-2.5
x̄ = 2.6 x̄ = 2.3 x̄ = 2.1
n=3 n=6
306 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Simulium Simulium Simulium Simulium Simulium


angrense auripellitum minuanum jujuyense bonaerense

† wing length (mm) 2.1-2.6 1.8-2.0 1.8-2.0 1.9-2.2 2.0-2.4


x̄ = 2.3 x̄ = 1.9 x̄ = 1.9
n=3
Scutum hair Tin to Tin to Silvery with Silvery- Tin to
coloration yellowish golden; male bright green grayish
green silvery-gray green to tin to tin
Frontal angle 103° 110-120° 100° 115-120° 110°
Ratio of length of 2.9-3.0 2.2-2.5 2.3-2.5 2.0-2.5 2.5-2.8
third palpomere to x̄ = 2.9 x̄ = 2.3 x̄ = 2.4 x̄ = 2.2 x̄ = 2.6
sensory vesicle n=2 n=4 n=3
Ratio of length/width of 6.2-8.9 7.0-9.0 7.3-8.2 6.4-7.7 6.0-7.0
hind basitarsus, female x̄ = 7.4 x̄ = 7.8 x̄ = 7.7 x̄ = 7.1 x̄ = 7.1
n=3 n=5
Ratio of length/width of 6.4-6.5 6.1-7.5 7.5-7.9 7.7-8.0 6.0-7.3
hind basitarsus, male = 6.4 = 6.6 = 7.7 = 7.9 = 6.5
n=4
Claw with subbasal tooth + + - + reduced + reduced
Pupa with tubercles on Very Relatively More Medium to From scarce
frontoclypeus and thorax numerous scarce scarce scarce to numerous
Cocoon ventral 3.0-3.7 2.5-3.0 2.5-2.9 2.8-3.0 3.8-4.0
length (mm)
Cocoon dorsal 2.7-3.2 2.0-2.7 1.7-2.0 2.5-2.7 3.2-3.5
length (mm)
Gill length 3.1-4.5 1.9-2.7 1.7-2.3 3.4-3.5 2.5-3.0
Larva antennal ratio of 1:1.0:1.0 1:1.0-1.1: 1:0.66-0.71: 1:1.0.0.9 1:1.0:0.9-1.0
segments: proximal, 1.0-1.2 1.15-1.27
medial and distal

Simulium (Psaroniocompsa) jujuyense PATERSON & SHANNON (Figs. 73 C; 74 A, B.)


Simulium jujuyense PATERSON & SHANNON, 1927: 739; WYGODZINSKY, 1951a: 206-214; COSCARÓN &
WYGODZINSKY, 1960: 1124-1125; VULCANO, 1967: 14.
Simulium (Psaroniocompsa) jujuyense: COSCARÓN & WYGODZINSKY, 1984: 67-69; COSCARÓN, 1987: 23;
COSCARÓN & COSCARÓN ARIAS, 1997: 115; CROSSKEY & HOWARD, 1997: 57; 2004: 51; STRIEDER
& COSCARÓN, 2000: 111; COSCARÓN, 2003: 5; COSCARÓN ARIAS, 2003: 198, 204, 208, 215.
Psaroniocompsa mendozana ENDERLEIN, 1936: 123.
Simulium spinifer KNAB, 1914b: 81.
Simulium hoffmanni VARGAS, 1943a: 138 (n. nom. Trichodagmia angustitarse ENDERLEIN, 1934b: 289).
Simulium sicuani SMART, 1944: 132 (n. nom. Ectemnaspis limbata ENDERLEIN, 1934b: 282).
Simulium incaicum VARGAS, 1945a: 148 (unjustified repl. name for E. limbata).
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 307

Female: Wing length 2.1-2.4 mm. Coloration general blackish. Frons and clypeus
velvety grayish. Scutum with silvery to bronze, silky pilosity, homogeneously distributed.
Legs dark brown to black on darkened areas. Abdomen with 1+1 lateral silvery spots on
tergite II. Wings without hair on Sc. Genitalia as in S. bonaerense.
Male: Wing length 1.9-2.2 mm; general coloration as in female. Genitalia as in S.
bonaerense.
Pupa: Cocoon yellowish whitish-gray, with straight border and without anterior
reinforcement (Fig. 73 C); weave superficially smooth, with thin threads. Length at base
2.8-3.0 mm; on dorsum 2.5 mm, gill 3.4 mm. Gill bifurcation of secondary branches more
proximal than in S. auripellitum, but more distal than in S. bonaerense. Primary ventral
branch bifurcation more proximal than secondary ventral (Fig. 73 C) to about same level
(Fig. 74 B). Frontoclypeal and thoracic tubercles relatively scarce, fewer than in S.
incrustatum. Abdominal chaetotaxy as in figure 73 A.
Larva: Maximum length 4.5-5.0 mm. General coloration light yellowish gray, with
greenish violaceous on abdominal segment bands. Ratio of antennal segments: proximal,
medial and distal = 1:1.0:0.9. Mandible, hypostomium and postgenal cleft as in S.
incrustatum. Anal gill with 8-10 diverticula on each lobe.
Distribution: Argentina: Buenos Aires, Chaco, Córdoba, Corrientes, Entre Ríos,
Formosa, Jujuy, La Pampa, La Rioja, Mendoza, Río Negro, Salta, Tucumán; Bolivia;
Colombia; Ecuador: Cotopaxi, Esmeraldas; Peru: Calca, Cuzco.
Bionomics: The preimaginal stages live on aquatic vegetation and submerged trees
branches, especially Salix humboldtiana in moderate to fast-flowing creeks and rivers.
Females attack humans, cows and horses.
Discussion: The closest species are S. auripellitum and S. bonaerense, which are
sympatric in some places, and can be differentiated by the yellowish green scutum pilosity
and the pupal gill branching, as explained in the description. Simulium bonaerense can be
distinguished by the big size and especially by the lower gill branching. This similarity
produced misidentification by COSCARÓN & WYGODZINSKY (1960: 1124) and COSCARÓN
(1968: 66) and in part by COSCARÓN & WYGODZINSKY (1984: 57) in relation to material
from Chaco, Corrientes, Entre Ríos and Formosa, which was corrected by STRIEDER &
COSCARÓN (2000: 111).

Simulium (Psaroniocompsa) bonaerense COSCARÓN & WYGODZINSKY


(Figs. 71 D-H; 72 B-D; 73 D; 74 C; 75 A, B.)
Simulium (Psaroniocompsa) bonaerense COSCARÓN & WYGODZINSKY, 1984: 59-62; COSCARÓN, 1987: 23;
1991: 183-184; COSCARÓN & COSCARÓN ARIAS, 1997: 115; CROSSKEY & HOWARD, 1997: 57;
2004: 51.
Simulium jujuyense PATERSON & SHANNON, 1927; COSCARÓN & WYGODZINSKY, 1960: 1124; COSCARÓN,
1968: 66 (misidentification).
Female: Wing length 2.2-2.5 mm. General coloration blackish. Frons and clypeus
gray pollinose, frons with velvety aspect. Scutum with hair bronze to grayish color,
homogeneously arranged, permitting subtriangular anterior silver spots to be seen. Leg
coloration in accordance with figures 71 D-F. Claw subbasal tooth present but small.
308 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

B
I
C

F
G

H
P

L
N O
Q

Figure 73. Pupa. A-N: General view; A: S. (Psaroniocompsa) auripellitum; B: S. (P.) minuanum; C: S. (P.)
jujuyense; D: S. (P.) bonaerense; E: S. (P.) angrense; F: S. (P.) incrustatum; G: S. (P.) limbatum; H: S. (P.)
anamariae; I: S. (P.) brevifurcatum; J: S. (P.) auristriatum; K: S. (P.) schmidtmummi (from WYGODZINSKY,
1973); L: S. (P.) siolii; M: S. (P.) damascenoi (L-M from PY-DANIEL, 1988); N: S. (P.) tergospinosum (from
HAMDA, 2000). O-Q Frontoclypeus; O: S. (P.) auripellitum; P: S. (P.) angrense; Q: S. (P.) incrustatum.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 309

I
C
B

H
G
K

A
D
E
F

Figure 74. Pupa. A: S. (Psaroniocompsa) jujuyense, abdominal chaetotaxy. B-I: Gills; B: S. (P.) jujuyense;
C: S. (P.) bonaerense; D: S. (P.) angrense; E: S. (P.) incrustatum; F: S. (P.) limbatum; G: S. (P.) anamariae; H:
S. (P.) schmidtmummi (H from WYGODZINSKY, 1973); I: S. (P.) damascenoi (I from PY-DANIEL, 1988). J-
K: Trichomes; J: S. (P.) auripellitum, trichomes and tubercles of thorax; K: S. (P.) schmidtmummi,
multibranched trichomes (from WYGOODZINSKY, 1973).
310 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Abdomen with 1+1 small silvery spots on tergite II. Frons relatively wide but narrower
than in S. auripellitum; fronto-ocular triangle very reduced. Wings with hair on Sc. Genitalia
as in figure 71 G. Eighth sternite with about 40 setae and gonapophyses with abundant
microtrichiae, internal border subparallel and scarcely sclerotized. Paraproct short (Fig. 71
H) and genital fork with stem well sclerotized.
Male: Wing 2.0-2.4 mm. General coloration as in female and with same pattern as in
other males of subgenus. Legs as in female with filiform and spatuliform hair (Fig. 72 B).
Genitalia as in figure 72 D.
Pupa: Cocoon light brown, straight and slowly reinforced on anterior border (Fig. 73
D); weave superficially smooth with evident threads; length at base 3.8-4.0 mm, at dorsum
3.2-3.5 mm, gills 2.5-3.0 mm. Gill branching very close to base (Fig. 74 C) and relatively
open. Frontoclypeus and exposed portion of thorax with platelets medially, abundant as
in S. incrustatum. Abdominal chaetotaxy as in S. jujuyense.
Larva: General coloration from greenish to yellowish brown. Morphology as in S.
incrustatum, only differentiated by scarce gill diverticula that only reach 4 on each lobe in S.
bonaerense (Fig. 75 B). Antenna as in figure 75 A, with ratio of proximal, medial and distal
segments = 1:1:0.9-1.0. Rectal papillae with 76-80 rows with 15-16 hooks each.
Distribution: Argentina: Buenos Aires (center and south west area), Río Negro
(Alto Valle).
Bionomics: This species lives in environments similar to those of S. jujuyense. It is
present all year, but the highest peak is in the spring (COSCARÓN et al., 2000). Females bite
humans and cattle, and are a problem for inhabitants.
Discussion: The closest species are S. auripellitum, S. jujuyense and S. minuanum. The
larger size and pupal gill branching close to the base allow it to be differentiated from
them. Also, S. auripellitum and S. minuanum show differences in scutum hair coloration,
which is bronze to golden with silvery greenish tones, and the latter species does not have
a subbasal tooth on the claws.

Simulium (Psaroniocompsa) angrense PINTO (Figs. 73 E, P; 74 D.)


Simulium angrensis PINTO, 1931: 683.
Simulium angrense: VARGAS, 1945a: 112; VULCANO, 1967: 6.
Simulium (Psaroniocompsa) angrense COSCARÓN & WYGODZINSKY, 1984: 52-54; COSCARÓN, 1987: 23;
1991: 179; COSCARÓN & COSCARÓN ARIAS, 1997: 115; CROSSKEY & HOWARD, 1997: 57; 2004: 51.
Psaroniocompsa angrense: PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 2004: 122.
The type of this species was not found at the Instituto Oswaldo Cruz by the first
author in 1979 and the description of COSCARÓN & WYGODZINSKY (1984) that is used as a
guide here, is based on material collected close to the type locality (Angra dos Reis, Rio de
Janeiro), and in accordance with the original description.
Female: Wing length 2.5-2.8 mm. Frons and clypeus from velvety grayish pollinose
to bronze to greenish yellow; scutum hair tin to yellowish green, arranged homogeneously,
difficult to see the 1+1 anterior subtriangular silvery spots. Sc with 10-12 hairs. Claw with
small subbasal tooth. Genitalia as in S. bonaerense.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 311

Male: Wing length 2.1-2.6 mm. General coloration as in female. Area around
antenna insertion velvety grayish. Scutum black, velvety with hair arranged as in female. Sc
without hairs. Genitalia similar to those of S. bonaerense.
Pupa: Cocoon with small anterior projection and reinforced on anterior border (Fig.
73 E), with resistant weave and visible threads. Length on base 3.0-3.7 mm, on dorsum
2.7-3.2 mm, gills 3.1-4.5 mm. Gill branching near base (Fig. 73 E), secondary branches
bifurcating frequently at same level (Fig. 74 D); frontoclypeus with abundant tubercles
(Fig. 73 P); trichomes bifid and a little shorter than in S. incrustatum.
Larva: Maximum length 4.5-5.5 mm. General coloration yellowish green. General
morphology close to that of S. incrustatum but mandible with thinner marginal teeth and
rectal papillae diverticula numbering 10-12 on each lobe.
Distribution: Brazil: Rio de Janeiro, Rio Grande do Sul, Santa Catarina, São Paulo.
Bionomics: Unknown.
Discussion: The closest species is S. auripellitum, which can be differentiated by its
smaller size, scutum pilosity more silky yellowish and pupal gill with secondary branches
bifurcating at a greater distance from the base. Simulium incrustatum, which is present in the
same area, can be differentiated by the scutum hair arranged in groups simulating scales,
and the pupal primary ventral and secondary branches bifurcating at greater distance from
the base.

Simulium (Psaroniocompsa) incrustatum LUTZ (Figs. 71 K-N; 73 F; 74 E, Q; 75 C-J.)

Simulium incrustatum LUTZ, 1910: 243-245; PINTO, 1931: 703; LANE & PORTO, 1940: 194; VARGAS,
1945a: 148; VULCANO, 1967: 13; COSCARÓN & WYGODZINSKY, 1973a: 170.
Simulium (Psaroniocompsa) incrustatum: COSCARÓN & WYGODZINSKY, 1984: 62-67; COSCARÓN, 1987: 23;
1991: 184-186; 2003: 5; COSCARÓN & COSCARÓN ARIAS, 1997: 115; CROSSKEY & HOWARD, 1997:
57; 2004: 51; SHELLEY et al., 1997: 24; COSCARÓN ARIAS, 2003: 198, 204, 208-215.
Psaroniocompsa incrustata: PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 2004: 133-135;
2004: 123-124.
Psaroniocompsa opalinifrons ENDERLEIN, 1934a: 192; 1936: 112.
Simulium opalinifrons: VARGAS, 1945a: 100; WYGODZINSKY, 1951a: 217; VARGAS & DIAZ NÁJERA,
1953b: 137.
Simulium (Psaroniocompsa) opalinifrons: COSCARÓN & WYGODZINSKY, 1973a: 161-172.
Simulium yarzabali RAMÍREZ PÉREZ, 1980: 66-68; 1983: 9; SHELLEY et al., 1987a: 463 (revalidated).
Female: Wing length 2.0-2.8 mm. General coloration blackish. Frons and clypeus
with silvery pruinosity, bluish-green sheen iridescent. Scutum velvety black with golden to
bronze hair grouped, simulating scales (Fig. 71 K) and arranged in longitudinal rows
specially at sides of anterior half (Fig. 71 L). Scutellum and metanotum black, the latter
with gray pollinosity. Legs brown with darkened areas blackish showing, with some
illumination, whitish fore tibia internally and 3/4 hind basitarsus externally. Abdomen
blackish with 1+1 silvery pollinose spots on tergite II. Fronto-ocular triangle very short;
Sc with 4-12 hairs, chaetotaxy of anterior wing veins as in figure 71 M. Claws with short
basal tooth (Fig. 71 N). Genitalia as in S. bonaerense; genital fork with well-sclerotized stem.
312 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

C
D

L
O

F
A

H E

G
N

J I B

Figure 75. Larva. A-B: S. (Psaroniocompsa) bonaerense; A: antenna; B: rectal papillae. C-J: S. (P.)
incrustatum; C: general aspect; D: head, dorsal view; E: antenna; F: mandible; G: trichomes of head,
dorsum; H: postgenal cleft and hypostomium; I: anal sclerite; J: distal portion of abdomen with
rectal papillae. K-N: S. (P.) auristriatum; K: general aspect; L: head, dorsal view; M: antenna; N:
hypostomium. O-Q: S. (P.) siolii ( from PY-DANIEL, 1988); O: general aspect; P: abdomen with more
magnification showing dorsal prominences and trichomes; Q: teeth of mandible with latero-
mandibular process (O-Q from PY-DANIEL, 1988).
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 313

Male: Wing length 1.9-2.8 mm. General coloration as in female. Area around
antennae insertion silvery with iridescent sheen. Scutum velvety black with golden hair
grouped as in female, but not arranged in longitudinal rows. Sc with 0-2 hairs. Abdominal
pattern as in subgenus. Genitalia as in S. bonaerense.
Pupa: Cocoon without dorsal anterior projection, slowly reinforced on rim, weave
not compacted, threads visible (Fig. 73 F). Length at dorsum 2.5-2.9 mm; at base 3.0-3.2
mm; gills 3.1-4.5 mm. Generally gill bifurcation of primary ventral and secondary
branches relatively far from base, variable distance among specimens, with secondary
branches bifurcating about 1/2-1/3 of total gill length, and primary ventral bifurcation at
half of anterior distance (Fig. 74 E). Cephalothoracic trichomes mostly bifid and relatively
elongated; platelets of frontoclypeus and exposed portion of thorax moderately abun-
dant, some grouped, leaving clear areas especially on thorax (Fig. 74 Q).
Larva: Maximum length 4.6-5.0 mm. General coloration light greenish-yellow.
General aspect as in figure 75 C. Body integument with microscopic hairs, like flattened
multibranched scales as seen with high magnification. Cephalic apotome with small,
discontinuous, median and sublateral positive spots (Fig. 75 D); Antennae surpassing stem
length of cephalic fan; ratio of proximal, medial and distal segments = 1:1.0-1.1:1.2 (Fig.
75 E). Mandible with thick first marginal tooth (Fig. 75 F). Head tegument with bifid hairs
(Fig. 75 G) Cephalic fan with about 35-40 rays. Hypostomium with 4 setae on each side;
postgenal cleft deep and rather rounded (Fig. 75 H). Anal ring with about 62-70 rows with
12-15 hooks each; anal sclerite with hairs single or with several branches (Fig. 75 I). Rectal
papillae with 5-8 diverticula on each lobe (Fig. 75 J).
Distribution: Argentina: Corrientes, Misiones; Brazil: Bahia, Federal District, Mato
Grosso do Sul, Minas Gerais, Paraná, Pernambuco, Rio de Janeiro, Rio Grande do Sul,
Roraima, Santa Catarina, São Paulo; Ecuador: Cotopaxi, El Oro, Esmeraldas, Guayas,
Los Rios, Manabi; Paraguay: Alto Paraná, La Cordillera; Venezuela: Amazonas, Bolivar.
Bionomics: This species breeds in small creeks with clean water, on submerged
aquatic vegetation. Females are haematophagous and the bite produces serious annoy-
ance in Brazil.
Discussion: The presence of hair grouped and forming scales on the scutum can
confuse the identification with S. limbatum, S. anamariae, S. auristriatum, S. brevifurcatum,
S. cuasiexiguum, S. exiguum and S. paraguayense. The three last-mentioned species
correspond to S. (Notolepria), with different genitalia and pupal gill and trichomes. From
limbatum, it can be differentiated by the presence of 1+1 silvery submedian stripes going
from anterior subtrianglular spots to the posterior border of the scutum, and grayish
pilosity. From S. anamariae, S. auristriatum and S. brevifurcatum it differs by the presence
of larger hair groups and greenish violaceous coloration, plus the cocoon with a dorsal
projection and different gill morphology. Pliodasina guttata (ENDERLEIN) from Paraguay
has some similarity with the female type of S. incrustatum, but since it is not well
preserved, it is not possible to define its identity; no other stages are known and we
prefer to maintain it as species inquirendae. Simulium yarzabali was synonymized by
RAMÍREZ PÉREZ (1983: 9), and rejected by SHELLEY (1987: 464). We consider it a
geographic variant and accept the RAMÍREZ PÉREZ criteria.
314 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Simulium (Psaroniocompsa) limbatum KNAB (Figs. 71 O, P; 72 E, F; 73 G; 74 F.)


Simulium limbatum KNAB, 1915c: 259; RAMÍREZ PÉREZ et al., 1982: 45; RAMÍREZ PÉREZ, 1983: 43;
SHELLEY et al., 1984: 143; SHELLEY et al., 1997: 27.
Simulium (Psaroniocompsa) limbatum: COSCARÓN, 1991: 188; CROSSKEY & HOWARD, 1997: 57; 2004 : 51.
Simulium (Psaroniocompsa) aequifurcatum LUTZ, 1910: 259, 263; COSCARÓN & WYGODZINSKY, 1984: 49;
COSCARÓN, 1987: 23; 1991: 186.
Simulium machadoi RAMÍREZ PÉREZ, 1971: 341.
Simulium machadoallisoni VULCANO, 1981: 276 (n. nom. to Simulium machadoi RAMÍREZ PÉREZ).
Simulium meruoca NUNES DE MELLO et al., 1973: 59.
Female: Wing length 1.8-2.2 mm. Color generally blackish. Frons and clypeus with
silvery pruinosity and blue sheen. Scutum black with one median and 1+1 submedian
silvery pollinose thin bands reaching posterior grayish area (Fig. 71 O), (evident with some
special illumination). Scutum hair pale gray-greenish, arranged in groups but fewer in
number than in S. incrustatum. Sc without hair, cibarium with evident spines arranged in
several rows (Fig. 71 P). Eighth sternite with about 20-25 setae on each side, and
gonapophysis with internal rim slightly divergent.
Male: Wing length 1.8-2.0 mm. General coloration as in female but gray pollinosity
of scutum bands shorter and thinner posteriorly (Fig. 72 E). Genitalia as in S. bonaerense
but gonostylus with wrinkled border (Fig. 72 F).
Pupa: Cocoon with slight dorsal anterior projection and reinforced border (Fig. 73 G).
Length on base 2.6-3.0 mm; on dorsum 2.4-2.6 mm. Gill with 6 branches emerging close to
base; primary ventral branch bifurcated about half of secondary dorsal and ventral
bifurcation level, (as shown in figure 74 F), or secondary dorsal branch bifurcating about two
times distance of secondary ventral bifurcation. Frontoclypeus and exposed portion of
thorax with numerous platelets, similar to condition in S. incrustatum, but with variation in
number and not arranged in groups. Trichomes bifid and shorter than in S. incrustatum.
Larva: Maximum length 4.0 mm. Body integument with scarce and short filiform,
generally single hairs. Cephalic apotome similar to that of S. incrustatum. Postgenal bridge
shorter than half of hypostomium length. Ratio of antennal segments: proximal, median
and distal = 1:0.8-0.9:1.1. Mandible with 2 latero-mandibular processes. Lateral sclerite of
thoracic proleg with 14-19 teeth arranged in 4-6 groups. Anal ring with 65-70 rows with
10-12 hooks each. Rectal papillae with 3 diverticula on each lobe.
Distribution: Brazil: Amapá, Bahia, Espírito Santo, Mato Grosso, Paraiba, Pará,
Pernambuco, Rio de Janeiro, Roraima, São Paulo; Guyana: Rupununi; Venezuela:
Amazonas, Anzoategui, Aragua, Barinas, Bolivar, Carabobo, Cojedes, Guarico, Lara,
Miranda, Monagas, Portuguesa, Táchira, Trujillo, Yaracuy.
Bionomics: This species breeds in small creeks, about 1-5 m wide and not more 1 m
deep, not torrential, flowing over aquatic vegetation. Females were collected biting
humans and horses.
Discussion: Simulium limbatum is very close to S. incrustatum. They can be differenti-
ated by the presence in S. incrustatum of 1+1 short silvery bands on the scutum that do not
continue from the anterior cuneiform spots nor surpass 1/4 of the scutum length, and
golden scutum hair arranged in groups a little bigger than in S. limbatum. Also, the pupal
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 315

gill branches more distal from the base. The latter character shows great variation in both
species and can produce confusion, especially when the material is not reared. Gill
branching similar to that in S. limbatum can be found in the range of variation for S.
incrustatum (SHELLEY et al., 1997). COSCARÓN & WYGODZINSKY (1984) and COSCARÓN (1987,
1991) used S. aequifurcatum LUTZ to describe this species, based on material deposited at
Institute Oswaldo Cruz. It corresponds to a pupa, without exact provenance. No reared
material from the LUTZ collection was found. The best characters to separate S. limbatum
from S. incrustatum are in the adults. It is best to use S. limbatum to name this taxon, which
was described from a female, and to accept the synonymy of S. aequifurcatum with S.
incrustatum proposed by SHELLEY et al. (1984). Material indicated as aequifurcatum by
COSCARÓN & WYGODZINSKY (1984), COSCARÓN (1987, 1991) & COSCARÓN et al. (1992)
corresponds to S. limbatum.

Simulium (Psaroniocompsa) fuliginis FIELD


Simulium fuliginis FIELD, 1969b: 281.
Simulium (Psaroniocompsa) fuliginis: COSCARÓN & WYGODZINSKY, 1984: 77; COSCARÓN, 1987: 24; 1991:
188; COSCARÓN & COSCARÓN ARIAS, 1997: 115; CROSSKEY & HOWARD, 1997: 57; 2004: 52.
This species was described based on females from Panama. The description, with
good illustrations, is similar to that for S. limbatum KNAB. It is differentiated only by its
small size (wing length 1.8 mm) and absence of 1+1 submedian thin silvery stripes on the
scutum, evident with some illumination. We could not see the types which are not in the
U.S.N.M. as indicated by the author. With the identity in doubt, we prefer not to
synonymize this species with S. limbatum, pending study of type material of all stages.

Simulium (Psaroniocompsa) anamariae VULCANO (Figs. 72 G; 73 H; 74 G.)


Simulium anamariae VULCANO, 1962: 246; 1967: 6.
Simulium (Psaroniocompsa) anamariae: COSCARÓN & WYGODZINSKY, 1984: 69-71; PY-DANIEL et al., 1985:
97-98; COSCARÓN, 1987: 23; 1991: 189-190; COSCARÓN & COSCARÓN ARIAS, 1997: 115;
CROSSKEY & HOWARD, 1997: 57; 2004: 52.
Psaroniocompsa anamariae: PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 2004: 122.
Female: Wing length 2.5-2.8 mm. Frons and clypeus grayish with bluish silvery
iridescent sheen. Scutum black without anterior spots, with abundant hair groups like
scales arranged in rows, one median and several laterals as in S. incrustatum. Humeral callus
light brown. Scutellum, metanotum and pleurae black; abdomen with tergite I brown with
nacreous sheen pollinosity at sides; tergite II with 1+1 lateral gray pollinose spots;
following tergites brownish with cereous sheen. Fronto-ocular triangle reduced, wider
than long. Legs with spatuliform hairs; claws with small subbasal tooth. Eighth sternite,
paraproct, cercus and gonapophysis as in S. bonaerense. Genital fork with enlarged stem
similar to that of S. auripellitum.
Male: Wing length 2.0-2.5 mm. General coloration blackish, ornamented as typical
for subgenus. Gonocoxite subtrapezoidal, about as long as wide; gonostylus half length of
gonocoxite, subtrapezoidal with spur subapical. Ventral plate as in figure 72 G.
316 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Pupa: Cocoon relatively flattened with two anterior, dorsal enlarged projections (Fig.
73 H); weave not compressed, with evident threads. Length on base 3.2-3.6 mm; on dorsum
without projections, 2.8-3.2 mm; gills 3.8-4.5 mm. Primary ventral and secondary dorsal and
median branch bifurcated far from base; the two latter about at half of total length (Fig. 74
G). Frontoclypeus with 2 pairs of frontal trichomes elongated and with 2-3 branches and 1
pair facials simple. Tubercles of cephalothorax abundant, but relatively small.
Larva (based on PY-DANIEL et al., 1985): Maximum length 5.1-5.3 mm. Body
integument with spiculae of 3-4 branches. Ratio of antennal segments: proximal, medial
and distal = 1:0.6:1.2. Cephalic fan with 45-46 rays. Mandibles with 2 sinuous latero-
mandiblular processes surpassing teeth apical border. Anal ring with about 66 rows with 9-
12 hooks each. Rectal papillae with 8-10 diverticula on each lobe.
Distribution: Brazil: Rio de Janeiro, Rio Grande do Sul, São Paulo.
Bionomics: Unknown.
Discussion: The most similar species are S. brevifurcatum and S. auristriatum, which
have similar scutum pilosity with grouped hairs arranged in longitudinal rows, and without
1+1 anterior silvery cuneiform spots. It is easy to separate them by the peculiar two
projections of the S. anamariae cocoon. Also, the gills have an open arrangement with very
basal branching in S. brevifurcatum, and the gill has two thick branches in S. auristriatum.

Simulium (Psaroniocompsa) auristriatum LUTZ (Figs. 71 Q, R; 72 H; 73 J; 75 K-N.)


Simulium auristriatum LUTZ, 1910: 245-246, 266-267; LUTZ & NUNEZ TOVAR, 1928: 131; PINTO, 1931:
684; D’ANDRETTA & D’ANDRETTA, 1949: 55-56; VULCANO, 1967: 7.
Simulium (Psaroniocompsa) auristriatum: COSCARÓN & WYGODZINSKY, 1984: 71-74; COSCARÓN, 1987: 23;
1991: 190-191; COSCARÓN & COSCARÓN ARIAS, 1997: 115; CROSSKEY & HOWARD, 1997: 56; 2004: 50.
Simulium infuscatum LUTZ, 1910: 236; PINTO, 1931: 704; PORTO, 1939: 369.
Psaroniocompsa auristriata: PY-DANIEL & MOREIRA SAMPAIO, 1995: 119.
Female: Wing length 2.5-2.6 mm. General coloration blackish. Scutum with hair
grouped as big scales greenish blue to violet and arranged in longitudinal rows (Fig. 71 Q).
Scutellum wide and fronto-ocular triangle reduced (Fig. 71 R). Sc without hairs. Genitalia
as in S. bonaerense; genital fork with stem well sclerotized and elongated, about 1/3 longer
than each posterior branch.
Male: Wing length 2.3-2.5 mm. Coloration and disposition of scutum hair as in
female. Gonocoxite similar to that of S. anamariae; gonostylus subquadrate with spur near
internal corner (Fig. 72 H).
Pupa: Cocoon projected dorsally (Fig. 73 J); weave soft with threads obvious. Length
at base 3.0-3.5 mm, at dorsum 2.8-3.0 mm, gills 4.3-5.4 mm. Gill with 2 thick branches,
emerging from short basal trunk; gills with some constrictions similar to pseudoannula-
tions (Fig. 73 J), surface of gills with small spiculae. Frontoclypeus and thorax with
abundant platelets. Trichomes bifid and some single.
Larva: Maximum length 5.5 mm; general coloration yellowish brown with transverse
band grayish green on abdominal segments, shape as in figure 75 K. Cephalic apotome
scarcely pigmented (Fig. 75 L). Antennae surpassing apex of cephalic fan stem; ratio of
proximal, medial and distal segments = 1:0.7-0.8:1.1-1.2 (Fig. 75 M). Cephalic fan with 45
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 317

rays. Mandible as in S. incrustatum. Hypostomium as figure 75 N. Postgenal bridge about as


long as hypostomium. Anal sclerite with hairs single or bifid, longer than in S. incrustatum.
Anal ring with about 65 rows of 12 hooks each. Rectal papillae unknown.
Distribution: Brazil: Mato Grosso do Sul, Minas Gerais, Rio de Janeiro, São Paulo.
Bionomics: This species breeds in small creeks, with moderate-flowing, clear water.
Female feeding habits are unknown.
Discussion: The adults are very close to those of S. (Notolepria) species, S.
incrustatum, S. anamariae and S. brevifurcatum. Simulium (Notolepria) species are smaller but
have a more acuminate paraproct. The S. incrustatum scutum is similar in appearance, but
the hair groups are not aligned in rows. Simulium anamariae has golden hair groups on the
scutum and S. brevifurcatum a grayish spot on the pronotal corner. The pupal gill shape
permits it to be differentiated from other Neotropical black fly species.

Simulium (Psaroniocompsa) brevifurcatum LUTZ (Figs. 71 S, T; 73 I.)


Simulium brevifurcatum LUTZ, 1910: 259, 263 , 264; PINTO, 1931: 694; VULCANO, 1962: 255; 1967: 8.
Simulium (Psaroniocompsa) brevifurcatum: COSCARÓN & WYGODZINSKY, 1984: 75; PY-DANIEL et al., 1985:
96-97; COSCARÓN, 1987: 23; 1991: 191-193; COSCARÓN & COSCARÓN ARIAS, 1997: 115;
CROSSKEY & HOWARD, 1997: 56; 2004: 50.
Psaroniocompsa brevifurcata: PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 2004: 123.
Female: Wing length 2.5-2.6 mm. Scutum, scutellum and metanotum velvety black;
scutum without anterior cuneiform silvery spots, and hair groups appearing like scales with
metallic golden violaceous coloration, ordered in longitudinal rows. Pronotum corners with
grayish spots. Abdomen black with I-IV segments velvety; subsequent segments with
cereous sheen; II-V with silvery large dorso-lateral spot on each side; Sc with 2-3 setae near
base. Legs yellowish with some portion darkened and abundant spatuliform hairs. Frons
relatively narrow and convergent (Fig. 71 S). Fronto-ocular triangle wider than high but
deeper than in remaining species in subgenus (Fig. 71 T). Genitalia as in S. bonaerense, but
paraproct shorter, slowly surpassing cercus ventral border; genital fork as in S. auristriatum.
Male: Wing length 2.3 mm. Coloration as in female; disposition of hair groups less
orderly than in female. Legs with abundant brownish spatuliform hairs. Gonostylus
subquadrate, with small median depression on distal rim and robust subapical spur
positioned as in S. limbatum.
Pupa: Cocoon laterally expanded and with dorsal projection (Fig. 73 I); weave thick
with visible threads; length at base 3.5-4.0 mm; at dorsum 3.8 mm; gills 3.4-4.0 mm. Gill
with 6 branches with dorsal secondary branch bifurcated very close to base (Fig. 73 I); all
filaments markedly expanded laterally. Frontoclypeus and exposed portion of thorax with
small and abundant platelets; frontoclypeus and trichomes bifid.
Larva (based on PY-DANIEL et al., 1985): Maximum length 5.7 mm. Body integument
with spiculae of 4-6 branches. Ratio of antennal segments: proximal, medial and distal =
1:0.9:0.9-1.0; cephalic fan with 52-53 rays. Mandible with latero-mandibular process
straight, surpassing apex. Rectal papillae with 68 rows and 10-12 hooks each.
Distribution: Argentina: Misiones; Brazil: Paraná, Rio Grande do Sul, Rio de
Janeiro, São Paulo.
318 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Bionomics: This species breeds in clear creeks of moderate to slow flow. Female
feeding habits are unknown.
Discussion: The adults closest to this species are those of S. anamariae and S.
auristriatum; differences were explained in those species. The cocoon and pupal gill
branching pattern permit differentiation of this species.

Simulium (Psaroniocompsa) schmidtmummi WYGODZINSKY (Figs. 72 I; 73 K; 74 H, K.)


Simulium (Psaroniocompsa) schmidtmummi WYGODZINSKY, 1973: 1-11; COSCARÓN & WYGODZINSKY,
1984: 77; COSCARÓN, 1987: 23; 1991: 193; COSCARÓN & COSCARÓN ARIAS, 1997: 115; CROSSKEY
& HOWARD, 1997: 57; 2004: 50.
Female (based on WYGODZINSKY, 1973): Wing length 3.0-3.3 mm. Frons and clypeus
from silver gray to nacreous. Scutum velvety black with 1+1 anterior silvery subtriangular
spots. Hair appressed, isolated or in small groups somewhat longitudinally arranged, but
not forming distinct lines. Pleurae silver gray to nacreous; scutellum velvety black.
Abdomen with tergite I from piceous to black; II-V blackish with III-V velvety black at
center. Tergite II with 1+1 large silver whitish spots; III-V with hind border silvery; VI-IX
piceous, polished. Frons relatively wide. Sc glabrous. Claws with minute subbasal tooth;
leg hairs spatuliform and single. Eighth sternite with 10 setae on each side; gonapophyses
with internal margin subparallel and abundant microtrichiae; paraproct very short; genital
fork with slender and well-sclerotized stem.
Male: Wing length 3.0 mm. General coloration as in female. Abdomen velvety black.
Segments II, IV-IX with 1+1 silver white lateral spots, segments IV and V not distinct.
Ratio of length/width of hind basitarsus = 5.8-6.6; spatuliform hair more abundant than
in female, especially on femora and tibiae. Gonostylus subquadrate (Fig. 72 I), remaining
genitalia parts similar to those of S. bonaerense.
Pupa: Cocoon with anterodorsal projection with blunt apex (Fig. 73 K). Cocoon
translucent with threads evident. Length at dorsum 3.5-4.0 mm; at base 4.0-4.2 mm; gills
4.3 mm. Gill with 4 filaments, branching from base, with very short primary branches (Fig.
74 H). Frontoclypeus and thorax with moderately abundant small irregularly arranged
platelets; head trichomes with simple to four branches; thoracic trichomes with 4-12
branches, some of them clavate at apex (Fig. 74 K).
Larva: Maximum length 6.5 mm. General coloration yellowish white slightly reddish
posteriorly. Body integument glabrous. Ninth segment with 1+1 ventral tubercles. Ratio
of antennal segments: proximal, medial and distal = 1:1.1:1.2. Mandible marginal teeth 2
in number, relatively thin, especially second, sometimes there is another very small third
tooth. Cephalic fan with 50-55 rays. Postgenal cleft deeper than postgenal bridge length.
Anal ring with about 66 rows of 14-15 hooks each. Anal sclerite with single hairs. Rectal
papillae with 12-16 diverticula on each lobe.
Distribution: Colombia: Boyaca, Cundinamarca, in the Páramo area between
3050-3450 m.
Bionomics: The larvae live on trailing grasses and stones in clear, small streams
about 1 m wide and 0.15 m deep, with temperatures of about 9-12 °C. Female feeding
habits are unknown.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 319

Discussion: Several species with hairs grouped and simulating scales on the scutum
can confuse identification. The golden to bronze hair coloration helps to separate this
species from S. incrustatum, the closest species. Simulium auristriatum and S. brevifurcatum
don’t show 1+1 anterior cuneiform spots. The pupa gill with 4 branches allows it to be
separated from S. (Psaroniocompsa) and S. (Notolepria) species.

Simulium (Psaroniocompsa) stellatum GIL ACEVEDO, FIGUEIRÓ & MAIA HERZOG


Simulium (Psaroniocompsa) stellatum GIL ACEVEDO et al., 2005 (922): 1-12.
(Based on original description).
Female: Wing length 2.5 mm. Scutum totally black with lateral and posterior margin
silver pruinose; pilosity golden, grouped hair arranged in longitudinal rows. Abdomen
silvery pruinose with segments I-IV velvety, remaining black shiny. Claw with small
subbasal tooth. Genitalia as in S. anamariae.
Male: Wing length 1.9 mm. Color and disposition of scutum hairs as in female.
Gonostyle subquadrate with tiny sub apical spur; ventral plate with smooth pointed apex.
Pupa: Cocoon with dorsal projection, surpassing cocoon base, weave compact and
thin, length at base 3.5-4.2 mm; gill length 3.4-4.1 mm. Gill with 4 branches emerging
from short trunk.
Larvae: Maximum length 6.0-7.2 mm. Cephalic apotome with one median longitudi-
nal 1+1 anterolateral and 1+1 sublateral positive spots. Antenna with distal segment
relatively long, ratio of proximal, medial and distal segments = 1:1:1.2: postgenal cleft
rounded apically. Ninth segment with 1+1 ventral tubercles; anal ring with 67 rows of 9-
13 hooks each; rectal papillae with 8-10 diverticula on each lobe.
Distribution: Brazil: Minas Gerais, Rio de Janeiro.
Bionomics: This species was collected in springs at about 2000 m in altitude,
attached to aquatic vegetation. No information about female feeding habits is available.
Discussion: Adults are similar to those of S. anamariae, which has 6 branches in the
pupa gill. The presence of 4 pupal gill branches makes S. stellatum similar to S.
schmidtmummi and S. (SIOLII) group species; these can be differentiated by the absence of
an anterodorsal process on the pupal cocoon and some other characters such as scutum
coloration, gonostyle shape and larval morphology. The closest species is S. schmidtmummi,
with few differences such as the scutum with 1+1 anterior silvery subtriangular spots, the
trichomes not arranged in rows and the pointed apex of the larval postgenal cleft.

SIOLII species group (Figs. 75 O-Q.)


The S. SIOLII species group has similarities with S. (Psaroniocompsa) s. str. (COSCARÓN,
1987), S. (Cerqueirellum) and S. (Coscaroniellum) and this is the reason that CROSSKEY &
HOWARD (1997, 2004) put all the species groups in S. (Psaroniocompsa) s. l.

The following diagnosis includes the characters that help to define the SIOLII
species group.
320 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Female: Small species (maximum wing length 1.7 mm). Scutum blackish with 1+1
silvery submedian stripes that join with posterior grayish area. Basal portion of
cibarium with sclerotized border and acute teeth with median area scarcely depressed,
with small plates. Fronto-ocular triangle relatively deep. Claws without subbasal tooth.
Genitalia as in S. (Psaroniocompsa) s. str.
Male: Scutum blackish with 1+1 submedian anterior silvery cuneiform spots from
narrow to wide joining with lateral grayish area. Gonostylus subtriangular, with one apical
strong and blunt spur, and a dorso-longitudinal carina.
Pupa: Cocoon slipper shaped. Frontoclypeus and thorax with trichomes single or
bifid and tubercles from rounded to acuminate, absent, on anterior and central area of
thorax; gill with 4-6 filiform branches.
Larva: Cephalic apotome with positive spots. First abdominal segment darkened
like band around body. Body segments II-VIII with subconical to rounded dorsal
protuberances and integument with abundant and large setae. Distal antennal segment
longer than other two. Mandible with 1-2 latero-mandibular processes; without supra-
marginal setae.
Five species are included: S. siolii, S. lourencoi, S. damascenoi, S. tergospinosum and S.
guaporense.

Simulium (Psaroniocompsa) siolii PY-DANIEL (Figs. 73 L; 75 O-Q.)


Simulium (Psaroniocompsa) siolii PY-DANIEL, 1988: 294; CROSSKEY & HOWARD, 1997: 57; 2004: 51.
Simulium siolii: HAMADA et al., 2003: 353.
Psaroniocompsa siolii: PY-DANIEL & MOREIRA SAMPAIO, 1995: 119.
(Based on PY-DANIEL, 1988).
Female: General coloration black. Scutum blackish bordered laterally and posterior-
ly by gray pollinose band; frontal illumination shows 1+1 silvery pollinose longitudinal
stripes from anterior to posterior border; pilosity composed of recumbent yellowish-
green hair. Pleura blackish with gray pollinosity. Frons relatively wide; fronto-ocular
triangle obsolescent. Sc and basal section of R without hair. Legs with hairs single and
spatuliform. Claw without subbasal tooth. Eighth sternite with 8-10 setae on each side;
gonapophyses with abundant microtrichiae and internal margin parallel; paraproct short.
Genital fork as in S. auripellitum.
Male: Scutum black with 1+1 anterior cuneiform grayish pruinose spot, that with
some illumination joins grayish posterior area; pilosity as in female. Abdomen brown.
Ratio of length/width of hind basitarsus = 6.3-7.0. Gonocoxite longer than gonostylus;
gonostylus subtrianglular with longitudinal carina and strong apical spur.
Pupa: Cocoon slightly projected ventrally, and with small dorsal projection (Fig. 73
L); anterior border reinforced. Length at base 2.5-3.3 mm; at dorsum 2.0-2.5 mm; gills 2.3-
2.5 mm. Gills with 4 terminal branches, with relatively short primary branch; bifurcation
of secondary branches at short distance from base. Frontoclypeus and exposed portion
of thorax with relatively low number of small tubercles, irregularly arranged in groups;
head trichomes single or bifid, thoracic trichomes single to trifid.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 321

Larva: Maximum length 3.4-4.1 mm. Cephalic apotome with positive spots diffuse,
and abundant single and bifid trichomes. Body with acuminate dorsal protuberances 2+2
on segments II-V and 1+1 on VI; ninth segment with 1+1 ventral tubercles (Fig. 75 O).
Body integument with abundant hair and scales with several acute projections (Fig. 75 P).
Antennae surpassing apex of cephalic fan stem; ratio of proximal, median and distal
segments = 1:0.9-1.2:0.8-1.1. Mandible with 2-3 very thin marginal teeth and one curved
latero-mandibular tooth surpassing teeth apical border (Fig. 75 Q). Hypostomium with 4-
7 setae on each side; postgenal cleft deep, with postgenal bridge about 1/4 hypostomium
length. Anal sclerite with hair single or bifid not obvious. Anal ring with about 63-66 rows
with 9-13 hooks each. Rectal papillae with 11-16 diverticula on each lobe.
Distribution: Brazil: Amapá, Mato Grosso, Rondônia.
Bionomics: This species breeds in small creeks flowing over deciduous leaves.
Discussion: Simulium tergospinosum is the closest species, but it has gill branches
bifurcated very close to the base and the gill is shorter than the cocoon. Simulium
schmidtmummi is also close, but can be differentiated by the gill branching very close to
the base, absence on the larval abdomen of dorsal prominences and abundant and long
trichomes, postgenal bridge longer than the hypostomium and a trapezoidal male
gonostylus.

Simulium (Psaroniocompsa) lourencoi PY-DANIEL


Simulium (Psaroniocompsa) lourencoi PY-DANIEL, 1988: 300 ; CROSSKEY & HOWARD, 1997: 57; 2004: 51.
Psasroniocompsa lourencoi: PY-DANIEL & MOREIRA SAMPAIO, 1995: 119.
(Based on PY-DANIEL, 1988).
Female and Male: A short abstract of a Congress communication (HAMADA &
AZEVEDO, 2002) is not sufficient to make a diagnosis.
Pupa: Cocoon elongated ventrally a little more than in S. siolii and with small
projection dorsally, anterior border scarcely reinforced; length at dorsum 1.7-1.9 mm, at
base 2.0-2.1 mm, gills 1.9-2.4 mm. Cephalic and thoracic trichomes long, single or bifid.
Tubercles of frontoclypeus and thorax rounded, somewhat abundant. Gills with 4
filaments, with branching similar to that of S. siolii.
Larva: Maximum length 3.2-3.7 mm. Head and body with hair and on abdomen on
acuminate dorsal projections as in S. siolii, except on prominences of segments VII or
VIII. Cephalic apotome with positive spots. Antennal length, proportion of segments,
mandible teeth, postgenal cleft and anal sclerite with abundant hairs sclerotized like scales
are very similar to those of S. siolii. Anal ring with 61-62 rows of 12-13 hooks each. Rectal
papillae with 6-8 diverticula on each lobe. Ninth sternite with 1+1 subtrianglular tubercles.
Distribution: Brazil: Goiás.
Bionomics: Unknown.
Discussion: This species is very similar to S. siolii. Small differences in the degree of
abdominal groove sclerotization in the pupa and the greater number of and shorter rectal
papillae diverticula, plus weakly defined cephalic apotome ornamentation in the larva, are
indicated by the species author as differences between the species.
322 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Simulium (Psaroniocompsa) damascenoi PY-DANIEL (Figs. 71 U; 72 J, 73 M; 74 I.)


Simulium (Psaroniocompsa) damascenoi PY-DANIEL, 1988: 304; 1989: 506-507; CROSSKEY & HOWARD,
1997: 57; 2004: 51; HAMADA et al., 2003: 353.
(Based on PY-DANIEL, 1988 and HAMADA et al., 2003).
Female: Wing length 1.6 mm. Scutum black with 1+1 longitudinal wide silvery
stripes (Fig. 71 U). Abdominal tergites I-II brown with silvery pollinosity, III-V brown
with tergal plates bright; VI-IX with peculiarly pale greenish sheen. Fronto-ocular triangle
absent. Claws without subbasal tooth. Eighth sternite setae 9 or 10 in number on each
side; gonapophysis, paraproct and genital fork as in S. siolii.
Male: Wing length 1.4 mm. Scutum black with 1+1 antero-lateral subquadrate silvery
spots (Fig. 72 J). Pleurae brown with silver pollinosity, scutellum and metanotum brown
with grayish pollinosity. Abdomen brown. Gonostylus subtriangular with apical spur.
Pupa: Cocoon with reinforced border and very small dorsal anterior projection (Fig.
73 M), threads weakly evident. Gill with 4 branches originating from short primary
branches (Fig. 74 I). Head without platelets; exposed portion of thorax with only small
and acuminate tubercles on posterior half. Trichomes of head and thorax single or bifid.
Larva: Maximum length 3.7-3.9 mm. Abdomen with acuminate dorsal protuberanc-
es and trichomes as in S. siolii. Protuberances on segment VIII rounded apically. Ninth
segment without ventral tubercles. Cephalic apotome with positive spots and scarce, single
trichomes. Antennae surpassing apex of cephalic fan stem; ratio of proximal, medial and
distal segments = 1:1.5-1.7:0.9-1.0. Postgenal cleft deep and postgenal bridge shorter than
half of hypostomium length. Mandible similar to that of S. siolii. Anal ring with 70-76
rows with 14-17 hooks each. Rectal papillae with 4-6 diverticula on each lobe.
Distribution: Brazil: Amapá.
Bionomics: This species breeds in rivers of about 25-27 °C and 80-100 m in width
in forests and savannas, with rocky bottoms, attaching to Podostomaceae, deciduous
leaves and streamside vegetation. Female feeding habits are undescribed.
Discussion: The species is similar to S. siolii, S. lourencoi, S. guaporense, S. schmidtmum-
mi and S. stellatum. The first three species show platelets on the pupal frontoclypeus and
the exposed portion of the thorax. Simulium schmidtmummi and S. stellatum also have a
dorsal anterior projection on the cocoon (but it is larger), absence of dorsal protuberances
in the larva, and a trapezoidal gonostylus in the male. In S. siolii and S. lourencoi, the medial
antennal segment is proportionally shorter and bifurcation of the gill secondary branches
are at a greater distance from the base. Simulium guaporense has 6 pupal gill branches.

Simulium (Psaroniocompsa) tergospinosum HAMADA (Figs. 72 K-M; 73 N.)


Simulium (Psaroniocompsa) tergospinosum HAMADA, 2000: 819; CROSSKEY & HOWARD, 2004: 51.
(Based on HAMADA, 2000).
Female: Wing length 1.7 mm. General color black. Frons and clypeus with silvery
blue pollinosity. Scutum black with 1+1 thick light silvery stripes, similar to that of S.
damascenoi, with thin golden reddish hair unevenly distributed. Pleurae dark brown with
silver pollinosity. Scutellum and metanotum brownish. Sc bare. Legs yellowish brown with
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 323

tarsi black on fore leg; hind leg with femur, tibial apex and terminal tarsomeres dark
brown. Abdomen brown with greenish membranous area and basal fringe highlighted
with gold. Tergite II with silvery blue pollinosity; VI-VIII slightly bright. Fronto-ocular
triangle obsolescent; claw without subbasal tooth. Eighth sternite with about 14 setae on
each side; gonapophysis, paraproct and genital fork similar to those of S. siolii.
Male: Wing length 1.7 mm. General coloration as in female. Antenna, hair and
insertion area with silvery-blue pollinosity. Scutum black with golden reddish hair and 1+1
silvery, anterior submedian cuneiform spots, reaching nearly middle of scutum length
(Fig. 72 K). Scutellum and metanotum brownish with silver pollinosity. Leg coloration as
in female. Abdomen dark brown with basal fringe gold; tergites II, V-VIII with 1+1 lateral
silvery spots. Gonostylus longer than half of gonocoxite length (Fig. 72 L), elongate,
subtriangular with dorsal internal longitudinal ridge and strong apical spur (Fig. 72 M).
Pupa: Cocoon with basal anterior enlargement and without dorsal projection (Fig. 73 N),
with reinforced anterior border, and weave with evident threads. Length at base 2.8 mm, at
dorsum 2.4 mm, gills 2.0 mm. Gills with 4 filaments bifurcated close to base with primary
branches very short, similar to those of S. damascenoi. Frontoclypeus and anterior portion of
thorax without tubercles; posterior exposed portion of thorax with rounded or pointed
tubercles. Head trichomes simple to trifid, thoracic trichomes from simple to with 4 branches.
Larva: Maximum length 4.3-4.6 mm. Cephalic apotome with light positive spots and
small simple trichomes. Antennae surpassing cephalic fan stem apex; ratio of proximal,
medial and distal segments = 1:0.8-1.2:1.0-1.1. Mandible with thin marginal teeth and 1-2
latero-mandibular processes, simple or bifid, reaching mandible teeth inferior margin.
Body with abundant scales multiply branched. Abdomen with 1+1 acuminate prominenc-
es dorsally and laterally on segments II-VI and only dorsally on segment VII. Ninth
segment with 1+1 ventral tubercles. Anal sclerite area with short branched setae. Anal ring
with 62-69 rows with 12-15 hooks each. Rectal papillae with 8-11 diverticula on each lobe.
Distribution: Brazil: Amazonas.
Bionomics: This species was collected in large black water river waterfalls and creeks
in the Amazonian region, on Podostemaceae and fallen leaves. Females bite humans.
Discussion: The closest species is S. damascenoi but it can be differentiated by the
presence in the larva of abdominal dorsal prominences on segment VIII and absence of
ventral tubercles on the ninth segment. Simulium siolii and S. lourencoi have the pupal gill
bifurcating far from the base, and gills as long as or longer than the cocoon. Simulium
schmidtmummi has a large dorsal projection on the cocoon and male subquadrate gonostylus.

Simulium (Psaroniocompsa) guaporense PY-DANIEL


Simulium (Psaroniocompsa) guaporense PY-DANIEL, 1989b: 502; CROSSKEY & HOWARD, 2004: 51.
Simulium (Psaroniocompsa) siolii PY-DANIEL, 1988: (in part).
Psaroniocompsa guaporense PY-DANIEL & MOREIRA SAMPAIO, 1995: 119.
(Based on PY-DANIEL, 1989).
Female and Male: Unknown.
Pupa: Cocoon with reinforced border and small antero-dorsal projection,
leaving free 2/3 portion of thorax; weave with evident threads. Gill with 6 terminal
324 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

branches; primary dorsal branch bifurcated close to base. Primary ventral and
secondary branches bifurcated near half of total length. Frontoclypeus and exposed
portion of thorax with moderate number of small tubercles rounded or acuminate,
arranged in irregular groups; tubercles of thorax acuminate on anterior half;
trichomes with 3-4 branches.
Larva: Maximum length 4.3 mm. Abdomen dorsally with hair and protuberances
as in S. siolii, but with 1+1 protuberances on tergite I, presence of well-sclerotized
scales, small ventral expansion of each abdominal segment and absence of protuber-
ances on segment VIII. Cephalic apotome scarcely ornamented but with positive
spots and abundant single elongated hairs. Postgenal cleft deep with postgenal bridge
about 1/5 of hypostomium length. Antennal apex reaching apex of cephalic fan stem;
ratio of proximal, medial and distal segments = 1:0.9:1.3. Mandible and anal sclerite
as in S. siolii. Anal ring with 79 rows with 10-15 hooks each. Rectal papillae diverticula
number unknown.
Distribution: Brazil: Rondônia.
Bionomics: Unknown.
Discussion: The SIOLII group species are very similar. The presence of six gill
branches allows the pupa to be readily separated from those of the other species. In the
larva, the other species have fewer integument scales that are less sclerotized on the
terminal portion of the abdomen, and also have dorsal protuberances on segment VIII.

Species inquirendae

Simulium catarinense PINTO


Simulium catarinense PINTO, 1931: 695; COSCARÓN, 1991: 193.
Simulium (Psaroniocompsa) catarinense: COSCARÓN & WYGODZINSKY, 1984: 77.
The description is based on a male, with illustrations that allow it to be placed in this
subgenus, but which makes it impossible to determine the species level.
Distribution: Brazil: Santa Catarina.

Simulium guttatum (ENDERLEIN)


Pliodasina guttata ENDERLEIN, 1936: 124.
Simulium guttatum: VARGAS & DIAZ NÁJERA, 1953b: 137; STONE, 1963: 14; COSCARÓN & WYGODZIN-
SKY, 1984: 77; COSCARÓN, 1987: 24; 1991: 193.

The type material is from Paraguay and deposited at the Berlin Museum. VARGAS &
DIAZ NAJERA stated that it is close to S. incrustatum, and STONE considered it as S.
incrustatum. We examined the female but it is not well preserved, the coloration is deficient
and does not permit us to determine to which species it corresponds. We prefer to
maintain it as species inquirendae.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 325

Simulium (Cerqueirellum) PY-DANIEL (Figs. 76-79; 143.)


Type-Species: Simulium amazonicum GOELDI, 1905: 138.
Female: Relatively small, maximum wing length 2.8 mm. General coloration black.
Frons and clypeus gray nacreous. Scutum black bordered laterally and posteriorly by
silvery pollinose area, and with 1+1 thick wide submedian silvery bands reaching posterior
grayish area. These silvery longitudinal bands generally have some light and anterior,
internal elongated darker, shadow-like spots. Abdomen with tergites I-IV velvety black,
with lateral and posterior borders light grayish, subsequent tergites black with cereous
sheen; tergite II with metallic greenish blue sheen at sides. Frons generally wider than
high; fronto-ocular triangle absent or very reduced. Female antennae as long as male
antennae. Infra-frontal sutures not developed. Sc and basal sector of R without hair; legs
with hairs single and spatuliform; calcipala well developed and claws generally without
subbasal tooth. Basal portion of cibarium well sclerotized, with abundant acute teeth in
several rows, which reach laterally to cornuae and leave (depressed) median area free.
Gonapophyses with abundant microtrichiae, straight distal border, and internal rim
slightly subparallel; paraproct shorter than high, cercus with subcircular border.
Male: Scutum ornamentation as in female or totally black with only 1+1 anterior
cuneiform silvery spots. Abdomen black with 1+1 gray nacreous spots on segments II, VI,
VII and very small on VIII. Gonocoxite subquadrate wider than long; gonostylus about as
long as gonocoxite, subtriangular or subtrapezoidal elongated with 1-2 (exceptionality to
5-7) subconical spurs on or near apex; endoparameres with abundant hooks.
Pupa: Cocoon generally slipper shaped, thick weave and frequently anteriorly
reinforced. Head with 2 pairs of frontal and 1 pair facial simple trichomes; 5-6 thoracic
trichomes simple, bifid, or up to 4 branches. Frontoclypeus and thorax with smooth
tubercles, not acute; gill with 4-10 branches.
Larva: Maximum length 5.5 mm. Body generally with light coloration showing dark
band around first abdominal segment. Body integument without scales or spines, but with
some hair. Cephalic apotome, generally scarcely ornamented and without positive spots;
with obvious hairs. Gular cleft deep with rounded margin, postgenal bridge about 1/4 or
more shorter than hypostomium. Hypostomium median tooth generally below lateral
teeth. Antenna as long as or longer than cephalic fan stem, medial article generally longer
or as long as distal article. Mandible with only one row of internal teeth; with or without
latero-mandibular process and with one row of setae on supramarginal inferior border.
Ninth segment with or without pair of ventral tubercle. Anal sclerite with relatively short
ventral branches, with enlarged union with dorsal branches. Anal ring with 50-70 rows of
8-14 hooks each. Rectal papillae with 3-9 diverticula.
Distribution: Found from central Argentina to Mexico, with greater distribution in
the intertropical region (Fig. 143).
Discussion: This subgenus includes 11 species. CROSSKEY & HOWARD (1997, 2004)
include these species in the AMAZONICUM species group of S. (Psaroniocompsa). This
subgenus is differentiated from Psaroniocompsa by the presence on the female scutum of 1+1
intervittae submedian silvery bands with cuneiform elongated interior spots slightly
darkened like a shadow; in the male by the subtriangular or subtrapezoidal elongated
326 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

gonostylus with 1-7 spurs; in the pupa by thoracic trichomes with 1-4 branches; and in the
larva by a supramarginal setal row on the mandibular inferior border. PY-DANIEL (1988) gave
a detailed description of the subgenus and of several species of this taxon, including a
species key for all life stages except the egg. SHELLEY et al. (1997, 2000, 2002) also treated
some of the species. Here we include a diagnosis of the species, with a key for the different
stages. We also include S. delponteianum, the southernmost species, noting several differences
from the common species, such as a more grayish colored scutum, greater number of spurs
on the gonostylus and the absence of supramarginal setae on the mandible; and S.
argentiscutum with different scutum ornamentation, and a shoe-shaped cocoon with non-
filamentous gill branches. In both cases, this is the closest subgenus in which they can be
placed. Also, can be objected the inclusion of S. minusculum, differentiated by a subquadrate
gonostylus and absence of supramarginal setae on the larval mandible. Another difficult
species is S. quadrivittatum, which has been placed here in S. (Coscaroniellum).

KEYS TO SIMULIUM (CERQUEIRELLUM) SPECIES

Females
1 Scutum with black, narrow median stripe reduced to thin line on anterior 2/3 and
1+1 black short sublateral stripes; with frontal light showing 1+1 cuneiform
anterior submedian intervittae blackish stripes reaching to half length of scutum
(Fig. 76 U); with posterior light, the black median and posterior sublateral stripes
shorter and intervittae cuneiform spots lighter and shorter, with nacreous sheen
(Fig. 76 V) ........................................................................................ Simulium argentiscutum
– Scutum with black, thick median stripe, and 1+1 black sublateral stripes covering
2/3 of scutum, anterior intervittae spots present or absent, if present, they can be
short (Fig. 76 F) or large surpassing half of scutum length (Figs. 76 C, H),
sometimes varying according to direction of light ..................................................... 2
2(1) Scutum with 1+1 black sublateral stripes short, divergent, covering about 1/2 to
1/3 of scutum, depending on direction of light; with anterior light, the 1+1 short
blackish diffuse intervittae spots (Fig. 76 F) evident; with posterior light,
intervittae generally not evident ................................................... Simulium amazonicum
– Scutum with 1+1 black sublateral stripes longer, surpassing 2/3 of scutum; with
or without anterior intervittae spots of different length and shape according to
light direction ...................................................................................................................... 3
3(2) Frons pollinose without sheen; scutum with abundant pilosity, sometimes not
allowing limit of stripes to be seen, giving them lead gray tonality (Fig. 76 Y);
claws generally with subbasal tooth (Fig. 76 Z); fronto-ocular triangle evident (Fig.
76 W) ............................................................................................... Simulium delponteianum
– Frons with nacreous or metallic sheen; scutum with sparse pilosity permitting full
extension of stripes to be seen; claws generally without subbasal tooth; fronto-
ocular triangle absent (Fig. 76 G) ................................................................................... 4
4(3) Anterior cuneiform intervittae spots of scutum shadowed anteriorly, short, not
surpassing anterior 1/3 of scutum, varying according to direction of light (Figs.
76 A, B) ...................... Simulium minusculum, Simulium roraimense, Simulium venezuelense
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 327

– Anterior cuneiform intervittae spots of scutum shadowed anteriorly, elongate,


surpassing anterior 1/2 of scutum, varying according to direction of light (Figs.
76 C, E, H, I, R) ................................................................................................................. 5
5(4) Wing length about 2.5 mm ................................................................ Simulium chaquense
– Wing length about 1.3-2.1 mm ....................................................................................... 6
6(5) Wing length about 1.7-2.1 mm ........ Simulium oyapockense, S. cuneatum, S. ganalesense
– Wing length about 1.3-1.5 mm ...................................................... Simulium sanguineum

Males
1 Scutum with 1+1 anterior silvery spots joining grayish posterior area (Fig. 77 F)
................................................................................................................................................ 2
– Scutum with 1+1 anterior silvery spots, not joining grayish posterior area (Figs. 77
A, C, D, L, R) ...................................................................................................................... 4
2(1) Wing length 2.2-2.3 mm; gonostylus generally with 2 apical spurs (Fig. 77 J) .......
.................................................................................................................. Simulium chaquense
– Wing length 1.2-1.6 mm; gonostylus generally with 1 apical spur as in Figure 77 O
................................................................................................................................................ 3
3(2) Scutum with 1+1 silvery submedian stripes relatively wide (Fig. 77 F) ...................
................................................................................................................ Simulium ganalesense
– Scutum with 1+1 silvery submedian stripes relatively thin, especially on posterior
1/2 (Fig. 77 E) ..................................................................................... Simulium roraimense
4(1) Gonostylus with 5-10 spurs (Fig. 77 T); scutum with silvery submedian stripes
thin anteriorly (Fig. 77 R) ............................................................ Simulium delponteianum
– Gonostylus with only 1 spur; scutum with silvery submedian stripes wide
anteriorly, subtriangular ..................................................................................................... 5
5(4) Scutum with 1+1 silvery submedian stripes not reaching 1/2 of scutum length
(Fig. 77 A) ............................................................................................................................ 6
– Scutum with 1+1 silvery submedian stripes surpassing 1/2 of scutum length
(Figs. 77 C, D, K-M) .......................................................................................................... 7
6(5) Scutum with 1+1 anterior silvery spots subtrianglular; gonostylus subtriangular
with apical spur .................................................................................. Simulium venezuelense
– Scutum with 1+1 anterior silvery spots comma shaped (Fig. 77 A); gonostylus
subtrapezoidal with subapical spur (Fig. 77 B) ........................... Simulium minusculum
7(5) Scutum with 1+1 silvery submedian stripes not reaching 1/2 of scutum length
.............................................................................................................. Simulium amazonicum
– Scutum with 1+1 silvery submedian stripes reaching nearly to grayish posterior
area of scutum (Figs. 77 C, D, K, Q) ............................................................................ 8
8(7) Wing length 1.7-2.0 mm ...................................................................... Simulium cuneatum
– Wing length 1.4-1.5 mm ................................................................................................... 9
9(8) Scutum with 1+1 black sublateral stripes wide anteriorly, reaching nearly to
anterior border (Figs. 77 C, D) ...................................................... Simulium oyapockense
– Scutum with 1+1 black sublateral stripes narrow anteriorly, not reaching (Fig. 77
Q), or only laterally reaching (Fig. 77 K) nearly to anterior border ...................... 10
328 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

10(9) Scutum with 1+1 silvery submedian stripes posteriorly reaching or very close to
posterior grayish area (Fig. 77 K) .................................................. Simulium sanguineum
– Scutum with 1+1 silvery submedian stripes posteriorly not reaching posterior
grayish area (Fig. 77 Q) ................................................................. Simulium argentiscutum

Pupae
1 Gill with 10 branches (Figs. 78 J, L) ................................................. Simulium cuneatum
– Gill with 4-8 branches ....................................................................................................... 2
2(1) Gill with 4 branches ......................................................................... Simulium venezuelense
– Gill with 6-8 branches ....................................................................................................... 3
3(2) Gill with 6 branches (Figs. 78 B, E, N, O) ................................................................... 4
– Gill with 8 branches (Fig. 78 H) ..................................................................................... 8
4(3) Cocoon shoe shaped (Figs. 78 M); weave structure spongy, without evident
threads; gill short with wrinkled border (Fig. 78 N) ............... Simulium argentiscutum
– Cocoon slipper shaped (Figs. 78 A, C, G, J), weave structure compact, with
evident threads; gill elongated with smooth border (Figs. 78 B, E, O) .................. 5
5(4) Species relatively large; cocoon length basally 3.6-4.6 mm; gill length 1.8-2.5 mm;
thoracic trichomes simple (Fig. 78 O) ...................................... Simulium delponteianum
– Species relatively small; cocoon length basally 2.0-2.2 mm; gill length 0.7-1.8 mm;
thoracic trichomes with 1-4 branches (Figs. 78 B, E) ................................................ 6
6(5) Frontoclypeus and exposed portion of thorax with few tubercles ..... Simulium minusculum
– Frontoclypeus and exposed portion of thorax with abundant tubercles (Fig. 78 D) ....... 7
7(6) Frontoclypeus and exposed portion of thorax with rounded tubercles ..................
........................................................................... Simulium ganalesense, Simulium oyapockense
– Frontoclypeus and exposed portion of thorax with some rounded but acuminate tubercles
on apex of frontoclypeus and dorsomedian area of thorax ................. Simulium roraimense
8(3) Species relatively large; cocoon length basally 3.0-3.5 mm, 2.5-2.7 mm dorsally;
gills 1.8-2.0 mm .................................................................................... Simulium chaquense
– Species relatively small, cocoon length basally 2.0-2.7 mm, dorsally 1.6-2.3 mm,
gills 1.0-1.8 mm .................................................................................................................. 9
9(8) Dorsal secondary branch single, ventral branch bifid (Fig. 78 I) ....... Simulium sanguineum
– Dorsal secondary branch bifid, ventral branch single .............. Simulium amazonicum

Larvae
Unknown: Simulium venezuelense.

1 Antenna not surpassing apex of cephalic fan stem; postgenal bridge about 1/5-1/6 of
hypostomium length; ninth segment without ventral tubercles ..... Simulium argentiscutum
– Antenna surpassing apex of cephalic fan stem; postgenal bridge about 1/2-1/6 of
hypostomium length; ninth segment generally with ventral tubercles .................... 2
2(1) Anal ring with 70-78 rows with 14-18 hooks each; mandible without row of
supramarginal setae ....................................................................... Simulium delponteianum
– Anal ring with 40-66 rows with 8-13 hooks each; mandible generally with row of
supramarginal setae ............................................................................................................ 3
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 329

3(2) Mandible without row of supramarginal setae ........................... Simulium minusculum


– Mandible with row of supramarginal setae .................................................................. 4
4(3) Cephalic fan with 17-22 rays ........................................................................................... 5
– Cephalic fan with 37-45 rays ........................................................................................... 8
5(4) Mandible without latero-mandibular process .............................................................. 6
– Mandible with 1-2 latero-mandibular processes ......................... Simulium sanguineum
6(5) Maximum length 4.5-5.0 mm; cephalic fan with 34-40 rays ....... Simulium chaquense
– Maximum length 3.6-4.3 mm; cephalic fan with 16-22 rays ..................................... 7
7(6) Postgenal bridge about 1/3 or shorter than hypostomium length; anal ring with
52-58 rows of hooks ....................................................................... Simulium amazonicum
– Postgenal bridge about 1/2 hypostomium length; anal ring with 60-70 rows of
hooks ........................................................................................................ Simulium cuneatum
8(4) Mandible with 2 latero-mandibular processes; cephalic fan with 27-38 rays ..........
............................................................................................................... Simulium oyapockense
– Mandible with 1 latero-mandibular process; cephalic fan with 37-42 rays .............
.............................................................................. Simulium roraimene, Simulium ganalesense

As the species of this subgenus are especially differentiated by the adult scutal
ornamentation and pupal gills, we will reduce the diagnosis to the principal characters.

CLAVES DE ESPECIES DE SIMULIUM (CERQUEIRELLUM)


Hembras
1 Escudo con una banda negra mediana angosta, reducida a una fina línea en 2/3
anteriores y 1+1 banda negra corta sublateral; con luz frontal presenta 1+1
bandas negruzcas cuneiformes anteriores submedianas, llegando hasta la mitad
del largo del escudo (Fig. 76 U); con luz posterior las bandas mediana y posterior
sublaterales, se presentan más cortas y las manchas submedianas anteriores
cuneiformes también cortas y más claras, con brillo nacarado (Fig. 76 V) .............
............................................................................................................ Simulium argentiscutum
– Escudo con banda negra mediana gruesa y 1+1 bandas negras sublaterales
cubriendo los 2/3 del largo del escudo; manchas anteriores entre bandas
presentes o ausentes, si están presentes pueden ser cortas (Fig. 76 F) o largas
sobrepasando la mitad del largo del escudo (Figs. 76 C, H), a veces variando según
la dirección de la luz .......................................................................................................... 2
2(1) Escudo con 1+1 bandas negras sublaterales cortas, divergentes, cubriendo alrededor
de la mitad o un tercio del largo del escudo de acuerdo a la iluminación; con luz
anterior presenta 1+1 manchas difusas cortas entre bandas (Fig. 76 F); con luz
posterior generalmente estas manchas no son evidentes ............. Simulium amazonicum
– Escudo con 1+1 bandas negras sublaterales sobrepasando 2/3 del largo del
escudo; con o sin manchas anteriores entre bandas presentando diferente longitud
y forma según la posición de la luz ................................................................................ 3
3(2) Frente polinosa sin brillo; escudo con abundante pilosidad, a veces impidiendo
ver el límite de las bandas dando un aspecto general gris plomizo (Fig. 76 Y); uñas
330 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

generalmente con diente subbasal (Fig. 76 Z); triángulo frontoocular evidente


(Fig. 76 W) ...................................................................................... Simulium delponteianum
– Frente con brillo nacarado o metálico; escudo con escasa pilosidad permitiendo
ver la extensión de las bandas; uñas generalmente sin diente subbasal; triángulo
frontoocular ausente (Fig. 76 G) .................................................................................... 4
4(3) Manchas del escudo sombreadas, anteriores, cuneiformes entre bandas de
acuerdo a la posición de la luz son cortas, no sobrepasando el 1/3 anterior (Figs.
76 A, B) ...................... Simulium minusculum, Simulium roraimense, Simulium venezuelense
– Manchas del escudo sombreadas, anteriores, cuneiformes entre bandas, de
acuerdo a la posición de la luz son alargadas sobrepasando la mitad del largo del
escudo (Figs. 76 C, E, H, I, R) ........................................................................................ 5
5(4) Largo del ala alrededor de 2,5 mm .................................................. Simulium chaquense
– Largo del ala alrededor de 1,3-2,1 mm ......................................................................... 6
6(5) Largo del ala alrededor de 1,7-2,1 mm ...........................................................................
........................................... Simulium cuneatum, Simulium ganalense, Simulium oyapockense
– Largo del ala alrededor de 1,3-1,5 mm ........................................ Simulium sanguineum

Machos
1 Escudo con 1+1 manchas anteriores plateadas uniéndose al área posterior
grisácea (Fig. 77 F) ............................................................................................................. 2
– Escudo con 1+1 manchas anteriores plateadas no uniéndose al área posterior
grisácea (Fig. 77 A, C, D, L, R) ....................................................................................... 4
2(1) Longitud ala 2,2-2,3 mm; gonostilo generalmente con dos espolones apicales
(Fig. 77 J) ................................................................................................ Simulium chaquense
– Longitud ala 1,2-1,6 mm; gonostilo generalmente con un espolón apical similar a
(Fig. 77 O) ........................................................................................................................... 3
3(2) Escudo con 1+1 bandas plateadas submedianas relativamente anchas (Fig. 77 F)
................................................................................................................ Simulium ganalesense
– Escudo con 1+1 bandas plateadas submedianas relativamente finas, especialmente
sobre la 1/2 posterior (Fig. 77 E) ..................................................... Simulium roraimense
4(2) Gonostilo con 5-10 espolones (Fig. 77 T); escudo con bandas plateadas
submedianas finas anteriormente (Fig. 77 R) .......................... Simulium delponteianum
– Gonostilo con solamente 1 espolón; escudo con bandas plateadas submedianas
anchas anteriormente en forma triangular ................................................................... 5
5(4) Escudo con 1+1 bandas plateadas submedianas no llegando a la mitad del largo
del escudo (Fig. 77 A) ....................................................................................................... 6
– Escudo con 1+1 banda plateada submediana sobrepasando la mitad del largo del
escudo (Figs. 77 C, D, K-M) ............................................................................................ 7
6(5) Escudo con 1+1 bandas plateadas submedianas subtriangulares; gonostilo de
forma subtriangular con espolón apical ....................................... Simulium venezuelense
– Escudo con 1+1 bandas plateadas en forma de coma (Fig. 77 A); gonostilo
subtrapezoidal con espolón subapical (Fig. 77 B) ...................... Simulium minusculum
7(5) Escudo con 1+1 bandas plateadas submedianas no alcanzando a la mitad del
largo del escudo ................................................................................ Simulium amazonicum
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 331

– Escudo con 1+1 banda plateada submediana llegando cerca del área posterior
grisácea (Figs. 77 C, D, K, Q) .......................................................................................... 8
8(7) Longitud ala 1,7-2,0 mm ..................................................................... Simulium cuneatum
– Longitud ala 1,4-1,5 mm .................................................................................................. 9
9(8) Escudo con 1+1 bandas negras sublaterales anchas anteriormente y llegando
cerca del borde anterior (Figs. 77 C, D) ....................................... Simulium oyapockense
– Escudo con 1+1 bandas negras sublaterales angostas anteriormente no llegando
(Fig. 77 Q), o sólo lateralmente (Fig. 77 K) cerca del borde anterior .................. 10
10(9) Escudo con 1+1 bandas plateadas submedianas llegando posteriormente o muy
cerca del área grisácea posterior (Fig. 77 K) ............................... Simulium sanguineum
– Escudo con 1+1 bandas plateadas submedianas no llegando posteriormente al
área posterior grisácea (Fig. 77 Q) ............................................. Simulium argentiscutum

Pupas
1 Branquias con 10 ramas (Figs. 78 J, L) ............................................. Simulium cuneatum
– Branquia con 4-8 ramas .................................................................................................... 2
2(1) Branquias con 4 ramas ..................................................................... Simulium venezuelense
– Branquias con 6-8 ramas .................................................................................................. 3
3(2) Branquia con 6 ramas (Figs. 68 B, E, N, O) ................................................................. 4
– Branquia con 8 ramas (Fig. 78 H) .................................................................................. 8
4(3) Capullo en forma de zapato (Fig. 78 M); estructura del tejido esponjoso, sin fibras
evidentes; branquia corta y de borde rugoso (Fig. 78 N) ..... Simulium argentiscutum
– Capullo en forma de chinela (Figs. 78 A, C, G, J) estructura del tejido compacta,
con fibras evidentes; branquia alargada con borde liso (Figs. 78 B, E, O) ............ 5
5(4) Especies relativamente grandes, largo del capullo en la base 3,6-4,6 mm; largo
de la branquia 1,8-2,5 mm; tricomas torácicos simples (Fig. 78 O) .......................
........................................................................................................... Simulium delponteianum
– Especies relativamente pequeñas; longitud del capullo en la base 2,0-2,2m, largo
branquia 0,7-1,8 mm; tricomas torácicos con 1-4 ramas (Figs. 78 B, E) ............... 6
6(5) Frontoclípeo y porción expuesta del tórax con escasos tubérculos ..........................
............................................................................................................... Simulium minusculum
– Frontoclípeo y porción expuesta del tórax con abundantes tubérculos (Fig. 78 D)
................................................................................................................................................ 7
7(6) Frontoclípeo y porción expuesta del tórax con tubérculos redondeados ................
........................................................................... Simulium ganalesense, Simulium oyapockense
– Frontoclípeo y porción expuesta del tórax con tubérculos algunos redondeados
pero aguzados en el ápice en frontoclípeo y área dorso-mediana del tórax ...........
................................................................................................................. Simulium roraimense
8(3) Especies relativamente grandes; largo del capullo en la base 3,0-3,5 mm; en el
dorso 2,5-2,7 mm; branquias 1,8-2,0 mm ....................................... Simulium chaquense
– Especies relativamente pequeñas; largo capullo en la base 2,0-2,7 mm; en el dorso
1,6-2,3 mm; branquias 1,0-1,8 mm ................................................................................. 9
9(8) Rama secundaria dorsal simple, ventral bífida (Fig. 68 I) ........ Simulium sanguineum
– Rama secundaria dorsal bífida, ventral simple ........................... Simulium amazonicum
332 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Larvas
Desconocidos: Simulium venezuelense.

1 Antena no sobrepasando el ápice del tallo del abanico cefálico; puente postgenal
aproximadamente 1/5-1/6 del largo del hipostomio; noveno segmento sin
tubérculos ventrales ....................................................................... Simulium argentiscutum
– Antena sobrepasando el ápice del tallo del abanico cefálico; puente postgenal
aproximadamente 1/2-1/6 del largo del hipostomio; noveno segmento
generalmente con divertículos ventrales ....................................................................... 2
2(1) Anillo anal con 70-78 hileras con 14-18 ganchos cada una; mandíbulas sin hileras
de cerdas supramarginales ........................................................... Simulium delponteianum
– Anillo anal con 40-66 hileras con 8-13 ganchos cada una; mandíbula generalmente
con una hilera de cerdas supramarginales ..................................................................... 3
3(2) Mandíbula sin hilera de setas supramarginales ........................... Simulium minusculum
– Mandíbula con hilera de setas supramarginales ........................................................... 4
4(3) Abanico cefálico con 17-22 rayos ................................................................................... 5
– Abanico cefálico con 37-45 rayos ................................................................................... 8
5(4) Mandíbula sin proceso latero-mandibular .................................................................... 6
– Mandíbula con 1-2 procesos latero-mandibular ......................... Simulium sanguineum
6(5) Longitud máxima 4,5-5,0 mm; abanico cefálico con 34-40 rayos ..... Simulium chaquense
– Longitud máxima 3,6-4,3 mm; abanico cefálico con 16-22 rayos ........................... 7
7(6) Puente postgenal aproximadamente 1/3 más corto que el largo del hipostomio;
anillo anal con 52-58 hileras de ganchos .................................... Simulium amazonicum
– Puente postgenal aproximadamente 1/2 del largo del hipostomio; anillo anal con
60-70 hileras de ganchos ..................................................................... Simulium cuneatum
8(4) Con 2 procesos latero-mandibulares; abanico cefálico con 27-38 rayos ..................
............................................................................................................... Simulium oyapockense
– Con 1 proceso látero-mandibular; abanico cefálico con 37-42 rayos .......................
............................................................................. Simulium roraimense, Simulium ganalesense

Siendo que las especies de este subgénero son diferenciados por la ornamentación
del escudo de los adultos y por las branquias de las pupas, reduciremos los diagnósticos a
los caracteres principales.

Simulium (Cerqueirellum) minusculum LUTZ (Figs. 76 A, B; 77 A, B; 78 A, B; 79 D.)


Simulium minusculum LUTZ, 1910: 253; NATHAN et al., 1982: 221-222; SHELLEY et al., 1982: 4-8, 22-25;
COSCARÓN, 1983: 90.
Psaroniocompsa minuscula ENDERLEIN, 1936: 123; PY-DANIEL & MOREIRA SAMPAIO, 1995: 119;
STRIEDER, 2004: 124.
Simulium (Psaroniocompsa) minusculum PY-DANIEL, 1983: 181; CROSSKEY & HOWARD, 1997: 56; 2004:
50; SHELLEY et al., 2000: 181-182.
Simulium (Cerqueirellum) minusculum: COSCARÓN, 1987: 24; 1991: 199-200.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 333

D F
A B C

P
E O
K
H I

R
Q
L M
G

Z
U V

Y
S

T
X N J

Figure 76. Female. A-B: S. (Cerqueirellum) minusculum, scutum with different illumination. C-D: S.
(C.) oyapockense, scutum with different illumination. E: S. (C.) ganalesense, scutum. F: S. (C.)
amazonicum, scutum (redesigned from SHELLEY et al., 1982). G-K: S. (C.) chaquense; G: fronto-ocular
triangle; H-I: scutum with different illumination; J: hair of legs; K: abdomen, lateral view. L-M: S.
(C.) sanguineum, scutum with different illumination (from TIDWELL et al., 1981). N-T: S. (C.) cuneatum;
N: basal portion of cibarium; O-R: scutum with different illumination (O-P: lateral view, Q-R
dorsal view); S: eighth sternite, gonapophyses, genital fork and spermatheca; T: cercus and
paraproct (left mounted in slide). U-V: S. (C.) argentiscutum, scutum with different illumination. W-Z:
S. (C.) delponteianum; X: basal portion of cibarium; W: fronto-ocular triangle; Y: scutum; Z: claws.
334 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

A C D E K

F G
H I

L M Q
S

B N

P
J
O
T

Figure 77. Male. A-B: S. (Cerqueirellum) minusculum; A: scutum; B: gonostylus. C-D: S. (C.)
oyapockense, scutum , with different illumination. E: S. (C.) roraimense, scutum. F: S. (C.) ganalesense,
scutum. G-J: S. (C.) chaquense; G: scutum; H-I: abdomen in dorsal and lateral position; J: gonocoxite
and gonostylus. K: S. (C.) sanguineum, scutum. L-P: S. (C.) cuneatum; L-M: scutum with different
illumination; N: genitalia in ventral view showing gonocoxite, gonostylus, ventral plate and
endoparameres; O: gonostylus dorsal view; P: ventral plate. Q: S. (C.) argentiscutum, scutum. R-T: S.
(C.) delponteianum; R-S: scutum (S: lateral view); T: gonocoxite and gonostylus.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 335

Female: Wing length 1.8-2.0 mm. Frons slightly nacreous; clypeus with grayish
pruinosity. Scutum velvety black with 1+1 submedian silvery stripes reaching grayish posterior
area (Figs. 76 A, B), anteriorly on intervittae area 1+1 short cuneiform spots (lighter with
posterior illumination and blackish with anterior illumination) that can be more elongated as in
figure 76 B. Fronto-ocular triangle small; cibarium and genitalia as in S. cuneatum.
Male: Wing length 1.4-1.6 mm. Scutum black with 1+1 anterior cuneiform silvery
spots with length 1/5 to 1/2 of scutum length (Fig. 77 A). Gonostylus subtrapezoidal
with submedian spur with truncate apex (Fig. 77 B).
Pupa: Cocoon weave with threads obvious, slightly reinforced border and small
dorsal projection (Fig. 78 A); length at base 2.0-2.3 mm, on dorsum 1.8-2.0 mm, gill 0.9-
1.3 mm. Gills with 6 branches bifurcated close to base (Fig. 78 B). Frontoclypeus without
tubercles, thorax with scarce tubercles acuminate on dorsum; cephalic trichomes single or
bifid; thoracic trichomes with 2-4 branches.
Larva: Maximum length 3.4 mm. Cephalic apotome with some positive spots; few
evident and scarce hairs. Antenna surpassing stem apex of cephalic fan as in figure 79 D;
ratio of proximal, medial and distal segments = 1:1:1. Postgenal bridge a little shorter than
hypostomium length. Mandible without latero-mandiblular process; row of supra-marginal
setae absent. Cephalic fan with 27-34 rays. Anal sclerite without hair or scales. Anal ring with
53-58 rows of 14-18 hooks each. Body integument with trichomes single or bifid.
Distribution: Argentina: Corrientes, Jujuy, Misiones, Salta; Brazil: Amazonas,
Bahia, Goiás, Maranhão, Mato Grosso, Minas Gerais, Pará, Piauí, Rio Grande do Sul,
Rondônia, Roraima, Santa Catarina, São Paulo, Tocantins; Guyana; Venezuela: Bolivar.
Bionomics: This species breeds in rivers in areas of relatively fast-flowing water, on aquatic
vegetation. It shows intense anthropophily and is implicated as a vector of Mansonella ozzardi in
Guyana (NATHAN et al., 1982) and Onchocerca volvulus in Toototobi, Brazil (SHELLEY et al., 1982: 24).
Discussion: Simulium oyapockense, S. roraimense and S. ganalesense have gills with 6 branches
and similar scutum ornamentation, but they have a subtriangular gonostylus with a spur on the
apex. They also have abundant frontoclypeal and thoracic tubercles. The absence of
supramarginal setae on the larval mandible and subtrapezoidal gonostylus suggest placing S.
minusculum in the INCRUSTATUM group of S. (Psaroniocompsa). The wide submedian vittae,
similar to those of Cerqueirellum species, support its inclusion in S. (Cerqueirellum).

Simulium (Cerqueirellum) oyapockense FLOCH & ABONNENC


(Figs. 76 C, D; 77 C, D; 78 C-F; 79 A-C.)
Simulium oyapockense: FLOCH & ABONNENC, 1946b: 4; RAMÍREZ PÉREZ, 1983: 60; SHELLEY, 1988: 341.
Simulium (Cerqueirellum) oyapockense: PY-DANIEL, 1983: 168; COSCARÓN, 1987: 25; COSCARÓN &
COSCARÓN ARIAS, 2000a: 179-182; COSCARÓN ARIAS, 2003: 196, 204, 208, 215.
Cerqueirellum oyapockense: PY-DANIEL & MOREIRA SAMPAIO, 1995: 118.
Simulium (Psaroniocompsa) oyapockense: SHELLEY et al., 1997: 19; CROSSKEY & HOWARD, 1997: 56; 2004:
50 (cytoforms).
Simulium pseudosanguineum RAMÍREZ PÉREZ & PETERSON, 1981a: 154 =159.
Simulium sanchezi RAMÍREZ PÉREZ, YARZABAL & PETERSON, 1982: 71.
Simulium cuasisanguineum RAMÍREZ PÉREZ, YARZABAL & PETERSON, 1982: 36.
336 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Simulium pseudoamazonicum RAMÍREZ PÉREZ & PETERSON, 1981a: 151-154.


Several cytoforms and morphoforms are mentioned by CROSSKEY & HOWARD, 1997: 56.
Female: Wing length 1.6-2.1 mm. General coloration blackish, with silver pruinosity.
Frons silver, iridescent, clypeus silver pruinose. Scutum with three black vittae bordered by
silver pruinosity, submedian silver stripes showing elongated shadow comma-shaped mark
with antero-lateral illumination, or when observed laterally with antero-superior light
position (Fig. 76 C); with anterior superior illumination black median vittae narrower and
more elongated posteriorly, submedian shadow intervittae elongated anteriorly and comma-
shaped marks shorter (Fig. 76 D); with posterior illumination black vittae slightly shorter and
silver area increased, with submedian bands reaching anterior scutal margin and lacking
anterior comma-shaped marks. Scutum pilosity golden, metanotum and scutellum black.
Abdomen dark brown to black with 1+1 silver spot on tergite II; tergites III-V velvety black
with narrow postero-lateral area silver pruinose, tergite VI-VIII shiny and waxy. Frons with
subparallel sides slightly convergent ventrally, fronto-ocular triangle absent, basal portion of
cibarium with 23-27 acuminate teeth on each side, extending from borders of deep central
area to cornua; sensory vesicle of palp 1/2 or nearly 1/3 length of palpomere; claws without
subbasal tooth; paraproct short with rounded border; sternite VIII with 14-18 hairs and
gonapophysis slowly divergent below with sclerotized borders.
Male: Wing length 1.7-1.8 mm. Coloration similar to that of female. Scutum with
1+1 black, submedian, silvery cunae extending for more than half of scutum length, but
generally not joining with silvery posterior area (Figs. 77 C-D); length of cunae change
with light position. Genitalia in accordance with the morphology of subgenus. Gonocox-
ite subquadrate, about as long as high; gonostylus subtriangular with strong club-shaped
spine; ventral plate with short basal arms and rounded border.
Pupa: Cocoon length on dorsum 2.1-2.6 mm (x̄ = 2.3 mm) on base 2.5-3.2 mm (x̄ =
2.7 mm) with light yellowish brown coloration. Cocoon slipper shaped, slightly protruding
dorsally and basally, and covering gill base (Fig. 78 C). Cocoon made of soft and thick silk
with closely woven threads obscuring pupa; gill length 1.2-1.4 mm (n = 5), (x̄ = 1.3 mm).
Frontoclypeus with moderately abundant rounded platelets; facial trichomes bifid with
1+1 or 2+2 single frontal trichomes (Fig. 78 D). Respiratory gill with six filamentous
branches subdivided near base, with short basal trunk and three short primary branches
(Fig. 78 E). Disc of thorax with numerous rounded platelets, but with some pointed on
posterior area and 5+5 trichomes mainly bifid (Fig. 78 F).
Larva: Maximum length 4.1-4.7 mm ( x̄ = 4.3 mm). Color (in alcohol) yellowish white
with greenish spots (Fig. 79 A) and circular band around anterior third of abdomen; head
yellowish grey; cephalic apotome light brownish at base, with pale median longitudinal
stripe and 1+1 submedian spots, covered by abundant long and single to bifid trichomes
leaving free only the pigmented area (Fig. 79 C). Deep subcircular postgenal cleft. Ratio of
hypostomium/postgenal bridge = 2.0-2.7; hypostomial teeth at same level, with 4+4 to
5+5 lateral setae and 7-8 lateral serrations on each side. Antenna thin, with ratio of
segments: proximal, medial and distal = 1:0.9-1.1:1-1.2. Mandible with 5-6 internal teeth,
2 marginal teeth, (the second small and isolated from anterior), fringe of supramarginal
setae and presence of 2 (3) latero-mandibular processes as Fig. 79 B. Cephalic fan with 27-
38 principal rays. Thoracic proleg lateral sclerite with about 40 teeth in 7-8 groups. Body
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 337

A M
D

N
J
H

B L
O

F
Figure 78. Pupa. A-B: S. (Cerqueirellum) minusculum; A: general view; B: gill and trichomes with high
magnification. C-F: S. (C.) oyapockense; C: general view; D: frontoclypeus and trichomes; E: gill; F:
thoracic trichomes with high magnification. G-H: S. (C.) chaquense: G: general view; H: gill and
thoracic trichomes. I: S. (C.) sanguineum, portion of thorax with trichomes and base of gill. J-L: S.
(C.) cuneatum; J: general view; K: frontoclypeus and trichomes; L: gill and thoracic trichomes. M-N:
S. (C.) argentiscutum (from SHELLEY & LUNA-DIAS, 1980); M: general view; N: gills. O: S. (C.)
delponteianum, portion of thorax with trichomes and gill.
338 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

A
B
1 mm

C K

D E

L M

F G

Figure 79. Larva. A-C: S. (Cerqueirellum) oyapockense; A: lateral view; B: apical portion of mandible,
showing teeth and supra-marginal setae; C: cephalic apotome. D: S. (C.) minusculum, antenna. E-G:
S. (C.) chaquense; E: antenna; F-G: rectal papillae in dorsal and lateral views. H-M: S. (C.) cuneatum; H:
head, dorsal view; I: antenna; J: postgenal cleft and hypostomium; K: anterior border of
hypostomium; L: lateral sclerite of thoracic proleg; M: anal sclerite and hairs of the area.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 339

integument with abundant palmate trichomes 0.02 mm in length. Anal sclerite with few
hairs. Anal ring with 57-63 rows with 12-15 hooks on each. Rectal papillae with 2-3
diverticula on each lobe.
Distribution: Argentina: Salta, Santiago del Estero, Tucumán; Brazil: Amapá,
Amazonas, Mato Grosso, Minas Gerais, Pará, Paraná, Rondônia, Roraima; Colombia:
Vaupes; Ecuador: Manabí; French Guiana; Guyana; Paraguay: Maracayú; Uruguay;
Venezuela: Amazonas, Bolivar, Guarico.
Bionomics: This species breeds in plains rivers with swift water, on aquatic vegetation.
Females bite humans and are considered vectors of Onchocerca volvulus (SHELLEY 1988: 341).
Discussion: The female and male, especially with regard to the scutum ornamenta-
tion, are very difficult to separate from the other species of the S. (Cerqueirellum) =
(amazonicum) group, such as S. minusculum, S. roraimense, S. ganalesense, S. amazonicum, S.
chaquense, S. sanguineum, S. venezuelense and S. cuneatum. The pupal gills with 4-8-10 filaments
allow S. oyapockense to be separated from five species. The closest species is S. ganalesense,
which can be differentiated only by the male scutum with submedian silvery stripes joining
with the grayish posterior area, which also occurs in S. roraimense (and exceptionally, too, in
some specimens of S. oyapockense). The female is less sclerotized on the dentate border of
the cibarium. Simulium minusculum and S. roraimense can be differentiated by the short
intervittae shadow stripe, which does not surpass the scutum anterior one third. The male of
S. minusculum has a subtrapezoidal gonostylus with a subapical spur; the pupal frontoclypeus
and thorax have sparse tubercles; and the larval mandible does not have supramarginal setae.
The S. roraimense pupal gills show primary ventral branch bifurcation at greater distances
from the base than do the secondary branches. The difficulties of separating S. oyapockense
from S. roraimense and S. ganalesense based on morphological characters make it necessary to
use other methods to separate them.

Simulium (Cerqueirellum) ganalesense VARGAS, MARTÍNEZ PALACIOS & DÍAZ NÁJERA


(Figs. 76 E; 77 F.)
Simulium (Byssodon) ganalesense VARGAS et al., 1946: 114, 165; VARGAS & DÍAZ NÁJERA, 1957b: 163,
253-254; COSCARÓN, 1987: 16.
Simulium (Cerqueirellum) ganalesense: COSCARÓN & IBAÑEZ-BERNAL, 1994: 1-7.
Simulium (Psaroniocompsa) ganalesense: CROSSKEY & HOWARD, 1997: 56; 2004: 50; SHELLEY et al., 2002:
155-158.
Female: Wing length 1.6-1.9 mm. General coloration (material discolored) dark
brown; cephalic appendages grayish-brown, frons iridescent. Scutum dark brown with
golden hair; silvery longitudinal band wide anteriorly and 1+1 elongated shadow bands (Fig.
76 E). Abdomen black with median posterior subrectangular tergite II and subquadrate dark
brown tergites III-V. Frons slightly convergent; fronto-ocular triangle absent and claw
without subbasal tooth. Basal portion of cibarium with blunt teeth on lateral area. Paraproct
very short and scarcely surpassing cercus. Eighth sternite with 8-10 setae on each side.
Male: Wing length 1.6 mm. Scutum black with 1+1 silvery bands reaching posterior
grayish area (Fig. 77 F). Ratio of length/width of hind basitarsus = 6.0-6.5. Gonostylus
subtriangular with big apical spur.
340 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Pupa: Cocoon with slight antero-ventral projection and reinforced border; thin
weave with evident threads; length on base 1.6 mm, on dorsum 1.4 mm; gills 1.2 mm.
Frontoclypeus and exposed portion of thorax with rounded tubercles; frontal trichomes
single, facial trichomes bifid, thoracic trichomes with 1-3 branches. Gill with 6 branches
bifurcated relatively close to base.
Larva: Maximum length 3.5-4.5 mm. Cephalic apotome scarcely ornamented, with
abundant single and bifid trichomes. Ratio of antennal articles: proximal, medial and distal
= 1:1.1:1.2-1.3. Mandible with one latero-mandibular process. Cephalic fan with 20-22
principal rays. Postgenal cleft acuminate distally, postgenal bridge half of hypostomium
length. Hypostomium median tooth surpassing lateral teeth. Anal sclerite with 1-3 rows of
scales and scarce hairs intermixed. Anal ring with 54-68 rows with 9-13 hooks each; rectal
papillae with 3-6 diverticula on each lobe.
Distribution: Belize: Cayo; Mexico: Morelos, San Luis de Potosí, Tabasco.
Bionomics: In Belize, S. guanalense was collected in rivers about 50 m wide to flat
creeks 2 m wide, flowing over submerged grasses. In this country, females bite humans
and are very voracious. No information is available from Mexico.
Discussion: The female thoracic ornamentation is very similar to that of S.
chaquense, S. sanguineum and S. cuneatum. Different numbers of gill branches permit S.
guanalense to be separated from the species that have a greater number of branches.
Simulium oyapockense and S. roraimense, which also have six branches, can be separated in the
male by the scutum with 1+1 submedian cuneiform silvery stripes not reaching the
posterior grayish area, and in the female by the scutum with 1+1 submedian silvery stripes
narrower with shorter shadow spots; in S. roraimense the shadow intervittae spots are
shorter. Belize material shows some differences in the male scutum, pupal size and
distances of gill bifurcation (SHELLEY et al 2002: 157). The authors believe that it is
necessary to have more material and to use cytological and molecular studies to determine
if these variations are inter- or intraspecific variation.

Simulium (Cerqueirellum) roraimense NUNES DE MELLO (Fig. 77 E.)


Simulium roraimense NUNES DE MELLO, 1974: 45; RAMÍREZ PÉREZ et al., 1982: 67; SHELLEY et al. , 1982:
22; RAMÍREZ PÉREZ, 1983: 66; SHELLEY et al., 1987: 22.
Simulium (Cerqueirellum) roraimense PY-DANIEL, 1983: 171; COSCARÓN, 1987: 25; 1991: 196, 198, 199.
Cerqueirellum roraimense: PY-DANIEL & MOREIRA SAMPAIO, 1995: 118.
Simulium (Psaroniocompsa) roraimense: CROSSKEY & HOWARD, 1997: 56; 2004: 50.
Description extracted in part from RAMÍREZ PÉREZ et al., 1982; SHELLEY et al., 1982;
PY-DANIEL, 1983.
Female: Wing length 1.5-1.6 mm. Color very close to that of S. oyapockense, S.
ganalesense and S. minusculum. Scutum with 1+1 silvery submedian stripes showing, with
some light, a shadow cuneiform inter-vittae spot, shorter in S. minusculum and a little
longer than in S. amazonicum. Claw without subbasal tooth. Genitalia as in S. cuneatum.
Male: Wing length 1.2-1.5 mm. Scutum black with 1+1 silvery submedian stripes,
reaching grayish posterior area (Fig. 77 E). Scutum hair yellowish. Genitalia as in S.
cuneatum.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 341

Pupa: Cocoon with small dorsal anterior projection, and anterior border reinforced;
weave with evident threads. Length at base 2.1-2.2 mm, at dorsum 1.8 mm, gills 1.7-1.8
mm. Gill with 6 branches showing primary ventral bifurcation at greater distance from
base than for secondary branches. Frontoclypeus and exposed portion of thorax with
tubercles rounded and acuminate. Head and thoracic trichomes simple and bifid.
Larva: Maximum length 3.4-3.7 mm. Cephalic apotome with positive spots and
short trichomes with 1-3 branches. Antenna surpassing apex of cephalic fan stem; ratio of
proximal, medial and distal segments = 1:1-1.5:1-1.4. Mandible with simple latero-
mandibular process. Cephalic fan with 40-42 rays. Postgenal cleft subcirclular, postgenal
bridge half of hypostomium length. Anal ring with about 68 rows with 11-13 hooks each.
Rectal papillae with 6-9 diverticula on each lobe.
Distribution: Brazil: Roraima; Venezuela: Amazonas.
Bionomics: The species was collected in rivers on aquatic vegetation and submerged
tree branches.
Discussion: Similarities in the female scutum and pupal gill branches with S.
oyapockense and S. minusculum can produce misidentifications, but the silvery submedian
stripes of the male scutum in the mentioned species do not reach the grayish posterior
area, and bifurcation of the pupal primary ventral branch is lower, not drawn away from
the bifurcations of the secondary branches. The closest species is S. ganalesense; only small
differences in length of the intervittae submedian shadow spots and proportionately
shorter pupal gill permit its separation. Here, also cytological or molecular studies will be
necessary to assure the differentiation of these species.

Simulium (Cerqueirellum) amazonicum GOELDI (Fig. 76 F.)


Simulium amazonicum GOELDI, 1905: 138; SHELLEY, 1982: 67; 1988: 203; SHELLEY et al., 1982: 8-21.
Simulium (Cerqueirellum) amazonicum: PY-DANIEL, 1983: 167.
Simulium (Psaroniocompsa) amazonicum: CROSSKEY & HOWARD, 1997: 56; 2004: 50.
Simulium tallaferroae RAMÍREZ PÉREZ, 1971: 339.
Cerqueirellum amazonicum PY-DANIEL & MOREIRA SAMPAIO, 1995: 118.
Description from SHELLEY et al., 1982 and PY-DANIEL, 1983 gives details.
Female: Wing length 1.2-1.9 mm. Frons and clypeus black with bluish silver pruinosity.
Scutum with anterior illumination showing black with silver gray pruinosity, leaving median
black and 1+1 sublateral bands, and 1+1 diffuse shadow-like spots on anterior border (Fig.
76 F); with posterior light, the median band is wider posteriorly and shorter sublaterally;
scutum with silver-yellow appressed hair. Abdomen with tergite I brownish black, tergites II-
V velvety black, anterior and posterior margins silver pruinose, tergites VI-IX shiny black.
Eighth sternite with about 12 setae on each side. Claws without subbasal tooth.
Male: Coloration generally as in female. Scutum black bordered with silver
pruinosity that extends as 1+1 wide triangular wedge submedially, similar to that of S.
argentiscutum, but not reaching half scutum length. Genitalia as in S. cuneatum.
Pupa: Cocoon with reinforced border and small dorsal projection. Length at base 2.0-2.7
mm; dorsally 1.6-2.3 mm; gills 1.0-1.5 mm. Gill with 8 filaments, with branching relatively close
to base, arrangement similar to that of S. chaquense but with some variations as shown by
342 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

SHELLEY et al. (1982). Frontoclypeus and thorax with small numerous acuminate tubercles;
trichomes bifid except frontals that can be single, those on thorax with 3 or 4 branches.
Larva: Maximum length 3.6-4.3 mm. Color yellowish to brown with gray mottling.
Cephalic apotome scarcely ornamented, with positive spots and with numerous hairs from
thin and single to thicker with 2-4 branches. Antenna surpassing stem apex of cephalic
fan. Ratio of antennal segments: proximal, medial and distal = 1:0.9-1: 1-1.1. Postgenal
cleft large with rounded margin; postgenal cleft 1/3 or shorter than hypostomium length.
Hypostomium with 3-4 lateral setae on each side. Mandible with 1 or 2 marginal teeth;
latero-mandibular process absent. Cephalic fan with 12-15 rays. Thoracic proleg lateral
sclerite with about 15-17 teeth arranged in about 8 process. Body integument with spines.
Ventral papillae obsolescent. Rectal papillae with about 8 diverticula on each lobe. Anal
ring with 52-58 rows with 9-12 hooks each.
Distribution: Brazil: Acre, Amazonas, Roraima; Colombia: Amazonas; Venezuela:
Amazonas, Apure, Barinas, Guarico.
Bionomics: Immature stages live in large rivers on herbaceous vegetation and
submerged bushes at places with fast current up to 15 m deep. This species is a very
dangerous pest, attacking humans, horses, cows and sheep in great quantity. It is
mentioned as a vector of Mansonella ozzardi and Onchocerca volvulus.
Discussion: The closest species are those with 8 filamentous branches in the pupa,
such as S. chaquense and sanguineum. Simulium sanguineum can be differentiated by the
branching of the gill (Fig. 78 I), and in S. amazonicum, as for S. chaquense, in the dorsal
branch (Fig. 78 H). Simulium chaquense can be differentiated in the pupa by the large size
and presence of small spines on abdominal tergites III-IV; in the female by the 1+1
anterolateral vittae more elongated and in the male by the 1+1 silvery submedian stripe
reaching the posterior grayish area. Based on the original description of S. tallaferroae, the
synonymy proposed by SHELLEY et al. (1982: 8) is accepted.

Simulium (Cerqueirellum) chaquense COSCARÓN (Figs. 76 G-K; 77 G-J; 78 G, H; 79 E-G.)


Simulium chaquense COSCARÓN, 1971: 33-41; SHELLEY et al., 1980: 785; SHELLEY et al., 1982: 8; PY-
DANIEL, 1983: 170; COSCARÓN, 1983: 81.
Simulium (Cerqueirellum) chaquense: PY-DANIEL, 1981: 170; COSCARÓN, 1987: 24; 1991: 201.
Simulium (Psaroniocompsa) chaquense: CROSSKEY & HOWARD, 1997: 56; 2004: 50.
As in other species of this group, large differences occur in the scutum ornamentation.
Female: Wing length 2.5 mm. Scutum black with 1+1 silvery submedian bands
reaching posterior grayish area, with longitudinal shadow elongated nacreous band over
silvery bands, showing variation in vittae shape, in accordance with illumination (Figs. 76
H, I). Abdomen blackish with pattern as in figure 76 K. Fronto-ocular triangle absent (Fig.
76 G); legs with single and spatuliform hairs (Fig. 76 J). Genitalia as in S. cuneatum.
Male: Wing length 2.2-2.3 mm. Scutum ornamentation very similar to that of
female, with 1+1 silvery submedian bands reaching posterior grayish area (Fig. 77 G), but
in some specimens not reaching posterior area, as in S. cuneatum. Abdomen coloration as
in figures 77 H, I. Gonostylus about as long as gonocoxite, subtriangular in shape with
blunt apex and 1-3 apical spurs (Fig. 77 J). Other genital pieces as in S. cuneatum.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 343

Pupa: Cocoon with very short ventral projection and reinforced anterior border
(Fig. 78 G); length at base 3.0-3.5 mm, dorsally 2.5-2.7 mm, gill 1.8-2.0 mm. Gill with 8
branches with branching near base, showing bifurcation in both secondary dorsal
branches (Fig. 78 H); this bifurcation can be at greater distances from base. Frontoclypeus
and thorax with rounded to acuminate tubercles, especially over median longitudinal area
of thorax in moderate to abundant quantity; trichomes from single to three branched.
Larva: Maximum length 4.5-5.0 mm. Cephalic apotome yellowish brown, scarcely
ornamented, with few evident spots and with hairs of 1-3 branches. Postgenal cleft deep,
subcirclular anteriorly; postgenal bridge about half of hypostomium length. Antenna
surpassing apex of cephalic fan, ratio of proximal, media and distal segments = 1:1:1.1
(Fig. 79 E). Mandible with marginal tooth and one latero-mandibular process (only visible
on the most southern specimen); cephalic fan with 36-40 ray; anal ring with about 62-66
rows of 11-12 hooks each. Rectal papillae with 4-6 diverticula on each lobe (Figs. 79 F, G).
Anal sclerite with scarce scales and single hair.
Distribution: Argentina: Buenos Aires, Chaco, Corrientes, Formosa, Salta, Santa
Fe, Santiago del Estero, Tucumán.
Bionomics: This species breed in small creeks with low to regular flow, but also in
medium (Salado River) to exceptionally large rivers (Paraná River) over grass or
submerged tree branches (also found on stones, plastic objects, etc.). In Paraná River, it
was found in small numbers only two times during one year of collections, in a place
where S. cuneatum is very abundant. In Salado River in the center of Buenos Aires
province it was very abundant during high river levels, which produce overflows; it
occurred in small currents at the sides of the Salado River. As it is an anthropophilic
species, it is a serious pest in several cities and towns such as Junin along the Salado River
where it also attacks cows and horses.
Discussion: The closest species are S. amazonicum and S. sanguineum, with 8 branches and
basal branching. Simulium sanguineum can be separated by the secondary branches bifurcating on
the ventral branch; in S. amazonicum and S. chaquense differentiation is on the dorsal branch.
Small differences are found in the pupa of both species: S. amazonicum is small and the few
tubercles on the head and thorax are acuminate; in S. chaquense, these are only conical on the
median area of the thorax. Simulium. amazonicum has a male scutum with 1+1 cuneiform silvery
bands not surpassing the median length of the scutum, and in S. chaquense, these reach the
posterior grayish area. Simulium amazonicum females have less expanded shadows in the
intervittae spots. The distinct environment where they breed is another difference of both
species. Non morphological methods probably can help to resolve this identification problem,
very common in Neotropical black flies. Simulium chaquense female coloration is similar to that
of S. cuneatum, but the pupal gill with 10 branches allows their separation.

Simulium (Cerqueirellum) sanguineum KNAB (Figs. 76 L, M; 77 K; 78 I.)


Simulium sanguineum KNAB, 1915c: 279; TIDWELL et al., 1981: 79; SHELLEY et al., 1982: 26; SHELLEY,
1988: 207.
Simulium (Cerqueirellum) sanguineum: PY-DANIEL, 1983: 168; COSCARÓN, 1991: 196.
Simulium (Psaroniocompsa) sanguineum: CROSSKEY & HOWARD, 1997: 56; 2004: 50.
344 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

This species was described in detail by TIDWELL et al. (1981), and PY-DANIEL (1983)
added more information on larval stages. Diagnosis was extracted from these authors.
Female: Wing length 1.3.0-1.5 mm. Scutum with 1+1 submedian bluish-silver to
golden-silver iridescent stripes reaching posterior grayish border and varying in shape in
accordance with light position, intervittae shadow spots elongated (Figs. 76 L, M). Claws
without subbasal tooth. Genitalia as in S. cuneatum.
Male: Wing length 1.4-1.5 mm. Scutum black with 1+1 silvery cuneiform anterior
spots projected by thin stripe reaching nearly posterior grayish area and sometimes
meeting it (Fig. 77 K). Genitalia as in S. cuneatum. Gonostylus subtriangular with single
spur as in S. amazonicum.
Pupa: Cocoon with small dorsal anterior projection. Cocoon length 1.7-2.5 mm; gills
1.4-1.8 mm. Gills with 8 branches; branching close to base; secondary ventral branches
bifurcated (Fig. 78 I). Frontoclypeus and thorax with rounded tubercles in regular quantity
and showing irregular pattern; frontal trichomes single, facial trichomes bifid, thoracic
trichomes with 2 to 4 branches.
Larva: Maximum length 3.8-4.5 mm. Antenna surpassing apex of cephalic fan stem;
ratio of proximal, medial and distal segments = 1:1:1. Mandible with 2 long, sinuous or
straight latero-mandibular processes. Anal ring with 52-55 rows of 8-12 hooks each.
Rectal papillae with 3-5 diverticula on each lobe.
Distribution: Colombia: Cauca, Choco; Panama: Darien.
Bionomics: Immature stages were collected in rivers with turbid water on sub-
merged branches about 0.5-3.0 m deep. The region is a tropical rain forest, below 200 m in
altitude, but the immatures have also been found at 1000 m in clear, fast-flowing streams.
This species is anthropophilic, and is implicated as a vector of Mansonella ozzardi.
Discussion: The closest species are S. amazonicum and S. chaquense. The bifurcation
of the secondary dorsal branches in the latter two species permits them to be separated
from S. sanguineum.

Simulium (Cerqueirellum) cuneatum (ENDERLEIN) (Figs. 76 N-T; 77 L-P; 78 J-L; 79 H-M.)


Psaroniocompsa cuneata ENDERLEIN, 1936: 124.
Simulium cuneatum VARGAS, 1945a: 129; VULCANO, 1967: 9; COSCARÓN, 1983: 90-95.
Simulium (Cerqueirellum) cuneatum: PY-DANIEL, 1983: 181; COSCARÓN, 1987: 24; 1991: 202; COSCARÓN
& COSCARÓN ARIAS, 2000: 25.
Simulium (Psaroniocompsa) cuneatum CROSSKEY & HOWARD, 1997: 57; 2004: 52.
Female: Wing length 1.7-2.1 mm. Coloration similar to that of S. chaquense,
complemented with small variations in relation to the illumination direction (Figs. 76 O-
R). Abdomen as in S. chaquense but lateral silver gray spots on segment III-V are less
evident. Frons relatively wide. Basal portion of cibarium with 2-3 rows of acute teeth (Fig.
76 N); claws without subbasal tooth. Eighth sternite with about 40 setae, gonapophysis
with abundant microtrichiae; genital fork with posterior branches relatively wide,
spermatheca subovoidal (Fig. 76 S); paraprocts relatively short (Fig. 76 T).
Male: Wing length 1.7-2.0 mm. Scutum black with 1+1 silvery submedian cuneiform
spots reaching or surpassing half of scutum length, but not joining with posterior grayish
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 345

area (Figs. 77 L, M). Scutum hair golden to silvery, homogeneously distributed. Ratio of
length/width of hind basitarsus = 5.7. Genitalia as in figure 77 N; gonostylus subtriangu-
lar with one apical spur (Fig. 77 O); ventral plate with slightly curved terminal border (Fig.
77 P); endoparameres with strong hooks.
Pupa: Cocoon with small anterior dorsal projection (Fig. 78 J), weave compact with
evident threads. Length on base 2.0-2.3 mm, at dorsum 2.5-2.7 mm; gills 1.2-1.8 mm.
Frontoclypeus and exposed portion of thorax with scarce and apically acute tubercles
(Fig. 78 K). Head trichomes single, thoracic trichomes mostly single but can be bifid. Gill
with 10 branches, with branching close to base, showing primary dorsal and median 4
branches each (Fig. 78 L).
Larva: Maximum length 3.5-4.0 mm. Cephalic apotome with scarce ornamentation
(Fig. 79 H) and numerous short single hairs. Antennae surpassing apex of head fan stem;
shape as in figure 79 I; ratio of proximal, medial and distal segments = 1:1.1:1.2 (Fig. 79 I).
Mandible without latero-mandibular process. Cephalic fan with 16-22 rays. Postgenal cleft
subcircular (Fig. 79 J); postgenal bridge about half of hypostomium length. Hypostomi-
um with central tooth lower than lateral teeth (Fig. 79 K). Lateral sclerite of thoracic
proleg with teeth in 3-5 groups with 2-5 teeth each (Fig. 79 L). Anal sclerite with wide and
short ventral branches with scarce single hairs (Fig. 79 M). Anal ring with 60-70 rows with
10-13 hooks each. Body integument with numerous single short hairs. Rectal papillae with
about 6 elongate diverticula on each lobe.
Distribution: Argentina: Chaco, Corrientes, Entre Ríos, Formosa, Misiones, Santa
Fe; Paraguay: Cordillera.
Bionomics: The species breeds in large rivers such as Paraná, and Paraguay, in fast-
flowing places, over leaves of tree branches, particularly of Salix humboldtiana. It is very
abundant during the dry season (August-October; COSCARÓN & COSCARÓN ARIAS, 2000).
Simulium cuneatum is an anthropophilic species and very dangerous near the river areas as in
Corrientes in the spring time.
Discussion: Simulium cuneatum is very close to S. chaquense and S. minusculum, which
sometimes are sympatric and also the other species of this group or subgenus. It can be
separated by the number of pupal gill branches, which in other species are of 4-8 branches.

Simulium (Cerqueirellum) venezuelense RAMÍREZ PÉREZ & PETERSON


Simulium venezuelense RAMÍREZ PÉREZ & PETERSON, 1981: 157; PY-DANIEL, 1983: 180; RAMÍREZ PÉREZ,
1983: 123.
Simulium (Psaroniocompsa) venezuelense: CROSSKEY & HOWARD, l997: 56; 2004: 50.
(Diagnosis based on RAMÍREZ PÉREZ & PETERSON, 1981).
Female: Wing length 1.6-1.7 mm. Blackish species with scutum and abdomen
ornamentation as in S. minusculum. Cibarium, fronto-ocular triangle, legs, claws and
paraprocts similar to those of S. cuneatum.
Male: Wing length 1.5-1.6 mm. Scutum black with 1+1 silvery short and wide
anterior cuneiform spots not reaching to half of scutum length (probably with another
illumination direction, they get longer). Genitalia as in S. cuneatum.
346 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Pupa: Cocoon slipper shaped with anterior reinforcement; gill length 1.7 mm. Gill
with 4 branches, bifurcated close to base, with primary ventral branch about 3 times
longer than dorsal branch. Frontoclypeus and thorax without platelets. Cephalic tri-
chomes single, thoracic trichomes with 2-3 branches.
Larva: Unknown.
Distribution: Venezuela: Bolivar; Brazil: Amapá, Tocantins.
Bionomics: Unknown.
Discussion: This is a peculiar species, with adults similar to the known species of
this group, but with a pupal gill of only 4 branches, which permits it to be readily
differentiated.

Simulium (Cerqueirellum) argentiscutum SHELLEY & LUNA DIAS (Figs. 76 U, V; 77 Q; 78 M.)


Simulium argentiscutum SHELLEY & LUNA-DIAS, 1980: 105; HAMADA & ADLER, 2001: 119.
Simulium (Cerqueirellum) argentiscutum: PY-DANIEL, 1982: 174.
Simulium (Psaroniocompsa) argentiscutum: CROSSKEY & HOWARD, 1997: 57; 2004: 52.
Cerqueriellum argentiscutum PY-DANIEL & MOREIRA SAMPAIO, 1995: 118.
The species was well described by the authors of the species name, and PY-DANIEL
(1982) added more information. The information for this diagnosis mostly was extracted
from the above-mentioned authors.
Female: Wing length 1.8-2.2 mm. Scutum black with intense silver-gray pruinosity,
leaving black narrow median and 1+1 sublateral bands, and 1+1 intervittae darkened
cuneiform shadow spots, elongated with frontal light and shorter with posterior
illumination (Figs. 76 U-V). Abdomen very similar to that of S. amazonicum. Claw without
subbasal tooth. Genitalia as in S. cuneatum.
Male: Wing length 1.9-2.0 mm. Scutum color and ornamentation very similar to that
of S. amazonicum, but 1+1 silvery submedian cuneiform bands surpassing half of scutum
length (Fig. 77 Q). Scutum hair yellowish. Genitalia as in S. cuneatum.
Pupa: Cocoon shoe shaped with gelatinous consistency, without evident threads
(Fig. 78 M). Length at base 2.8-3.4 mm, on dorsum 1.9-2.2 mm, gills 0.8-0.9 mm. Gill with
6 branches bifurcated close to base, but ventral primary branch bifurcated at greater
distance, about 4-5 times its length from base (Fig. 78 N); filaments with winkled margins.
Larva: Maximum length 4.4-5.3 mm. Cephalic apotome with positive spots and
single or bifid trichomes. Antenna reaching apex of cephalic fan stem; ratio of proximal,
medial and distal segments = 1:1.1-1.3:1.3-1.5. Mandible without latero-mandibular tooth;
well-developed marginal teeth and presence of supramarginal setae. Postgenal cleft very
deep, reaching near hypostomium base, interrupting postgenal cleft centrally. Cephalic fan
with 25-28 rays. Ninth segment without ventral tubercles. Anal ring with 52-65 rows of 8-
16 hooks each. Rectal papillae with 5-8 diverticula on each lobe.
Distribution: Brazil: Acre, Amazonas, Bahia, Rondônia; Colombia: Amazonas; Peru.
Bionomics: The species breeds in large, fast-flowing rivers. It bites humans and is
considered a vector of Mansonella ozzardi (SHELLEY, 1980).
Discussion: Simulium argentiscutum differs from similar species in the female by the
extensive silvery gray pruinosity of the scutum, the narrow median and 1+1 sublateral
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 347

black vittae, and very enlarged intervittae shadow spots, with reddish brown to black
tones in some specimens when illumination changes. The cocoon is shoe shaped and has
a gelatinous consistency. The larva has a deeper postgenal cleft, presence of multi-
branched scales on the body integument and absence of ventral papillae.

Simulium (Cerqueirellum) delponteianum WYGODZINSKY (Figs. 76 W-Z; 77 R-T; 78 O.)


Simulium delponteianum WYGODZINSKY, 1961: 30; COSCARÓN, 1983: 95; PY-DANIEL, 1983: 181; SHELLEY
et al., 1980: 785; SHELLEY et al., 1982: 4.
Simulium (Cerqueirellum) delponteianum: COSCARÓN, 1987: 24; 1991: 203.
Simulium (Psaroniocompsa) delponteianum: CROSSKEY & HOWARD, 1997: 57; 2004: 52.
Cerqueirellum delponteianum: STRIEDER, 2004: 125.
Female: Wing length 2.5-2.8 mm. Frons blackish with gray pruinosity. Mesonotum
blackish gray-pruinose with hair grayish to tin colored; scutum with silvery light antero-
lateral angles and 1+1 submedian, subovoidal spots projected posteriorly as gray-matte
stripes each joining posterior grayish area (Fig. 76 Y). Scutellum, metanotum and pleurae
black, the latter two with silver pruinosity. Legs yellowish gray, with darker portion
blackish. Abdomen black with tergites I-V velvety and VI-IX with cereous sheen, segment
II with 1+1 silvery spots. Fronto-ocular triangle rather deep (Fig. 76 W). Basal portion of
cibarium with teeth shorter than in other species of this group, and median thicker
invaginated area (Fig. 76 X). Claw with or without small subbasal tooth (Fig. 76 Z).
Genitalia as in S. cuneatum.
Male: Wing length 2.2-2.7 mm. Scutum silvery black with 1+1 anterior submedian
cuneiform silvery spots, which with changing light direction, are continued as thin grayish
stripes surpassing half of scutum length (Figs. 77 R-S). Abdomen blackish with 1+1
silvery spots on segments II-VIII. Gonostylus slightly longer than half of gonocoxite
length, triangular with blunt apex and 5-10 spurs on dorsal distal area (Fig. 77 T). Ventral
plate as in S. cuneatum.
Pupa: Cocoon with straight and reinforced border, with weave thin and homoge-
neous and evident threads; length at base 3.6-4.6 mm; on dorsum 3.0-4.0 mm; gills 1.8-2.5
mm. Frontoclypeus and exposed portion of thorax with medially abundant rounded
tubercles; cephalic and thoracic trichomes single and well developed. Gills with 6
branches with branching near base (Fig. 78 O).
Larva: Maximum length 5.0 mm. Cephalic apotome with few, weakly evident
positive spots, and slightly darkened at base, hair of head dorsally with 1-3 branches.
Antenna as long as or scarcely surpassing apex of cephalic fan stem; ratio of proximal,
medial and distal segments = 1:1.1:1.3. Mandible with one latero-mandibular tooth; supra-
marginal setae absent. Cephalic fan with 28-32 rays. Postgenal cleft subcircular, postgenal
bridge about half of hypostomium length. Anal ring with 70-78 rows, with 14-18 hooks
each; hair of anal sclerite area scarce and single. Rectal papillae with 3-5 diverticula on
each lobe. Body integument with scarce and short single hairs.
Distribution: Argentina: Buenos Aires, Corrientes, Entre Ríos; Brazil: Rio Grande
do Sul; Uruguay: Artigas.
348 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Bionomics: The species breed in creeks of plains, with little current in places with
slight gradient over submerged aquatic vegetation. Females can bite humans but they are
not aggressive; they also attack cattle.
Discussion: Simulium delponteianum has similarities with the S. amazonicum group, but
can be separated by the grayish thorax, small size of the cibarium teeth, male gonostylus
with large number of spurs, and larval mandible without supramarginal setae. These
differences make it difficult to include it in this supraspecific taxon. Simulium (Cerqueirel-
lum) argentiscutum has scutum ornamentation and cocoon shape and texture different from
other species. Simulium minusculum has the gonostylus shape and absence of mandibular
supramarginal setae not concordant with the other species of the group. Simulium chaquense
has 2 or 3 spurs on the gonostylus. All of these species show variations from the typical
subgeneric characters.

Species inquirendae

Simulium quadristrigatum ENDERLEIN


Simulium quadristrigatum ENDERLEIN, 1934b: 285-286; VARGAS, 1945a: 189; VULCANO, 1967: 19;
COSCARÓN, 1983: 96; 1987: 25.
Psaroniocompsa quadristrigata: ENDERLEIN, 1936: 123.
Simulium (Psaroniocompsa) quadristrigatum: CROSSKEY & HOWARD, 1997: 58; 2004: 52; HERNÁNDEZ &
SHELLEY, 2005: 9-10.
This species is not easy to identify. The type is a female in the Berlin Museum. It has a
type label and “Paraná”. It is not well preserved; it is pinned through the scutum, and is
attacked by fungus and has lost its claws. The original description does not help much.
Apparently, it is close to S. delponteianum in the gray scutum tones and leg coloration and wing
length, but the cibarium (HERNANDEZ & SHELLEY, 2005; Fig. 42) is like that of S. chaquense
and S. cuneatum. It would be very useful to know the shape of the claws, as well as the other
life stages, especially the larva to best define the species identity. The type locality (Paraná)
does not identify clearly the place it was collected, which can refer to the Brazilian state or to
the Paraná River that crosses several states in southern Brazil, Paraguay and Argentina.
ENDERLEIN (1937: 13) mentions another two females, from Paraguay but without adequately
specifying the locality. We prefer to maintain S. quadristrigatum as species inquirendae.

Simulium lutzi KNAB


Simulium lutzi KNAB, 1913: 155 (n. nom. to Simulium minutum SURCOUF & GONZÁLEZ RINCONES);
SHELLEY et al., 1982: 27; COSCARÓN, 1987: 25.
Simulium exiguum: LUTZ, 1909: 141 (preoc.).
Simulium minutum SURCOUF & GONZÁLEZ RINCONES, 1911: 290 (repl. name preoc. for exiguum).
SHELLEY et al. (1982) explained, in detail, the history of S. lutzi and the material
studied by LUTZ (determined as S. exiguum) from Rio Grande at the limit of São Paulo and
Minas Gerais (Brazil). As this material is not well preserved and does not permit proper
determination, we prefer to maintain S. lutzi as species inquirendae.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 349

Subgenus Simulium (Coscaroniellum) PY-DANIEL (Figs. 80-83; 144.)


Type-species: Simulium quadrifidum LUTZ, 1917: 66.
Female: Small species, maximum wing length 2.5 mm. Coloration generally blackish
to reddish dark brown. Scutum with 1+1 silvery submedian bands that can reach posterior
grayish area or only on anterior half as cuneiform spots. Abdomen similar to that of S.
(Cerqueirellum). Frons wider than high, with infrafrontal sutures well developed. Legs with
filiform and petaloid hair; claw with small subbasal tooth. Basal portion of cibarium with
median invaginated area well sclerotized, with small blunt projections at sides and teeth
mostly apically blunt. Paraprocts short; genital fork stem distally wider. Gonapophysis
distally surpassing eighth sternite; internal border slightly concave and well sclerotized;
surface with abundant spiculae. Spermatheca relatively large.
Male: Coloration blackish to reddish dark brown with 1+1 anterior submedian
cuneiform silvery spots. Abdomen similar to that of S. (Cerqueirellum). Gonostylus about
as long as gonocoxite, subtrapezoidal, longer than wide, with large spur distally and
slightly flattened; ventral plate and endoparameres as in S. (Cerqueirellum).
Pupa: Cocoon compact with threads showing in some species, poorly defined weave
with spongy aspect. Gill with 4-8 branches. Frontoclypeus and thorax mostly with apically
acute tubercles, and trichomes with 2-3 branches.
Larva: Maximum length 4.0 mm. Body generally with dark band on first abdominal
segment; integument with elongate multibranched trichomes. Cephalic apotome with
median and lateral spots generally weak. Antenna surpassing apex of cephalic fan stem,
with distal article proportionately long. Postgenal cleft deep, with postgenal bridge shorter
than hypostomium length. Mandibles with latero-mandibular process and without
supramarginal setae. Rectal papillae with diverticula. Ninth abdominal segment ventrally
with 1 pair of tubercles.
Distribution: This taxon is a heterogeneous group of exclusively Amazonian or
periamazonian species, composed of S. quadrifidum, S. cerradense, S. cauchense and, with
some restrictions, S. goeldii, S. ulyssesi, S. daltanhani and S. quadrivittatum (Fig. 144).

KEYS TO SIMULIUM (COSCARONIELLUM) SPECIES

Females
1 Scutum black with 1+1 silvery submedian stripes reaching or not reaching silvery
posterior area (Figs. 80 E, Q, R, W) .............................................................................. 2
– Scutum reddish brown to blackish without 1+1 silvery submedian stripes or only
1+1 anterior rectangular thin silvery spots ................................................................... 5
2(1) Scutum with 1+1 anterior submedian comma-shaped silvery stripes not reaching
half scutum length ................................................................................ Simulium cauchense
– Scutum with 1+1 submedian silvery stripes reaching silvery posterior area ......... 3
3(2) Scutum with anteriorly wide silvery submedian stripes, and posteriorly wide drop-
shaped median black stripe (Fig. 80 E); 1+1 cibarium submedian protuberances
blunt (Fig. 80 D) ............................................................................... Simulium quadrifidum
350 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

– Scutum with submedian silvery stripes not as wide anteriorly and median black
stripe not as wide posteriorly, 1+1 cibarium submedian protuberances acute (Figs.
80 P-Y) ................................................................................................................................. 4
4(3) Black median scutum stripes wider posteromedially, and with median thin silvery
stripe (Fig. 80 Q) .................................................................................. Simulium cerradense
– Black median scutum stripes not wider posteromedially, and without median
silvery stripe (Fig. 80 W) ............................................................ Simulium quadrivtittatum
5(1) Scutum black with 1+1 anterior submedian thin rectangular silver spots
connected with silver lateral stripe; wing length 2.0 mm; ratio of palpomere
length/sensory vesicle = 1.4 .............................................................Simulium daltanhani
– Scutum reddish without anterior silver spots (Fig. 80 S); wing length 1.8; ratio of
palpomere length/sensory vesicle = 2.3-3.1 .......... Simulium goeldii, Simulium ulyssesi

Males
1 Scutum with 1+1 anterior submedian stripes (Fig. 81 A) ......................................... 2
– Scutum without anterior submedian stripes or only small and thin rectangular
anterior spot (Fig. 81 D) ................................................................................................... 4
2(1) Scutum with 1+1 submedian silvery stripes joining with posterior grayish area
(Fig. 81 K); gonostylus about 1.5 longer than wide at base (Fig. 81 L) ...................
......................................................................................................... Simulium quadrivtittatum
– Scutum with 1+1 submedian silvery stripes not joining with posterior grayish area
(Fig. 81 A); gonostylus about 2 times longer than wide at base (Fig. 81 C) ......... 3
3(2) Scutum with 1+1 anterior submedian thin stripes reaching about 1/2 scutum
length (Fig. 81 A) ............................................................................. Simulium quadrifidum
– Scutum with 1+1 anterior submedian cuneiform stripes reduced to anterior 1/4
of scutum length ................................................................................... Simulium cauchense
4(1) With some illumination, silvery lateral band of scutum joining anteriorly to 1+1
submedian silvery spots (Fig. 81 D); gonostylus with 1 spur; gonostylus length/
width ratio at base = 2.5 ..................................................................... Simulium cerradense
– With no illumination, silvery lateral band of scutum joining 1+1 submedian
silvery spots; gonostylus with 1-3 spurs; gonostylus length/width ratio at base =
1.8-2.1 ................................................................................................................................... 5
5(4) Gonostylus with 2 or 3 spurs (Fig. 81 J) ........................................ Simulium daltanhani
– Gonostylus with 1 spur (Fig. 81 H) ......................... Simulium goeldii, Simulium ulyssesi

Pupae
1 Gill with 4 branches .......................................................................................................... 2
– Gill with 6-8 branches ....................................................................................................... 3
2(1) Gill branches longer than cocoon base (Fig. 82 A); length 3.3-3.4 mm and 2.8 mm
respectively; secondary gill branches emerging close to primary gill branches (Fig. 82
C); abdominal tergite V with denticles on anterior border .......... Simulium quadrifidum
– Gill branches and cocoon base length about same size (length 2.2-2.6 and 2.0-2.6
respectively); secondary gill branches emerging distal to 1/2 gill length; abdominal
tergite V without denticles on anterior border ............................... Simulium cauchense
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 351

3(1) Gill with 6 branches .......................................................................................................... 4


– Gill with 8 branches .......................................................................................................... 6
4(2) Basal trunk of gill elongated and bifurcation of secondary branches relatively far
from base (Fig. 82 K) .............................................................................. Simulium ulyssesi
– Basal trunk of gill short and bifurcation of secondary branches near base (Figs. 82
F, L) ....................................................................................................................................... 5
5(4) Cocoon weave fibrous, with obvious threads; cocoon with anterodorsal projection
(Fig. 82 L); thoracic tubercles rounded apically; abdominal tergite V with denticles
on anterior border ...............................................................................Simulium daltanhani
– Cocoon weave spongy, without obvious threads; cocoon without anterodorsal
projection (Fig. 82 E); thoracic tubercles acuminate apically; abdominal tergite V
without denticles on anterior border ............................................... Simulium cerradense
6(3) Cocoon expanded laterally and with open loops (Fig. 82 M); bifurcation of
tertiary branches relatively close to base ................................. Simulium quadrivittatum
– Cocoon not expanded laterally and with compact weave (Fig. 82 G); bifurcation
of tertiary branches relatively far from base ........................................ Simulium goeldii

Larvae
1 Subesophageal ganglion pigmented; body length 4.8-5.5 mm; postgenal cleft
acuminate apically ...............................................................................Simulium daltanhani
– Subesophageal ganglion not pigmented; body length 3.1-4.7 mm; postgenal cleft
apically rounded or not (Figs. 83 D, S) .......................................................................... 2
2(1) Anal ring with 80-84 rows of hooks; ratio of postgenal bridge length/
hypostomium height = 3.9; ratio of proximal, medial and distal antennal articles =
1:1.0:1.0 (Fig. 83 N) ............................................................................. Simulium cerradense
– Anal ring with 57-66 rows of hooks; ratio of postgenal bridge length/
hypostomium height = 1.4-2.9; ratio of proximal, medial and distal antennal
articles = 1:0.3-0.9:1.1-1.8 ................................................................................................ 3
3(2) Rectal papillae with 10-13 diverticula on each lobe; latero-mandibular process
curved and thin (Fig. 83 C); ratio of hypostomium length/postgenal bridge length
= 2.0-2.9 ................................................... Simulium quadrifidum, Simulium quadrivittatum
– Rectal papillae with 4-7 diverticula on each lobe; latero-mandibular process
straight or sinuous, relatively thick; ratio of hypostomium length/postgenal
bridge length = 1.4-2.1 ..................................................................................................... 4
4(3) Latero-mandibular process sinuous, thin and simple apically; proportion of length
of proximal, medial and distal antennal articles = 1:1.5:1.6; ratio of hypostomium
length/postgenal bridge length = 1.9-2.1 ........................................ Simulium cauchense
– Latero-mandibular process straight, thick and apically bifid or trifid; proportion
of length of proximal, medial and distal antennal articles = 1:1.1:1.4; ratio of
hypostomium length/postgenal bridge length = 1.4-1.6 ........................................... 5
5(4) Latero-mandibular process apically bifid; cephalic fan rays 38-47; body length =
3.6-3.8 mm ................................................................................................... Simulium goeldii
– Latero-mandibular process apically trifid; cephalic fan rays 30-36; body length =
4.1-4.2 mm ................................................................................................. Simulium ulyssesi
352 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

CLAVES DE ESPECIES DE SIMULIUM (COSCARONIELLUM)


Hembras
1 Escudo negro con 1+1 bandas plateadas submedianas llegando o no al área
plateada posterior (Figs. 80 E, Q, R, W) ....................................................................... 2
– Escudo castaño rojizo a negruzco sin 1+1 bandas submedianas plateadas o solamente
reducidas a 1+1 manchas plateadas anteriores rectangulares finas ............................... 5
2(1) Escudo con 1+1 bandas plateadas submedianas en forma de coma no llegando a la
mitad del largo del escudo ................................................................... Simulium cauchense
– Escudo con 1+1 bandas plateadas submedianas llegando al área plateada posterior ....... 3
3(2) Escudo con bandas submedianas plateadas más anchas anteriormente, y banda
mediana negra más ancha posteriormente en forma de gota (Fig. 80 E); cibario con
1+1 protuberancia submedianas romas (Fig. 80 D) ....................... Simulium quadifidum
– Escudo con bandas submedianas plateadas no tan anchas anteriormente y banda
mediana negra no tan ancha posteriormente; cibario con 1+1 protuberancias
submedianas aguzadas (Figs. 80 P, Y) ............................................................................ 4
4(3) Banda negra mediana del escudo más ancha medio-posteriormente y con una
banda mediana plateada fina (Fig. 80 Q) ........................................ Simulium cerradense
– Banda negra mediana del escudo no más ancha medio-posteriormente y sin una
banda mediana plateada (Fig. 80 W) ........................................ Simulium quadrivittatum
5(1) Escudo negro con 1+1 mancha plateada anterior submediana rectangular
conectada con banda lateral plateada; longitud ala 2,0 mm; relación del largo del
palpómero/ vesícula sensorial = 1,4 ...............................................Simulium daltanhani
– Escudo castaño rojizo sin manchas anteriores plateadas (Fig. 80 S); longitud ala
1,8 mm; relación del largo del palpómero / vesícula sensorial = 2,3-3,1 ................
.......................................................................................... Simulium goeldii, Simulium ulyssesi

Machos
1 Escudo con 1+1 banda plateada submediana (Fig. 81 A) ......................................... 2
– Escudo sin 1+1 banda plateada submediana o solamente una mancha anterior
rectangular fina (Fig. 81 D) .............................................................................................. 4
2(1) Escudo con 1+1 banda plateada submediana uniéndose con el área grisácea
posterior (Fig. 81 K); gonostilo aproximadamente 1,5 más largo que ancho en la
base (Fig. 81 L) ............................................................................. Simulium quadrivittatum
– Escudo con 1+1 banda plateada submediana no uniéndose con el área grisácea
posterior (Fig. 81 A); gonostilo aproximadamente 2 veces más largo que ancho en
la base (Fig. 81 C) ............................................................................................................... 3
3(2) Escudo con 1+1 banda fina anterior submediana, llegando cerca de la mitad del
largo del escudo (Fig. 81 A) ........................................................... Simulium quadrifidum
– Escudo con 1+1 banda cuneiforme anterior submediana reducida al 1/4 anterior
del largo del escudo .............................................................................. Simulium cauchense
4(1) Con cierta iluminación las bandas plateadas del escudo se unen con 1+1 manchas
plateadas anteriores (Fig. 81 D); gonostilo con 1 espolón; relación largo/ancho en
la base del gonostilo = 2,5 ................................................................. Simulium cerradense
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 353

– Con ninguna iluminación las bandas plateadas laterales del escudo se unen con
1+1 manchas plateadas submedianas; gonostilo con 1- 3 espolones; relación del
largo/ancho en la base del gonostilo = 1,8-2,1 ........................................................... 5
5(4) Gonostilo con 2 o 3 espolones (Fig. 81 J) ..................................... Simulium daltanhani
– Gonostilo con 1 espolón (Fig. 81 H) ....................... Simulium goeldii, Simulium ulyssesi

Pupas
1 Branquia con 4 ramas ........................................................................................................ 2
– Branquias con 6-8 ramas .................................................................................................. 3
2(1) Branquias más largas que el capullo en la base (Fig. 82 A); largo 3,3-3,4 mm y 2,8
mm respectivamente; ramas secundarias de la branquia emergen cerca de las
ramas primarias (Fig. 82 C); V tergito abdominal con dentículos sobre el borde
anterior ............................................................................................... Simulium quadrifidum
– Largo de las branquias y de la base del capullo aproximadamente del mismo
tamaño (largo 2,2-2,6 mm y 2,0-2,6 mm respectivamente; ramas secundarias
emergiendo lejos de la mitad del largo de la branquia. V tergito abdominal sin
dentículos sobre el borde anterior ..................................................... Simulium cauchense
3(1) Branquia con 6 ramas ........................................................................................................ 4
– Branquia con 8 ramas ........................................................................................................ 6
4(3) Tronco basal de la branquia alargada y bifurcación de las ramas secundarias
relativamente lejos de la base (Fig. 82 K) ........................................... Simulium ulyssesi
– Tronco basal de la branquia corto y bifurcación de las ramas secundarias próxima
a la base (Figs. 82 F, L) ..................................................................................................... 5
5(4) Tejido del capullo fibroso, con fibras evidentes; capullo con prolongación dorsal
anterior (Fig. 82 L); tubérculos del tórax redondeados apicalmente; tergito
abdominal V con dentículos sobre el borde anterior .................. Simulium daltanhani
– Tejido del capullo esponjoso, sin fibras evidentes; capullo sin prolongación
anterior dorsal (Fig. 82 E); tubérculos del tórax aguzados apicalmente; tergito
abdominal V sin dentículos sobre el borde anterior .................... Simulium cerradense
6(3) Capullo extendido lateralmente y con agujeros (Fig. 82 M); bifurcación de las
ramas terciarias relativamente cercanas a la base .................. Simulium quadrivittatum
– Capullo no expandido lateralmente y con tejido compacto (Fig. 82 G); bifurcación
de las ramas terciarias relativamente lejos de la base ......................... Simulium goeldii

Larvas
1 Ganglio subesofágico pigmentado; longitud del cuerpo 4,8-5,5 mm; hendidura
postgenal aguzada apicalmente ........................................................ Simulium daltanhani
– Ganglio subesofágico no pigmentado; longitud del cuerpo 3,1-4,7 mm; hendidura
postgenal redondeada o no apicalmente (Figs. 83 D, S) ............................................ 2
2(1) Anillo anal con 80-84 hileras de ganchos; relación puente postgenal / altura del
hipostomio = 3,9; relación de los artejos antenales proximal, mediano y distal =
1:1:1 (Fig. 83 N) .................................................................................... Simulium cerradense
354 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

– Anillo anal con 57-66 hileras de ganchos; relación puente postgenal / altura del
hipostomio = 1,4-2,9; relación de los artejos antenales proximal, mediano y distal
= 1:0,3-0,9:1,1-1,8 .............................................................................................................. 3
3(2) Papilas rectales con 10-13 divertículos en cada lóbulo; proceso látero-mandibular
curvo (Fig. 83 C); relación largo del hipostomio/ puente postgenal = 2,0-2,9 ......
.................................................................... Simulium quadrifidum, Simulium quadrivittatum
– Papilas rectales con 4-7 divertículos en cada lóbulo; proceso látero-mandibular
recto o sinuoso; relación largo hipostomio/ puente postgenal = 1,4-2,1 ............. 4
4(3) Proceso látero-mandibular sinuoso, fino y apicalmente simple; relación de los
artejos antenales proximal, mediano y distal = 1:1,5:1,6 relación largo
hipostomio/ puente postgenal = 1,9-2,1 ......................................... Simulium cauchense
– Proceso látero-mandibular recto, grueso y apicalmente bífido o trífido; relación de
los artejos antenales proximal, mediano y distal = 1:1,1:1,4; relación largo
hipostomio/ puente postgenal = 1,4-1,6 ...................................................................... 5
5(4) Proceso látero-mandibular apicalmente bífido; rayos del abanico cefálico en
número de 38-47; longitud del cuerpo = 3,6-3,8 mm ....................... Simulium goeldii
– Proceso látero-mandibular apicalmente trífido; rayos del abanico cefálico en
número de 30-36; longitud del cuerpo = 4,1-4,2 mm ..................... Simulium ulyssesi

QUADRIFIDUM species group

Simulium (Coscaroniellum) quadrifidum LUTZ (Figs. 80 A-M; 81 A-C; 82 A-D; 83 A-K.)


Simulium quadrifidum LUTZ, 1917: 66; PINTO, 1931: 709; VULCANO, 1967: 19; TIDWELL et al., 1982: 62;
SHELLEY et al., 1984: 152; HAMADA & GRILLET, 2001: 34; HAMADA & FOUQUE, 2001: 955;
HAMADA & ADLER, 2001: 119; ALVAN-AGUILAR et al., 2005: 250-256.
Simulium (Coscaroniellum) quadrifidum: PY-DANIEL, 1983: 184; COSCARÓN, 1987: 25; 1991: 207.
Coscaroniellum quadrifidum PY-DANIEL & MOREIRA SAMPAIO, 1995: 118.
Simulium (Psaroniocompsa) quadrifidum: SHELLEY et al., 1997: 15; CROSSKEY & HOWARD, 1997: 57, 2004:
51; COSCARÓN ARIAS, 2003: 196, 205, 208, 215.
?Simulium rasii RAMÍREZ PÉREZ, 1980: 60.
Simulium torrealbai RAMÍREZ PÉREZ, 1980: 64.
Female: Wing length 1.7-2.0 mm. Frons and clypeus with bluish-green nacreous
pruinosity. Scutum black with 1+1 silvery submedian stripes becoming thinner posteri-
orly, reaching grayish posterior area; median black stripe wide posteriorly (Fig. 80 E);
with changing light position, silvery submedian stripe becomes thinner and does not
reach grayish posterior area. Scutum silvery. Pleurae dark grayish with silvery sheen.
Abdomen velvety black on tergites I-V, VI-IX with cereous sheen; tergite II with 1+1
large lateral silvery spots; III-V with thin grayish pruinose band on posterior border
(Fig. 80 J). Frons relatively wide (Fig. 80 A); fronto-ocular triangle relatively short (Fig.
80 B). Sensory vesicle of maxillary palp about half of palpomere and with numerous
tubercles (Fig. 80 C). Basal portion of cibarium well sclerotized apically with large
rounded submedian process and short teeth (Fig. 80 D). Leg shape and ornamentation
as in figures 80 F, G, H. Claws with short subbasal tooth (Fig. 80 I). Genitalia with
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 355

A Y

D
C

B
O
F I

G
H
L K

R
M Z
S Q

P T

V U X
Figure 80. Female. A-M: S. (Coscaroniellum) quadrifidum; A: frons; B: fronto-ocular triangle; C:
maxillary palp ; D: basal portion of cibarium; E: scutum; F-H: fore, mid and hind leg, respectively;
I: claw; J: abdomen, K: eighth sternite, gonapophyses and genital fork; L: spermatheca; M: cercus
and paraproct. N-R: S. (C.) cerradense; N: frons; O: fronto-ocular triangle; P: basal portion of
cibarium; Q-R: scutum with different position of light. S-U: S. (C.) goeldii; S: scutum; T: basal
portion of cibarium; U: genital fork. V: S. (C.) cauchense, frons. W-Z: S. (C.) quadrivittatum; W:
scutum; X; abdomen; Y: basal portion of cibarium; Z: cercus and paraproct.
356 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

general aspect as in figure 80 K. Eighth sternite with about 20 setae; spermatheca


subspherical (Fig. 80 L); paraproct short (Fig. 80 M).
Male: Wing length 1.6-1-7 mm. Scutum black with 1+1 silvery thin anterior
submedian cuneiform spots that can reach distally to half of scutum length (Fig. 81 A);
(this submedian spot can be absent in some specimens (SHELLEY et al., 1997)). Genitalia as
in figure 81 B; gonostylus subtriangular in shape with large spur (Fig. 81 C); ventral plate
slowly acuminated apically.
Pupa: Cocoon with reinforced border and slightly projected distally (Fig. 82 A), with
compact weave and evident threads; length at base 2.8-2.9 mm, on dorsum 2.0-2.5 mm,
gills 3.3-3.8 mm with 4 filiform branches, bifurcating near base (Fig. 82 C). Frontoclypeus
and exposed portion of thorax with regular quantity of acuminate tubercles (Fig. 82 B);
cephalic and thoracic trichomes bifid (Fig. 82 D). Abdominal tergite V with denticles on
anterior border.
Larva: Maximum length 4.0 mm. Cephalic apotome light brown with median
and lateral spots not evident (Fig. 83 A). Antenna with distal article elongated (Fig. 83
B), ratio of proximal, medial and distal articles = 1:0.7-0.9:1.2-1.3. Cephalic fan with
35-40 rays. Mandible as in figure 83 C. Postgenal cleft wide and deep, postgenal
bridge about 1/3 of hypostomium length (Fig. 83 D). Hypostomium with lateral teeth
acuminate and longer than median tooth (Fig. 83 E). Body integument with abundant
multibranched hairs as shown in figures 83 G-K. Ninth sternite with 1+1 ventral
papillae. Anal ring with 55-60 rows with 10-12 hooks each. Anal sclerite with
abundant hair with 1-4 branches and irregular size (Fig. 83 F). Rectal papillae with 10-
13 diverticula on each lobe.
Distribution: Bolivia: Cochabamba; Brazil: Amapá, Amazonas, Pará, Rondônia,
Roraima; Colombia; Ecuador: Esmeraldas, Napo, Pastaza; French Guiana; Guyana;
Suriname; Venezuela: Amazonas, Bolívar.
Bionomics: This species breeds in small creeks with clear water, frequently in
shaded places. Females have zoophilic habits (SHELLEY et al., 1997).
Discussion: This species is similar to S. cauchense and S. cerradense. The first species
can be distinguished by the silvery submedian stripes of females and males that are
shorter and do not reach beyond half the scutum length. In the pupa, the gills are
proportionally shorter, and they have denticles on abdominal tergite V. The larva has a
shorter distal antennal article and longer medial article. Simulium cerradense can be
differentiated from S. quadrifidum in the female by the presence of intervittae stripes on
the scutum and the acuminate submedian process of the cibarium, in the male by the
anterior subtrapezoidal spots, in the pupa by the spongy cocoon weave and the gills with
6 branches, and in the larva by the cephalic apotome with evident positive spots and the
medial antennal article longer than the proximal and the distal. Simulium rasii was
synonymized with some doubt, because material from Venezuela, Monagas and San
Antonio, determined by RAMÍREZ PÉREZ, has pupal gills with thicker and shorter primary
branches and more tubercles on the frontoclypeus and thorax.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 357

A
B

C
F

L
G

K I

Figure 81. Male. A-C: S. (Coscaroniellum) quadrifidum; A: scutum; B: genitalia, ventral view showing
gonocoxite, gonostylus, ventral plate and endoparameres; C: gonostylus, dorsal view (mounted on
slide). D-G: S. (C.) cerradense; D: scutum; E: ventral plate, median sclerite and endoparameres; F:
gonocoxite and gonostylus, ventral view; G: gonostylus (mounted on slide). H-I: S. (C.) goeldii, H:
gonocoxite and gonostylus on dorsal view; I: ventral plate. J: S. (C.) daltanhani, gonostylus (from
HAMADA & ADLER, 1998). K-L: S. (C.) quadrivittatum; K: scutum, L: gonostylus.
358 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

A
E

G
M

D F

I
C J

B H
Figure 82. Pupa. A-D: S. (Coscaroniellum) quadrifidum; A: lateral view; B: frontoclypeus with
trichomes and tubercles; C: portion of thorax with trichomes and gill; D: thoracic trichomes and
tubercles. E-F: S. (C.) cerradense; E: dorsolateral view; F: portion of thorax with trichomes and gill.
G-J: S. (C.) goeldii; G: dorsal view; H: frontoclypeus trichomes and sparse tubercles; I: portion of
thorax with trichomes and gill; J: thoracic tubercles with high magnification. K: S. (C.) ulyssesi, basal
portion of gill. L: S. (C.) daltanhani, lateral view (from HAMADA & ADLER, 1998). M: S. (C.)
quadrivittatum, dorsal view.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 359

N L

O
A
M

D
B

E
F

R
S Q
G
T
J I K

Figure 83. Larva. A-K: S. (Coscaroniellum) quadrifidum; A: cephalic apotome; B: antenna; C: teeth of
mandible; D: postgenal cleft and hypostomium; E: hypostomium; F: anal sclerite; G-K: different
body trichomes. L-P: S. (C.) cerradense; L: lateral view; M: head, dorsal view; N: antenna; O: teeth of
mandible; P: lateral sclerite of thoracic proleg. Q-S: S. (C.) goeldii; Q: antenna; R: teeth of mandible;
S: postgenal cleft and hypostomium. T: S. (C.) ulyssesi, latero-mandibular process.
360 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Simulium (Coscaroniellum) cerradense COSCARÓN, CERQUEIRA, SATO & LA SALVIA


(Figs. 80 N-R; 81 D-G; 82 E, F; 83 L-P.)
Simulium (Coscaroniellum) cerradense COSCARÓN et al., 1992: 111-113.
Simulium (Coscaroniellum) sp. A COSCARÓN, 1991: 206.
Simulium (Psaroniocompsa) cerradense: CROSSKEY & HOWARD, 1997: 57; 2004: 51.
Coscaroniellum cerradense PY-DANIEL & MOREIRA SAMPAIO, 1995: 118.
Simulium cerradense: HAMADA et al., 2004: 1-8.
Female: Wing length 1.8 mm. Scutum black with 1+1 silvery submedian stripes that
join with grayish posterior area, variation in width of stripes in accordance lighting (Figs.
80 Q, R); hair golden to silvery. Abdomen with tergites I-IV velvety black, V-IX grayish
with cereous sheen. Frons with weakly developed infra-frontal sutures (Fig. 80 N).
Fronto-ocular triangle wider than high (Fig. 80 O). Basal portion of cibarium with 1+1
submedian process distally acuminate and teeth of invaginated area very small (Fig. 80 P);
claws with subbasal tooth; leg hair filiform and petaloid. Genitalia with general aspect as
in S. quadrifidum; eighth sternite with about 20 setae on each side.
Male: Wing length 1.7 mm. Scutum velvety black with 1+1 anterior submedian
silvery spots, subtrapezoidal (Fig. 81 D); changing light position shows narrow band
joining to grayish lateral area. Ventral plate, median sclerite and endoparameres as in
figure 81 E. Gonostylus subtrianglular, twice as long as wide (Figs. 81 F, G).
Pupa: Cocoon weave spongy, thick not transparent, with reinforcement on
anterior border (Fig. 82 E). Length at base 2.6-2.8 mm, at dorsum 2.1-2.4 mm, gills 3.0-
4.0 mm. Frontoclypeus and exposed portion of thorax with abundant acuminate spots;
trichomes bifid and large (Fig. 82 F). Gill with 6 branches bifurcated near base.
Abdominal tergite V without denticles on anterior border. Abdominal terminal spines
short and with slightly acute apex.
Larva: Maximum length 4.3-4.7 mm. General aspect as in figure 83 L. Cephalic
apotome with positive spots evident (Fig. 83 M). Hairs of dorsum of head single. Antenna
surpassing apex of cephalic fan stem. Ratio of proximal, medial and distal articles =
1:1:1.1 (Fig. 83 N). Mandible with 2 marginal teeth and strong latero-mandibular process
(Fig. 83 O). Postgenal bridge longer than half of hypostomium length. Cephalic fan with
40-46 rays. Body integument with short hairs with 2.5 branches. Lateral sclerite of
thoracic proleg with 22-24 teeth arranged in 7-9 groups (Fig. 83 P). Ninth sternite with
1+1 ventral tubercles, slightly acuminate apically. Anal sclerite area with abundant
unbranched hairs. Anal ring with 80-84 rows with 12-13 hooks each. Rectal papillae with
5-6 diverticula on each lobe.
Distribution: Brazil: Bahia.
Bionomics: This species was collected in creeks 5-10 m wide and 0.5-2.0 m deep, in
crystalline waters with little to moderate flow over aquatic vegetation. Female habits are
unknown. The distributional area corresponds to the Cerrado region of northeastern Brazil.
Discussion: The ornamentation of the female scutum is similar to that of S.
quadrifidum and S. (Cerqueirellum) species, S. (Notolepria) and some species of S. (Psilopelmia)
and S. (Aspathia). Differentiation from S. quadrifidum was explained under that species. It can
be distinguished from the Cerqueirellum species group by the shape of the male gonostylus,
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 361

from S. (Notolepria) and S. (Psilopelmia) by the shape of the female paraprocts, and from S.
(Aspathia) by the shapes of the male gonostylus and ventral plate. The spongy cocoon weave
of S. cerradense permits it to be differentiated from these similar species. Only S. argentiscutum
has a similar cocoon, but the pupal gill and cocoon shape permit separation.

Simulium (Coscaroniellum) cauchense FLOCH & ABONNENC (Fig. 80 V.)


Simulium cauchense FLOCH & ABONNENC, 1946a: 1; SHELLEY et al., 1984: 144-145; HAMADA & GRILLET,
2001: 34; HAMADA & ADLER, 2001: 119; ALVAN-AGUILAR et al., 2005: 250-256.
Simulium (Coscaroniellum) cauchense: PY-DANIEL, 1983: 188; COSCARÓN, 1987: 25; 1991: 206.
Coscaroniellum cauchense PY-DANIEL & MOREIRA SAMPAIO, 1995: 118.
Simulium (Psaroniocompsa) cauchense: SHELLEY et al., 1997: 17; CROSSKEY & HOWARD, 1997: 57; 2004: 51.
Simulium sextobecium NUNES DE MELLO, 1974: 15.
Simulium rangeli RAMÍREZ PÉREZ, RASSI & RAMÍREZ, 1977: 163.
(Based on description by PY-DANIEL, 1983).
Female: Wing length 1.5-1.8 mm. Frons and clypeus with pruinosity sheen bluish
green. Scutum black with 1+1 silvery anterior submedian cuneiform spot not reaching
beyond half length of scutum with anterior or posterior illumination, abdomen coloration
very close to quadrifidum. Frons with pronounced infra-frontal sutures (Fig. 80 V); basal
portion of cibarium, claws and genitalia shape as in S. quadrifidum.
Male: Wing length 1.4-1.7 mm. Scutum black with 1+1 small cuneiform anterior
submedian silvery spots. Genitalia as in S. quadrifidum.
Pupa: Cocoon similar to that of S. quadrifidum; length along base 1.6-2.5 mm, on
dorsum 1.7-2.4 mm, gills 2.2-3.1 mm. Gills with 4 branches bifurcated near the base,
dorsal primary branch slightly longer than ventral; dorsal bifurcation can be near midpoint
of gill. Frontoclypeus and thorax with tubercles and trichomes as in S. quadrifidum.
Abdominal tergite V without denticles on anterior border.
Larva: Maximum length 3.0-3.8 mm. Color and morphology similar to S. quadrifidum.
Cephalic apotome with obvious positive spots. Ratio of proximal, medial and distal
antennal articles = 1:0.4-0.5:1.5-1.6. Mandible teeth as in S. cerradense; cephalic fan with
36-38 rays. Anal sclerite as in S. quadrifidum but with hair more evident. Anal ring with 60-
72 rows with about 13 hooks each. Rectal papillae with 3-8 diverticula each. Body
integument with single to multibranched trichomes.
Distribution: Brazil: Amapá, Amazonas, Pará, Roraima; French Guiana; Guyana;
Venezuela: Amazonas, Bolivar.
Bionomics: Females are zoophilic (SHELLEY et al., 1984: 144).
Discussion: The most similar species are S. quadrifidum and S. cerradense, but the
more elongated submedian silvery stripes that join with the posterior grayish area on the
scutum of females separate them. The male of S. quadrifidum has the silvery submedian
stripes thicker and more elongated, reaching half the length of the scutum, and in S.
cerradense the silvery submedian stripes are reduced to anterior subtrapezoidal spots.
Simulium cerradense can be separated also by the pupal gill with 6 branches. Simulium goeldii
and S. ulyssesi have some similarities, but their scutum is reddish brown and their pupal
gills have 8 and 6 branches, respectively.
362 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Simulium (Coscaroniellum) goeldii CERQUEIRA & NUNES DE MELLO


(Figs. 80 S-U; 81 H, I; 82 G-J; 83 Q-S.)
Simulium goeldii CERQUEIRA & NUNES DE MELLO, 1967: 125; PY-DANIEL, 1981: 176; RAMÍREZ PÉREZ,
1983: 47; CROSSKEY & HOWARD, 1997: 81; HAMADA & ADLER, 2001: 119; RÍOS-VELÁSQUEZ et al.,
2002: 114-117.
Simulium (Coscaroniellum) goeldii: COSCARÓN, 1987: 25.
Simulium (Psaroniocompsa) goeldii: CROSSKEY & HOWARD, 2004: 51.
Simulium scorzai RAMÍREZ PÉREZ, 1980b: 59.
Female: Wing length 1.7 mm. Frons and clypeus with silvery pruinosity and
nacreous sheen; scutum dark and scutellum dark reddish brown; scutum bordered
laterally and posteriorly by silvery pruinose area, with 1+1 sublateral, transverse silvery
spots that with some lighting join lateral grayish area (Fig. 80 S); with posterior lighting,
spots reduced to small oblique bands; pleurae and metanotum blackish with nacreous
pruinosity; scutum hair golden to tin colored. Abdomen blackish with 1+1 lateral gray
nacreous area on segment II-V. Basal portion of cibarium with median invagination and
1+1 submedian projections (Fig. 80 T); all this area with small acuminate teeth. Fronto-
ocular triangle wider than high. Claws with subbasal tooth. Genitalia as in quadrifidum;
eighth sternite with about 12 setae on each side; paraproct short, spermatheca enlarged;
genital fork with well sclerotized stem (Fig. 80 U).
Male: Wing length 1.7 mm. General coloration as in female; scutum without obvious
1+1 anterolateral silvery spots. Gonostylus surpassing gonocoxite by half length,
subtrapezoidal elongate shape and with 1 subapical spur (Fig. 81 H). Ventral plate slightly
acuminate apically (Fig. 81 I).
Pupa: Cocoon with thin weave and slight anterodorsal projection (Fig. 82 G); threads
weakly evident. Length along base 2.2-2.3 mm, on dorsum 1.7-2.0 mm, gills 2.3-2.5 mm.
Frontoclypeus and exposed portion of thorax with sparse (Fig. 82 H) and elongate tubercles
with blunt apices (Fig. 82 J). Trichomes long, frontal bifid, facial with 3-7 branches, thoracic
with 2-5 branches. Gills with 8 filamentous branches, relatively long basal trunk and 2 short
primary branches; primary dorsal branch bifurcated at short distance; dorsal secondary
branches bifurcated at short distance, and both dorsal tertiary branches bifurcated near half
of gill length (Fig. 82 I); abdominal tergite V with barely evident denticles.
Larva: Maximum length 3.5 mm. Color light reddish brown, with darker ring around first
abdominal segment. Cephalic apotome not ornamented with positive spots. Cephalic
integument without hair. Antenna thin with short median article (Fig. 83 Q); ratio of proximal,
medial and distal articles = 1:0.3-0.4:1.4-1.8. Mandible with robust latero-mandibular process
(Fig. 83 R). Cephalic fan with 33-38 rays. Postgenal cleft dome shaped; postgenal bridge about
3/4 of hypostomium height (Fig. 83 S); teeth of hypostomium evident. Lateral sclerite of
thoracic proleg elongated, with about 17 teeth arranged in 5-6 groups. Anal ring with 54-56
rows with about 9 hooks each. Anal sclerite with sparse single hairs on posterior area.
Distribution: Brazil: Amazonas, Pará, Rondônia; Venezuela: Amazonas.
Bionomics: This species breeds in small creeks, generally on leaves in fast flow, in
clear or dark acidic waters, with temperatures of about 25 °C, which are typical in the
Amazonian region. Females were collected biting humans.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 363

Discussion: This peculiar species is easy to distinguish by the arrangement of the


pupal gill, tubercle shape of the frontoclypeus and thorax, coloration and ornamentation
of the scutum, and gonostylus shape, all of which make it difficult to include it in the S.
quadrifidum species group. The closest species are S. ulyssesi, which can be distinguished by
the pupal gill with 6 branches and the latero-mandibular process that is trifid distally, and
S. daltanhani, which has 6 branches in the pupal gill and a male gonostylus with 3 spurs.

Simulium (Coscaroniellum) ulyssesi PY-DANIEL & COSCARÓN (Figs. 82 K; 83 T.)


Coscaroniellum ulyssesi PY-DANIEL & COSCARÓN, 2001: 52-57.
Simulium 6 B1 HAMADA & ADLER, 1999: 44.
Simulium ulyssesi: RÍOS-VELÁSQUEZ et al., 2002: 114-117.
Female and male coloration and morphology are similar to that of S. goeldii.
Female: Fronto-ocular triangle reduced, basal portion of cibarium well sclerotized
with teeth bordering median invagination short and blunt, and laterally more developed
and apically acute.
Male: Gonostylus subtrapezoidal with flattened spur subapically positioned.
Pupa: Cocoon shape as in S. goeldii, length along base 1.8-2.7 mm; on dorsum 2.1-2.9
mm; gills 4.5-5.2 mm. Thoracic tubercles subconical; trichomes elongated, facial with 3
branches, frontal bifid, thoracic with 3-4 branches. Gill with 6 filamentous branches and
long basal trunk, primary branches bifurcated at short distance, and dorsal secondary
branches bifurcated at twice length of primaries (Fig. 82 K) Abdominal tergite V with
small teeth on anterior border.
Larva: Maximum length 4.1-4.2 mm. Coloration as for S. goeldii. Cephalic apotome
with small single hairs. Antennae surpassing apex of cephalic fan stem; ratio of proximal,
medial and distal articles = 1:0.4-0.5:1.1-1.2. Mandible with 1 strong latero-mandibular
process straight and apically with 3 branches (Fig. 83 T). Cephalic fan with 30-36 rays.
Postgenal cleft shape, ratio of postgenal bridge to hypostomium, anal sclerite and number
of anal ring hook rows similar to S. goeldii.
Distribution: Brazil: Amazonas.
Bionomics: Simulium ulyssesi lives in the same environment as S. goeldii.
Discussion: This species is very similar to S. goeldii with which it is sympatric. It is
possible to differentiate it because S. goeldii has a pupa with a shorter gill of 8 branches, a
shorter cocoon, and a shorter larva with a relatively shorter medial antennal article and the
apex of the latero-mandibular process with 2 branches.

Simulium (Coscaroniellum) daltanhani HAMADA & ADLER (Figs. 81 J; 82 L.)


Simulium (Coscaroniellum) daltanhani HAMADA & ADLER, 1998a: 317; 2001: 119.
Simulium (Psaroniocompsa) daltahani: CROSSKEY & HOWARD, 2004: 51.
Female: Wing length 2.0 mm. General coloration similar to that of S. goeldii; scutum
blackish to reddish brown with 1+1 rectangular silvery anterolateral spots joining with
lateral silvery area, 4 short submedian pruinose bands faintly visible; scutum hair golden;
scutellum and metanotum blackish, the latter with silvery pruinosity. Abdomen similar to
364 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

that of S. goeldii. Fronto-ocular triangle evident, rather shallow, infra-frontal sutures short;
basal portion of cibarium well sclerotized and with submedian projections and teeth as in
S. goeldii; sensory vesicle of maxillary palp elongated, about 2/3 of palpomere length;
claws with subbasal tooth; genitalia similar to those of S. goeldii.
Male: Wing length 1.9 mm. Scutum blackish, with 1+1 narrow anterolateral silver
spots connected with silver lateral band, and 1+1 faintly visible silvery submedian stripes;
hair reddish gold. Gonostylus about half of gonocoxite length, subtrapezoidal with 1-3
subapical spurs (Fig. 81 J). Ventral plate and endoparameres as in S. goeldii.
(Pupa and larva based on HAMADA & ADLER, 1998a).
Pupa: Cocoon similar to that of S. goeldii. Length along base 2.7-3.4 mm, on dorsum
3.0-3.4 mm, gills 3.9-4.0 mm. Frontoclypeal and thoracic tubercles rounded distally. Head
and dorsum of thorax with bifid trichomes. Gills with 6 filamentous branches, with trunk
short and subdivided at short distance from base, with secondary dorsal branches very
short (Fig. 82 L).
Larva: Maximum length 4.8-5.5 mm. Cephalic apotome slightly darkened on median
and basal area. Postgenal cleft dome shaped. Postgenal bridge 0.5-0.8 times hypostomium
length. Subesophageal ganglion heavily pigmented. Antenna about same length as
cephalic fan stem; ratio of proximal, medial and distal articles = 1:0.7-0.9:1.2-1.5.
Cephalic fan with 49-51 rays; latero-mandibular process single. Anal sclerite with single to
trifid hairs. Anal ring with 60-66 rows of 12-13 hooks each. Rectal papillae with 9-10
diverticula on each lobe.
Distribution: Brazil: Amazonas.
Bionomics: Simulium daltanhani breeds in small, shallow, acidic streams with sandy
bottoms and warm temperatures (26-29 °C) flowing over fallen leaves and grasses. Female
feeding habits are unknown.
Discussion: The similar scutal ornamentation and cocoon with an anterior projec-
tion can confuse S. daltanhani with S. goeldii and S. ulyssesi, but the latter two don’t have the
submedian faintly visible silvery stripes. Simulium goeldii also can be differentiated by the 8
pupal gill branches; S. ulyssesi has 6 branches but they are directed forward, not laterally
expanded, and the larval latero-mandibular process has 3 branches.

QUADRIVITTATUM species group


The S. quadrivittatum group has particular characters, such as narrow straight silvery
submedian stripes joined posteriorly by a transverse straight, narrow band before joining
the posterior grayish area on the scutum, and a cibarium with 1+1 submedian elongated
thin processes with acute teeth. The following characters indicate the similarity of this
group to Simulium (Psaroniocompsa), S. (Cerqueirellum) and S. (Coscaroniellum): female scutum
with 1+1 submedian stripes and, in some illumination, with shadow-like intervittae spots,
short rounded paraproct, subconical gonostylus, wing with basal sector of R without hair;
long bifid cephalic and thoracic trichomes on the pupa, a single dark ring on the first
abdominal segment of the larva, and the cephalic apotome with positive median, lateral
and posterior spots, Simulium (Coscaroniellum) is the closest group, as evidenced in the
female by 1+1 submedian processes on the cibarium, relatively deep fronto-ocular
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 365

triangle, and infra-frontal sutures; in the male by the subconical gonostylus; and in the
larva by the hypostomium with intermediate teeth about 1/3-1/2 the height of the
median and lateral teeth, and the distal antennal article nearly two times longer than the
medial article. This species group consists of only one species.

Simulium (Coscaroniellum) quadrivittatum LOEW (Figs. 80 W-Z; 81 K, L; 82 M.)


Simulium quadrivittatum LOEW, 1862: 186; WYGODZINSKY, 1953b: 334-336; SHELLEY & ARZUBE, 1985:
285; COSCARÓN, 1987: 22.
Simulium (Psilopelmia) quadrivittatum. SHELLEY et al., 1989: 336; CROSSKEY & HOWARD, 1997: 60; 2004:
54; SHELLEY et al., 2002: 167-169.
Simulium (Psaroniocompsa) quadrivittatum COSCARÓN ARIAS, 2003: 196, 204, 209, 215.
Psilopelmia quadrivittatum: RUBTSOV & GARCIA AVILA, 1972: 19-23.
Wilhelmia mallochi ENDERLEIN, 1925: 208.
Simulium fairchildi VARGAS, 1942b: 458.
Simulium (Byssodon) quadrivittatum: VARGAS & DIAZ-NÁJERA, 1948b: 349; 1957b: 162, 262.
Simulium metallicum MALLOCH, 1914: 25 part (nec BELLARDI, 1859).
?Simulium (Byssodon) benjamini DALMAT, 1952: 339-344; 1955: 121-126 (n. syn.).
Female: Wing length 1.6-2.0 mm. Frons and clypeus blackish with bluish pruinosity.
Scutum black with 1+1 silvery pruinose submedian stripes (Fig. 80 W), joining with dark
grayish posterior area; vittae when posteriorly illuminated show anterior small shadow-like
intervittae spots. Fronto-ocular triangle deep, wider than high. Abdomen blackish with
posterior border grayish, wide on segments III-V (Fig. 80 X). Cibarium basal portion with
sclerotized rim, slightly depressed medially and with 1+1 thin and sclerotized submedian
process and acute teeth (Fig. 80 Y). Basal portion of R without hair, claws with small subbasal
tooth. Gonapophysis with straight distal edge; paraproct short with rounded edge (Fig. 80 Z).
Male: Wing length 1.8 mm. Scutum black with 1+1 silvery thin submedian stripes
that surpass half of scutum length (Fig. 81 K); (in specimens without hair, submedian
silvery stripes are wider with less silvery color). Gonostylus subconical with blunt apex
and elongated subapical spur (Fig. 81 L).
Pupa: Cocoon slipper shaped with basal and dorsal protuberances, lateral sides
expanded with open loops and softly reinforced on anterior edge (Fig. 82 M), length along
base 2.4-2.6 mm, on dorsum 1.6-1.8 mm, gills with 2.8-3.2 mm. Gill with 8 branches with
secondary and tertiary branches bifurcated closer to base than in S. goeldii. Frontoclypeus
and thorax with sparse rounded tubercles; cephalic and thoracic trichomes bifid and
relatively long.
Larva: Cephalic apotome with positive median, mediolateral and posterolateral
spots. Antenna longer than cephalic fan stem; median article shorter than other two
articles; cephalic fan with 40-42 rays. Ratio of proximal, medial and distal article = 1:0.6-
0.7:1.4-1.5. Hypostomium with intermedian teeth lower by at least one third median and
lateral teeth. Postgenal bridge about as long as hypostomium height. Mandible with 1
latero-mandibular process; supramarginal setae absent. Body with dark band around first
abdominal segment; integument with sparse hair dorsally. Ninth abdominal segment with
ventral tubercles. Anal ring with 62-65 rows with 11-13 hooks each.
366 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Distribution: Belize: Belize, Cayo, Stan Creek, Toledo; Colombia: Choco; Costa
Rica: Cartago, Limón; Cuba; Dominican Republic; Ecuador: Cotopaxi, El Oro,
Esmeraldas, Manabi, Pichincha; Guatemala; Mexico: Campeche, Chiapas, Oaxaca,
Puebla, Veracruz; Nicaragua; Panama; Puerto Rico; Venezuela.
Bionomics: This species breeds in small creeks with clear water and slow current, on
leaves of aquatic vegetation in forested areas of lowlands (Ecuador). Females bite cattle
and humans and are considered vectors of Onchocerca volvulus in Ecuador.
Discussion: Simulium quadrivittatum can be confused with species of S. (Cerqueirel-
lum), S. quadrifidum, S. metallicum and S. haematopotum by its scutum, but the peculiar
cibarium shape allows it to be separated easily.

Simulium (Ectemnaspis) ENDERLEIN (Figs. 84-90; 145-148.)


Simulium (Ectemnaspis) ENDERLEIN, 1934b: 281.
Type-species: Ectemnaspis macca ENDERLEIN, 1934b = Simulium bicoloratum MALLOCH, 1912.
Female: Medium to large size, wing length 1.5-3.0 mm. General coloration yellowish
to blackish brown. Scutum yellowish-orange without silvery longitudinal bands, or if
present, not reaching posterior border, frequently with median and sublateral areas brown
to blackish, bordered by yellow extended in 1+1 areas over humeral plates and 1 median
posterior area. Scutellum yellow to yellowish brown. Metanotum generally blackish.
Abdomen generally yellow to brownish on tergites I-IV, blackish on posterior area.
Pleurae from yellow to black pollinose. Legs from yellow to blackish with some portions
black. Fronto-ocular triangle generally wider than high. Basal portion of cibarium with
sclerotized rim, from untoothed to covered by abundant teeth with median area slightly
covered by small tubercles or flanked by 1+1 submedian large process covered by small
teeth. Basal sector of R generally haired. Claws generally with stout subbasal tooth.
Paraproct subtrianglular, apically elongated and frequently with rounded basal lobe.
Gonapophysis divergent below, lightly sclerotized internally, without hair or spiculae.
Genital fork with stem as long as length of posterior branches joined.
Male: Wing length 1.8-3.0 mm. Pigmentation as in female. Scutum unbanded, rarely
with 1+1 silvery anterior wedges. Posterior basitarsus ratio length/width 3.5-7.5. Gono-
stylus about same length as gonocoxite, subcylindrical, with slight distal curvature and 1
terminal spur. Ventral plate in ventral view generally about as long as wide with slight
lateral concavity; endoparameres with strong hooks.
Pupa: Cocoon slipper shaped sometimes with dorsal anterior projection and
longitudinal dorsal crest; weave from resistant with evident threads to soft like felt or
spongy and without obvious threads. Gill generally with 6-8 thin filaments, but can be
from 2-250, and in some cases with thick and irregular filaments. Two pairs of trichomes
on frontoclypeus and 5 on thorax, well developed with 3-45 branches. Abdomen with
terminal spur short.
Larva: Maximum length 4.8-7.0 mm. Tegument with or without trichomes. Cephalic
apotome generally with pigmentation on basal or subbasal area with light median subbasal
window. Postgenal cleft deep, nearly reaching hypostomium, with postgenal cleft shorter
than hypostomium height. Antenna with medial and distal segment about same length and
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 367

both longer than proximal. Cephalic fan with 35-50 rays. Anal sclerite without scales. Anal
ring with 68-113 rows of hooks. Diverticula of rectal papillae generally 3-13 but can be 1-20.
Simulium (Ectemnaspis) can be subdivided into 4 species groups: BICOLORATUM,
ROMANAI, PERFLAVUM and DINELLII.

SIMULIUM (ECTEMNASPIS) species groups


BICOLORATUM species group: yellowish scutum and scutellum, scutum frequent-
ly slightly darkened to blackish median and laterally, but bordered by yellow, metanotum
blackish; abdomen with segments yellowish to brown. Cibarium smooth with thin
sclerotized rim between cornuae. Paraproct subtriangular, nearly as long as wide at base,
blunt, rounded apically. Gonostylus subcylindrical, slightly curving distally; ventral plate with
slight lateral constriction. Cocoon frequently with thick spongy or felt-like weave, reinforced
anterior border and sometimes with dorsal ridge; frontoclypeus and thorax sometimes with
big trichomes with about 50 branches; gill with 2-10 branches; larval cephalic apotome
generally darkened on posterior half, with negative median, lateral and basal spots; rectal
papillae diverticula of normal size. Fifteen species in the group: S. anaimense, S. antonii, S.
arcabucense, S. bicoloratum, S. bicornutum, S. cormonsi, S. furcillatum, S. ignescens, S. jaimeramirezi, S.
pautense, S. pifanoi, S. roquemayu, S. rubiginosum, S. tolimaense, S. tunja.
ROMANAI species group: scutum with variable coloration from yellowish to
totally black but bordered by yellow, scutellum yellowish, metanotum black; abdomen
with segment I (II-III) yellowish, the following ones dark-brown; cibarium with median
area slightly depressed and covered with tubercles or plaques laterally to cornuae; base
with thin teeth-like spines; paraproct subtriangular, nearly as long as wide at base, with
blunt apex. Male coloration mostly blackish, gonostylus subcylindrical and slightly curved
distally; ventral plate with marked lateral constriction. Pupal cocoon from compact to
thick with spongy or felt-like weave; gill with 6-8 branches. Larval ornamentation of
cephalic apotome and general morphology as in S. bicoloratum species group. Six species in
the group: S. albanense, S. gabaldoni, S. lutzianum, S. mayuchuspi, S. romanai, S. wolffhuegeli.
PERFLAVUM species group: yellowish scutum, scutellum and metanotum; abdo-
men with segments I (II-III) yellowish and following ones dark brown; cibarium with
median portion elevated, sometimes scarcely invaginated at center and with space
between cornuae, covered by acute teeth-like spines; paraproct subtriangular nearly 2
times longer than wide at base, acute distally and with abundant robust spiculae
posteriorly. Male coloration similar to that of female; gonostylus subcylindrical, about as
long as or longer than gonocoxite. Ventral plate as long as wide, with lateral constriction.
Pupal cocoon frequently with irregular weave, leaving open areas anteriorly, and with 8-20
branches; larval cephalic apotome without peculiar ornamentation and rectal papillae with
10-15 diverticula elongated. Six species in the group: S. kabanayense, S. perflavum, S.
rorotaense, S. suarezi, S. trombetense, S. maroniense.
DINELLII group: scutum, scutellum and metanotum yellowish, sometimes with 1+1
submedian silvery nacreous stripes on anterior half; abdomen with segments yellowish, V-IX
yellowish brown; fronto-ocular triangle wider than high; cibarium with 1+1 submedian
relatively thick process covered with small teeth leaving median curved smooth or wrinkled
368 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

concavity; paraproct slightly longer than wide at base, and rounded distally. Male with same
coloration as female; gonostylus subcylindrical with slight curvature, slightly shorter than
gonocoxite; ventral plate nearly wider than long with or without very slight lateral concavity.
Pupal cocoon with compact weave, threads evident; frontoclypeal and thoracic trichomes
with 2-8 branches; gill with 8 branches. Larval cephalic apotome mostly uniformly colored
without particular ornamentation; postgenal cleft deep ,reaching base of hypostomium. Five
species in the group: S. antillarum, S. dinellii, S. ochraceum, S. nuneztovari, S. shewellianum.

KEYS TO SIMULIUM (ECTEMNASPIS) SPECIES

Females
Simulium flavipictum not included.

1 Scutum yellowish with median and 1+1 sublateral areas orange to blackish brown
(Figs. 84 A, B); metanotum predominately blackish brown or totally blackish gray
................................................................................................................................................ 2
– Scutum totally yellowish; metanotum predominately yellowish ............................. 16
2(1) Cibarium untoothed and with thin sclerotized rim (Figs. 84 I, J) .............................
....................................................................................... BICOLORATUM sp. group......3
– Cibarium toothed with border of median area depressed and covered with
tubercles, and with acute lateral teeth (Figs. 84 K-P) ...................................................
................................................................................................ ROMANAI sp. group......11
3(2) Basal sector of R without hair (Fig. 85 A) ................................................................... 4
– Basal sector of R with hair .............................................................................................. 8
4(3) Scutum yellowish orange, without brown color (Fig. 84 A) ..................................... 5
– Scutum yellowish to brown dark on disc ...................................................................... 6
5(4) Wing length 3.0 mm; abdominal tergites I-IV yellowish; paraproct length about 1/
4 longer than cerci width at base ........................................................... Simulium antonii
– Wing length 2.4-2.6 mm; abdominal tergites I-III yellowish; paraproct as long as
cerci width at base (Figs. 85 G, H) ........................ Simulium pautense, Simulium pifanoi
6(4) Wing length 2.8-2.9 mm; paraproct longer than basal width (Fig. 85 I) ..................
......................................................................................................................... Simulium tunja
– Wing length 2.4-2.7 mm; paraproct about as long as basal width (Fig. 85 E) ...... 7
7(6) Abdominal tergites I-IV totally grayish yellow ..............................................................
......................................... Simulium furcillatum, Simulium bicornutum, Simulium tolimaense
– Abdominal tergites I-IV yellow with tergal plates III-IV brownish on posterior
border ...................................................................................................Simulium arcabucense
8(3) Scutellum and abdominal tergites I-IV brownish yellow .......................................... 9
– Scutellum yellowish or greenish orange; abdominal tergites II-IV yellow to
yellowish brown ................................................................................................................ 10
9(8) Maxillary palp sensory vesicle length less than 1/3 palpomere length (Fig. 84 Y)
.................................................................................................................... Simulium ignescens
– Maxillary palp sensory vesicle length nearly 1/2 palpomere length (Fig. 84 X) ....
...............................................................................................................Simulium rubiginosum
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 369

10(8) Scutum totally yellow; abdominal tergites I-III yellowish, IV brown on tergal
plate; length of paraproct about as long as basal width (Fig. 85 F) ..........................
.................................................................................................................Simulium roquemayu
– Scutum generally yellow with extensive black (Fig. 84 B), but central area from
brown to orange; abdominal tergites I-II yellowish orange, III-IV blackish brown;
length of paraproct about 1/3 longer than basal width ..............................................
...................................... Simulium bicoloratum, Simulium cormonsi, Simulium jaimeramirezi
11(2) Basal portion of R glabrous (Fig. 85 A); basal portion of cibarium with rounded
tubercles medially and sparse and diminutive spiculae laterally (Fig. 84 K) ...........
................................................................................................................... Simulium albanense
– Basal portion of R with hair; basal portion of cibarium with tubercles or small
plates of different shape medially and generally with abundant small teeth-like
spines laterally (Figs. 84 L-P) ......................................................................................... 12
12(11) Scutum, scutellum, metanotum and abdomen blackish (Fig. 84 C); claw without
subbasal tooth (Fig. 85 B) ............................................................... Simulium mayuchuspi
– Scutum, scutellum, metanotum and abdomen yellowish in variable proportion;
claw with subbasal tooth (Fig. 85 C) ............................................................................ 13
13(12) Scutum orange; median area of cibarium with subovoidal tubercles (Fig. 84 L)
...................................................................................................................Simulium gabaldoni
– Scutum yellowish orange but frequently with blackish brown spots on median and
1+1 sublateral areas; cibarium with subquadrate to subspheroidal small plates on
median area (Figs. 84 M-P) ............................................................................................ 14
14(13) Abdominal tergites I-II (III) yellowish ........................................... Simulium lutzianum
– Abdominal tergites I yellowish ...................................................................................... 15
15(14) Abdominal tergites II-IV yellowish to yellowish brown with median tergal plate
on III-V dark brown (Fig. 84 G); Sc with 0-3 hairs ................... Simulium wolffhuegeli
– Abdominal tergites II-IV blackish brown to black; Sc with 6-9 hairs ......................
.................................................................................................................... Simulium romanai
16(1) Abdominal tergites IV-VIII generally blackish brown with tergal plate yellowish;
cibarium median area with 1+1 submedian wide and short elevations covered by
strong teeth followed by minute teeth (Figs. 84 Q, R); paraproct apically
acuminate (Figs. 85 M, N) to moderately blunt (Fig. 85 K) .......................................
......................................................................................... PERFLAVUM sp. group......17
– Abdominal tergites I-IV yellowish (Fig. 84 H); tergites V-VII from yellow to
yellowish brown or yellowish to light gray; cibarium median area with 1+1
submedian elongated processes flanking median depressed well-sclerotized area,
superficially from smooth to wrinkled (Figs. 84 T-W); paraproct blunt apically
(Figs. 85 O-Q) ...................................................................... DINELLII sp. group......21
17(16) Legs and abdominal tergites all yellowish; fronto-ocular triangle higher than wide;
paraproct apically blunt (Fig. 85 K); wing length 2.5-2.7 mm ......... Simulium perflavum
– Legs and abdominal tergites, at least IV-VII, blackish brown, paraproct generally
acuminate apically (Figs. 85 M, N); wing length 2.2-2.3 mm ................................. 18
18(17) Metanotum yellow to yellowish brown ....................................................................... 19
– Metanotum blackish ........................................................................................................ 20
370 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

19(18) Sensory vesicle of maxillary palp about 1/3-1/4 length of palpomere (Fig. 84 b)
........................................................................... Simulium maroniense, Simulium trombetense
– Sensory vesicle of maxillary palp about 1/2 length of palpomere (Fig. 84 c) .......
.................................................................................................................. Simulium rorotaense
20(18) Abdominal tergite III yellowish; ratio of paraproct length/width at base = 1.3
(Fig. 85 L) .......................................................................................... Simulium kabanayense
– Abdominal tergite III blackish; ratio of paraproct length/width at base = 1.7 (Fig.
85 N) ........................................................................................................... Simulium suarezi
21(16) Scutum homogeneously pigmented without silvery submedian longitudinal bands
.............................................................................................................................................. 22
– Scutum with 1+1 short silvery submedian longitudinal bands (Fig. 84 E) ......... 23
22(21) Sensory vesicle of maxillary palp shorter than 1/2 of palpomere length, (Fig. 84
a); cibarium with 1+1 acuminate submedian processes (Fig. 84 U); wing length 1.9
mm ..................................................................................................... Simulium shewellianum
– Sensory vesicle of maxillary palp longer than 1/2 of palpomere length; cibarium
with 1+1 submedian processes blunt (Fig. 84 T); wing length about 2.1 mm .......
.................................................................................. Simulium dinellii, Simulium nuneztovari
23(21) Wing 2.2-2.3 mm; scutum matte yellow with 1+1 wedges connected with silvery
anterolateral area (Fig. 84 E); cibarium with median depressed area wrinkled or
with small spines (Fig. 84 V); metanotum dark brown ................ Simulium ochraceum
– Wing 1.9 mm; scutum reddish with 1+1 wedges not well connected with
anterolateral silvery area (Fig. 84 D); cibarium with median depressed area
smooth, without spines or wrinkling and bordered basally by thick rim (Fig. 84
W); metanotum yellowish .................................................................. Simulium antillarum

Males
Unknown: Simulium anaimense, S. antonii, S. roquemayu, S. tolimaens.

1 Scutum, scutellum, metanotum and abdomen dark grayish; scutum with posterior
1/4 black (Fig. 86 C); ventral plate with strong lateral constrictions (Fig. 87 G)
............................................................................................................... Simulium mayuchuspi
– Scutum and abdomen all yellowish, or having this pigment reduced to border of
scutum and abdominal tergite I (Figs. 86 A, D; ventral plate with or without lateral
constrictions (Figs. 87 B, H) ............................................................................................ 2
2(1) Scutum yellowish orange, with black in different proportions from small dark
brown to blackish spots on median and sublateral areas of disc (Fig. 86 B);
metanotum blackish ........................................................................................................... 3
– Scutum yellowish; metanotum brownish yellow to dark brown ............................ 14
3(2) Gonostylus about 1/2 to 2/3 of gonocoxite length, subquadrate with distal
border truncate (Figs. 86 M, O); ventral plate wide with length/width ratio = 3.0-
4.0 (Figs. 87 E, I) ................................................................................................................ 4
– Gonostylus about 3/4 or more of gonocoxite length, subcylindrical with
acuminate distal border (Figs. 86 K, N); ventral plate not as wide as in above, with
length/width ratio = 1.1-1.9 (Figs. 87 D, H) ............................................................... 5
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 371

4(3) Scutum black, with yellow area restricted to pronotal angles, laterally and narrow
posterior border; scutellum dark brown and abdomen blackish; gonostylus with 8-
10 terminal spurs (Fig. 86 O) .......................................................... Simulium wolffhuegeli
– Scutum yellowish with blackish pigmentation on disc as in Simulium romanai (Fig.
86 D), in some specimens; abdominal tergites I-III yellowish; gonostylus with 1
apical spur (Fig. 86 M) ......................................................................... Simulium albanense
5(3) Lateral areas of ventral plate strongly concave (Fig. 87 H) ...................................... 6
– Lateral areas of ventral plate slightly concave (Figs. 87 A, C, D) ........................... 7
6(5) Length/width ratio of hind basitarsus = 4.8-5.1; scutum frequently mostly black
as in figure 86 D ..................................................................................... Simulium romanai
– Length/width ratio of hind basitarsus = 6.5-7.6; scutum yellowish orange ...........
................................................................................................................. Simulium lutzianum
7(5) Basal sector of R haired ................................................................................................... 8
– Basal sector of R glabrous ............................................................................................... 9
8(7) Abdominal tergites I-IV yellowish brown, the following tergites black ...................
.............................................................................. Simulium rubiginosum, Simulium ignescens
– All abdominal tergites black ................... Simulium bicoloratum, Simulium jaimeramirezi
9(7) Wing length 2.9-3.0 mm; abdominal tergites I-III or IV yellowish brown to yellow
(Fig. 86 F); ventral plate with slight lateral concavity ............................................... 10
– Wing length 2.5 mm; abdominal tergites I-II or I-III yellowish, the following
tergites blackish; ventral plate laterally convex or with slight concavity .............. 11
10(9) Abdominal tergites I-IV greenish yellow with small dark brown area on tergite IV (Fig.
86 F); length/width ratio of posterior basitarsus = 3.5 ................. Simulium bicornutum
– Abdominal tergites I-III yellowish brown with large brown dark area on tergite IV
(Fig. 86 G); length/width ratio of posterior basitarsus = 4.0 ......... Simulium furcillatum
11(9) Ventral plate laterally convex (Fig. 87 B); gonostylus shorter than gonocoxite
(Figs. 86 I-J) ....................................................................................................................... 12
– Ventral plate not convex laterally (Figs. 87 A, D); gonostylus about same length as
gonocoxite (Figs. 86 H, L) ............................................................................................. 13
12(11) Scutum with 1+1 elongated lateral blackish stripe and median T shaped one on
anterior 1/3 (Fig. 86 B) ......................................................................... Simulium pautense
– Scutum with 1+1 short lateral brownish stripes ................................ Simulium pifanoi
13(11) Length/width ratio of posterior basitarsus = 4.3; wing length 2.7 mm; abdominal
tergite III yellowish with tergal plate brownish to black .................... Simulium tunja
– Length/width ratio of posterior basitarsus = 7.5; wing length 2.5 mm; abdominal
tergite III yellow with tergal plate brownish ................................ Simulium arcabucense
14(2) Ventral plate with strong lateral concavity (Fig. 87 F); abdominal tergites black
...................................................................................................................Simulium gabaldoni
– Ventral plate generally without strong lateral concavity (Fig. 87 O); abdominal
tergites all yellow or reddish-yellowish brown ........................................................... 15
15(14) Scutum with short silvery anterior submedian bands (Fig. 86 E); ventral plate
laterally convex (Fig. 87 O) ............................................................................................ 16
– Scutum homogeneously ornamented, without bands; ventral plate generally with
small lateral concavity (Figs. 87 J-N) ........................................................................... 17
372 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

16(15) Scutum yellowish orange, with 1+1 silvery stripes on anterior third .......................
.................................................................................................................. Simulium ochraceum
– Scutum yellowish-reddish brown, with 1+1 silvery stripes on anterior half or
surpassing it .......................................................................................... Simulium antillarum
17( 15) Ventral plate distal portion narrowing slowly (Fig. 87 N); gonostylus shorter than
gonocoxite (Fig. 86 S) ..................................................................................................... 18
– Ventral plate distal portion narrowing abruptly after 1/2 its length (Figs. 87 K-M);
gonostylus not shorter than gonocoxite (Figs. 86 P-R) ...............................................
......................................................................................... PERFLAVUM sp. group......20
18(17) Abdomen blackish brown; wing length 1.8 mm ....................... Simulium shewellianum
– Abdomen all reddish yellow to greenish yellow; wing length 2.0-2.5 mm .......... 19
19(18) Abdomen reddish yellow; wing length 2.5 mm .................................. Simulium dinellii
– Abdomen greenish yellow; wing length 2.0 mm ........................ Simulium nuneztovari
20(17) Metanotum and abdomen yellowish; gonostylus longer than gonocoxite; wing
length 2.6-2.7 mm ................................................................................ Simulium perflavum
– Metanotum and abdomen brownish; gonostylus not longer than gonocoxite; wing
length 1.8-2.6 mm ............................................................................................................ 21
21(20) All abdominal tergites brown to blackish brown; wing length 2.0-2.1 mm .............
................................................................................................................ Simulium trombetense
– Abdominal tergites I or II yellowish to yellowish brown, remaining tergites
blackish brown; wing length 1.8-2.6 mm .................................................................... 22
22(21) Abdominal tergite I brown or yellowish brown, remaining tergites dark brown
............................................................................... Simulium suarezi, Simulium kabanayense
– Abdominal tergites I-II yellowish brown, remaining tergites dark brown ...............
.............................................................................. Simulium maroniense, Simulium rorotaense

Pupae
Unknown: Simulium antonii.

1 Gill composed of 2-5 branches; cocoon with prominent anterodorsal projection


(Figs. 88 C, D, F) ................................................................................................................ 2
– Gill composed of 6-250 filaments; cocoon without prominent anterodorsal
projection ............................................................................................................................. 4
2(1) Gill branches relatively thin (Fig. 88 F) ........................................... Simulium tolimaense
– Gill composed of 2-4 very thick branches ................................................................... 3
3(2) Gill with 2 similar branches and small internal protuberance on each (Fig. 88 C)
................................................................................................................ Simulium bicornutum
– Gill with 4 branches, 1 thick inferior branch and 3 thin ones positioned overhead
(Fig. 88 D) .............................................................................................Simulium furcillatum
4(1) Cocoon with anterior open loops (Fig. 88 L); gill with 17-250 relatively thick
branches (Figs. 89 N, O) ................................................................................................... 5
– Cocoon without anterior open loops (Fig. 88 G); gill with 6-12 thin branches .......... 8
5(4) Gill with 7 thick and short basal branches with 100 to 250 thin, short filaments
(Fig. 89 O) ........................................................................................... Simulium trombetense
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 373

– Gill with 14-23 non arborescent branches, without thin filaments ......................... 6
6(5) Gill with 20 relatively thick branches emerging from enlarged short common
trunk (Fig. 89 N) ...................................................................................... Simulium suarezi
– Gill with 10-23 thin branches basally emerging from primary, secondary, tertiary
or more subdivided branches .......................................................................................... 7
7(6) Gill branch length changing gradually from top to bottom, dorsal branches
shorter than ventral branches (Fig. 89 L); in frontal view, filaments form rosette
basally; frontoclypeus and thorax with abundant tubercles and trichomes with
robust basal portion (Fig. 88 V) ...................................................... Simulium maroniense
– Gill branches all about same length (Fig. 89 M); in frontal view not forming
rosette basally; frontoclypeus and thorax with sparse tubercles and trichomes with
basal portion not robust ..................................................................... Simulium rorotaense
8(4) Gill with 10-12 filaments .................................................................................................. 9
– Gill with 6-8 filaments .................................................................................................... 11
9(8) Frontoclypeus without tubercles except for small patch on base; frontoclypeal
trichomes, with 3-6 branches; gills with 10 filaments .................... Simulium ignescens
– Frontoclypeus with regularly spaced tubercles; frontoclypeal trichomes with 1-2
branches (Fig. 88 Q); gills with 10-12 filaments (Fig. 89 K) ................................... 10
10(9) Frontoclypeus and thorax with few tubercles; gill about as long as cocoon length;
gill with 10-12 filaments (Fig. 89 K) ............................................ Simulium kabanayense
– Frontoclypeus and thorax with abundant tubercles (Fig. 88 Q); gill shorter than
cocoon length (Fig. 88 G); gill with 10 filaments ................................. Simulium tunja
11(8) Gill with 6 branches ........................................................................................................ 12
– Gill with 8 branches ........................................................................................................ 17
12(11) Cocoon whitish with enlarged anterior margin and with dorsal longitudinal ridge
(Fig. 88 H) .......................................................................................................................... 13
– Cocoon not light colored, without enlarged anterior margin, without dorsal ridge
.............................................................................................................................................. 15
13(12) Cephalic and thoracic trichomes curled at base (Fig. 88 Y); gill branches very open
(Fig. 89 F) ..............................................................................................Simulium anaimense
– Cephalic and thoracic trichomes straight at base (similar to Fig. 88 P); gill
branches not very open .................................................................................................. 14
14(13) Cocoon anterior edge strongly oblique ..................................... Simulium jaimeramirezi
– Cocoon anterior edge slightly oblique ............................................... Simulium cormonsi
15(12) Gill about same length as cocoon length (Figs. 88 A, E) ........................................ 16
– Gill shorter than cocoon length ......................................................... Simulium gabaldoni
16(15) Cocoon without anteroventral projection (Fig. 88 A); bifurcation of tertiary
branches near base (Fig. 89 A) ........................................................ Simulium arcabucense
– Cocoon with anteroventral projection (Fig. 88 E), bifurcation of median and
ventral tertiary branches not near base (Fig. 89 B) ......................... Simulium pautense
17(11) Gill terminal branches emerging very close to base, very short primary and
secondary branches (Fig. 89 J) .......................................................... Simulium perflavum
– Gill terminal branches not emerging close to base, longer primary and secondary
branches (Figs. 89 C, D) ................................................................................................. 18
374 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

18(17) Frontoclypeal and thoracic tubercles longer than wide and acuminate distally (Fig.
88 Z) ........................................................................................................ Simulium albanense
– Frontoclypeal and thoracic tubercles shorter than wide and blunt distally ......... 19
19(18) Cocoon usually soft and thick; threads not very evident, with felt-like or spongy
texture, generally with thickened anterior border and dorsal longitudinal ridge
(Figs. 88 B, I, J) ................................................................................................................. 20
– Cocoon compact; threads thin, evident, anterior border can be reinforced but not
thickened and without dorsal ridge (Fig. 88 K) ......................................................... 25
20(21) Frontoclypeus and thorax with 35-40 branches (Fig. 88 M); gill with minute
tubercles giving wrinkled appearance ........................................... Simulium bicoloratum
– Frontoclypeus and thorax with 3-8 branches; gill branches without minute
tubercles, having smooth appearance .......................................................................... 21
21(20) Gill branches relatively open at base (Fig. 89 H) ...................................................... 22
– Gill branches relatively closed at base (Fig. 89 G) .................................................... 23
22(21) Frontoclypeus and thorax with sparse tubercles .......................................................
............................................................................................ (in part)......Simulium lutzianum
– Frontoclypeus and thorax with abundant tubercles (Fig. 88 T) ...........................
.................................................................................................................... Simulium romanai
23(21) Frontoclypeus and thorax with sparse tubercles (Fig. 88 R) ..... Simulium lutzianum
– Frontoclypeus and thorax with moderate (Fig. 88 O) to abundant tubercles (Fig.
88 S) .................................................................................................................................... 24
24(23) Frontoclypeus and thorax with sparse tubercles (Fig. 88 O); trichomes with 3-7
branches ...................................................................................................... Simulium pifanoi
– Frontoclypeus and thorax with abundant tubercles (Fig. 88 S); trichomes with 7-
12 branches ......................................................................................... Simulium mayuchuspi
25(19) Tertiary branches emerging beyond midlength of gill (Fig. 89 D) ...........................
.................................................................................................................Simulium roquemayu
– Tertiary branches emerging at basal 1/3 of gill (Figs. 89 E, I) .............................. 26
26(25) Frontoclypeal and thoracic trichomes with 2-4 branches relatively elongated (Fig.
88 X) .................................................................................................. Simulium shewellianum
– Frontoclypeal and thoracic trichomes with variable number of not so elongated
branches (Fig. 88 W) ........................................................................................................ 27
27(26) Frontoclypeus and thorax with sparse tubercles (Figs. 88 P, W) ........................... 28
– Frontoclypeus and thorax with abundant tubercles (Fig. 88 U) ............................ 29
28(27) Cocoon prolonged dorsally and basally; gill length 2.7-3.5 mm; tubercles of
frontoclypeus absent or sparse (Fig. 88 P) ..................................Simulium rubiginosum
– Cocoon without anterior projections; gill length 2.0-2.2 mm; tubercles more
abundant than described above (Fig. 88 W) ........................................ Simulium dinellii
29(27) Cocoon with open weave ................................................................ Simulium nuneztovari
– Cocoon compact, without open weave (Fig. 88 K) .................................................. 30
30(29) Thoracic trichomes with 4-9 branches; arrangement of primary and secondary
branches not close to base (Fig. 89 I) ............................................ Simulium wolffhuegeli
– Thoracic trichomes with 2-6 branches; arrangement of primary and secondary
branches close to base ..................................... Simulium ochraceum, Simulium antillarum
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 375

Larvae
Unknown: Simulium antonii, S. cormonsi, S. gabaldoni, S. jaimeramirezi, S. nuneztovari, S.
roquemayu, S. tolimaense.

1 Medial antennal segment similar or smaller than proximal one (Fig. 90 O); anal
ring with 84-100 rows of hooks; cephalic apotome darkened on basal border and
with median longitudinal band (Fig. 90 H) .................................................................. 2
– Medial antennal segment usually longer than proximal one (Fig. 90 N); anal ring,
usually with fewer than 84 rows of hooks; cephalic apotome with different
ornamentation ..................................................................................................................... 5
2(1) Ratio of length of postgenal bridge/hypostomium = 0.1-0.2 (Fig. 90 K) ..............
.............................................................................. Simulium maroniense, Simulium rorotaense
– Ratio of length of postgenal bridge/hypostomium = 0.3-0.4 ................................. 3
3(2) Anal ring with about 84 rows of hooks; ratio of length of postgenal bridge/
hypostomium = 0.4 ......................................................................... Simulium kabanayense
– Anal ring with 96-110 rows of hooks; ratio of length of postgenal bridge/
hypostomium = 0.3 ........................................................................................................... 4
4(3) Ratio of antennal segments: proximal, medial and distal = 1:0.9-1.0:1.1-1.3;
cephalic fan with 45-50 rays ............................................................ Simulium trombetense
– Ratio of antennal segments: proximal, medial and distal = 1:0.8:1.2; cephalic fan
with 40 rays ............................................................................................... Simulium suarezi
5(1) Anal ring with over 80 rows of hooks; body length usually over 6.0 mm ............ 6
– Anal ring with less than 80 rows of hooks; body length usually less than 6.0 mm
.............................................................................................................................................. 11
6(5) Ratio of length of postgenal bridge/hypostomium = 0.3-0.4 (Fig. 90 J); anal ring
with 83-85 rows of hooks ................................................................................................ 7
– Ratio of length of postgenal bridge/hypostomium = 0.2; anal ring with variable
number of rows .................................................................................................................. 8
7(6) Body length 5.7 mm; cephalic fan with 35 rays ........................... Simulium furcillatum
– Body length 6.0-6.7 mm; cephalic fan with 45-50 rays ................................................
......................................................................... Simulium rubiginosum, Simulium bicoloratum
8(6) Anal ring with 86-88 rows with 17-20 hooks each ......................... Simulium ignescens
– Anal ring with 77-85 rows and 12-16 hooks each ...................................................... 9
9(8) Cephalic fan with about 30-32 rays; rectal papillae with 8-11 diverticula on each
lobe; anal sclerite without hairs .............................................................. Simulium pifanoi
– Cephalic fan with about 37-40 rays; rectal papillae with 10-16 diverticula on each
lobe (Fig. 90 T); anal sclerite with hairs ...................................................................... 10
10(9) Medial segment of antenna longer than distal one (Fig. 90 N); ratio of antennal
segments: proximal, medial and distal =1:1.7:1.4-1.6; body length 6.5 mm ...........
.................................................................................................................. Simulium perflavum
– Medial segment of antenna shorter than distal one (Fig. 90 M); ratio of antennal
segments: proximal, medial and distal = 1:1.7:2.5; body length 5.8 mm .................
................................................................................................................ Simulium bicornutum
376 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

11(5) Ratio of length of postgenal bridge/hypostomium 0.2-0.3; cephalic apotome


darkened subbasally with lighter submedian area (Fig. 90 F) ........ Simulium wolffhuegeli
– Ratio of length of postgenal bridge/hypostomium less than 0.10-0.25; cephalic
apotome darkened mediobasally with lighter subbasal window (Figs. 90 B, D, E)
.............................................................................................................................................. 12
12(11) Anal ring with 70-79 rows of hooks ............................................................................ 13
– Anal ring with fewer than 69 rows of hooks ............................................................. 17
13(12) Cephalic fan with 42-48 rays ............................................................... Simulium pautense
– Cephalic fan with 27-39 rays ......................................................................................... 14
14(13) Cephalic fan with 27-35 rays ......................................................................................... 15
– Cephalic fan with 36-39 rays ......................................................................................... 16
15(14) Body length 4.8-5.2 mm; postgenal cleft subtrianglular (Fig. 90 I); cephalic apotome
without darkened areas and with very few hairs (Fig. 90 C) .......... Simulium arcabucense
– Body length 6.0 mm; postgenal cleft dome shape (Fig. 90 J); cephalic apotome slightly
darkened on basal border and with abundant hairs (Fig. 90 D) ....... Simulium albanense
16(14) Body length about 6.0 mm ..................................................................... Simulium dinellii
– Body length about 4.6 mm ............................ Simulium ochraceum, Simulium antillarum
17(12) Cephalic fan with about 47 rays; anal ring with 52 rows of hooks ...................... 18
– Cephalic fan with about 35 rays; anal ring with 55-74 rows of hooks ....................
................................................................................................................. Simulium lutzianum
18(17) Anal sclerite with trichomes and scales (Fig. 90 R) ............................. Simulium tunja
– Anal sclerite with only trichomes, without scales ..................................................... 19
19(18) Body length about 6.0 mm .................................................................. Simulium romanai
– Body length 4.3-4.7 mm ................................................................................................. 20
20(19) Body integument with foliaceus trichomes (Fig. 90 P); ratio of length of postgenal
bridge/hypostomium = 0.2 ...............................................................Simulium anaimense
– Body integument without foliaceus trichomes; ratio of length of postgenal
bridge/hypostomium = 0.1 ..................... Simulium mayuchuspi, Simulium shewellianum

CLAVES DE ESPECIES DE SIMULIUM (ECTEMNASPIS)


Hembras
1 Escudo amarillento con área mediana y 1+1 áreas sublaterales anaranjadas hasta a
veces castaño negruzco (Figs. 84 A, B); metanoto predominantemente castaño
negruzco o totalmente gris negruzco ............................................................................ 2
– Escudo totalmente amarillento; metanoto predominantemente amarillento ...... 16
2(1) Cibario sin dientes y con borde fino esclerotizado (Figs. 84 I, J) .............................
....................................................................................... grupo sp. BICOLORATUM......3
– Cibario dentado con área mediana de borde deprimido, cubierto con tubérculos y
lateralmente con dientes aguzados (Figs. 84 K-P) ......... grupo sp. ROMANAI......11
3(2) Sector basal de R sin pelos (Fig. 85 A) .......................................................................... 4
– Sector basal de R con pelos ............................................................................................. 8
4(3) Escudo naranja amarillento, sin color castaño (Fig. 84 A) ........................................ 5
– Escudo amarillento a castaño oscuro sobre el disco .................................................. 6
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 377

5(4) Largo ala 3,0 mm; tergitos abdominales I-IV amarillentos; largo de paraprocto
aproximadamente 1/4 del ancho del cerco en la base ...................... Simulium antonii
– Largo ala 2,4-2,6 mm; tergitos abdominales I-III amarillentos; paraprocto tan largo
como el ancho del circo en la base (Figs. 85 G, H) ......Simulium pautense, Simulium pifanoi
6(4) Largo ala 2,8-2,9 mm; paraprocto más largo que ancho en la base (Fig. 85 I) ......
......................................................................................................................... Simulium tunja
– Largo ala 2,4-2,7 mm; paraprocto aproximadamente similar al ancho en la base
(Fig. 85 E) ............................................................................................................................ 7
7(6) Tergitos abdominales I-IV totalmente amarillo grisáceo .............................................
......................................... Simulium bicornutum, Simulium furcillatum, Simulium tolimaense
– Tergitos abdominales I-IV amarillo con placas tergales III-IV castaño sobre el
borde posterior ................................................................................... Simulium arcabucense
8(7) Escutelo y tergitos abdominales I-IV castaño amarillento ........................................ 9
– Escutelo naranja amarillento o naranja verdoso; tergitos abdominales II-IV
amarillo a castaño amarillento ....................................................................................... 10
9(8) Longitud vesícula sensorial del palpo maxilar menos de 1/3 de la longitud del
palpómero (Fig. 84 Y) ........................................................................... Simulium ignescens
– Longitud vesícula sensorial del palpo maxilar menos de 1/2 de la longitud del
palpómero (Fig. 84 X) ......................................................................Simulium rubiginosum
10(8) Escudo totalmente amarillo, tergitos abdominales I-III amarillentos, IV castaño
sobre placa tergal; longitud de paraprocto aproximadamente tan largo como
ancho en la base (Fig. 85 F) ...............................................................Simuliun roquemayu
– Escudo generalmente amarillo con abundante negro (Fig. 84 B), con área central
desde castaño a anaranjado; tergitos abdominales I-II amarillo-anaranjado, III- IV
castaño negruzco; longitud del paraprocto aproximadamente 1/3 mas largo que el
ancho basal tan largo como ancho en la basal ...............................................................
....................................... Simuliun bicoloratum, Simulium cormonsi, Simulium jaimiramirezi
11(2) Porción basal de R glabro (Fig. 85 A); porción basal del cibario con tubérculos
redondeados medianamente, y escasos y diminutas espículas lateralmente (Fig. 84
K) .............................................................................................................. Simulium albanense
– Porción basal de R con pelos; porción basal del cibario con tubérculos y pequeñas
placas de diferente forma medianamente, y generalmente con dientes abundantes
y pequeños como espinas lateralmente (Figs. 84 L-P) ............................................. 12
12(11) Escudo, escudete, metanoto y abdomen negruzcos (Fig. 84 C); uña sin diente
subbasal (Fig. 85 B) ........................................................................... Simulium mayuchuspi
– Escudo, escudete, metanoto y abdomen amarillentos en proporción variable; uña
con diente subbasal (Fig. 85 C) ..................................................................................... 13
13(12) Escudo naranja; área mediana del cibario con tubérculos subovoidales (Fig. 84 L)
...................................................................................................................Simulium gabaldoni
– Escudo naranja amarillento pero frecuentemente con manchas castaño negruzcas
medialmente y 1+1 áreas sublaterales; cibario con pequeñas placas subcuadradas a
subesferoidales sobre el área mediana (Figs. 84 M-P) ............................................. 14
14(13) Tergitos abdominales I-II (III) amarillentos ................................. Simulium lutzianum
– Tergitos abdominales I amarillento .............................................................................. 15
378 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

15(14) Tergitos abdominales II-IV amarillentos a castaño amarillentos, con la porción


mediana de las placas tergales del III-V castaño oscuro (Fig. 84 G); Sc con 0-3
pelos ...................................................................................................... Simulium wolffhuegeli
– Tergitos abdominales II-IV castaño negruzco a negro; Sc con 6-9 pelos ...............
.................................................................................................................... Simulium romanai
16(1) Tergitos abdominales IV-VIII generalmente castaño negruzco con placa tergal
amarillenta; área mediana del cibario con 1+1 elevación submediana ancha, corta
y cubiertas por fuertes dientes seguidos por pequeños dientes (Figs. 84 Q, R);
paraprocto apicalmente desde aguzado (Figs. 85 M, N) a moderadamente romo
(Fig. 85 K) ..................................................................... grupo sp. PERFLAVUM......17
– Tergitos abdominales I-IV amarillentos (Fig. 84 H); tergitos V-VII desde amarillo
a castaño amarillento o amarillo a gris claro; área mediana del cibario con 1+1
prolongaciones submedianas alargadas, flanqueando un área mediana deprimida
bien esclerotizada, superficialmente desde lisa a rugosa (Figs. 84 T-W); paraprocto
romo apicalmente (Figs. 85 O-Q) .................................. grupo sp. DINELLII......21
17(16) Patas y tergitos abdominales totalmente amarillentos; triángulo fronto-ocular más
alto que ancho; paraprocto apicalmente romo (Fig. 85 K); largo ala 2,5-2,7 mm
.................................................................................................................. Simulium perflavum
– Tergitos abdominales presentando por lo menos IV-VII castaño negruzco; paraprocto
apicalmente generalmente aguzado (Figs. 85 M, N); largo ala 2,2-2,3 mm ............... 18
18(17) Metanoto amarillo a castaño amarillento .................................................................... 19
– Metanoto negruzco .......................................................................................................... 20
19(18) Vesícula sensorial del palpo maxilar alrededor de 1/3-1/4 del largo del palpómero
(Fig. 84 b) ........................................................ Simulium maroniense, Simulium trombetense
– Vesícula sensorial del palpo maxilar alrededor de 1/2 del largo del palpómero
(Fig. 84 c) ............................................................................................... Simulium rorotaense
20(18) Tergito abdominal III amarillento; relación del largo/ancho en la base del
paraprocto = 1,3 (Fig. 85 L) .......................................................... Simulium kabanayense
– Tergito abdominal III negruzco; relación del largo/ancho en la base del
paraprocto = 1,7 (Fig. 85 N) ................................................................. Simulium suarezi
21(16) Escudo homogeneamente pigmentado sin bandas plateadas submedianas ........ 22
– Escudo con 1+1 bandas plateadas submedianas cortas (Fig. 84 E) ..................... 23
22(21) Vesícula sensorial del palpo maxilar más corta que lo mitad del largo del
palpómero (Fig. 84 A); cibario con 1+1 prolongación submediana aguzada (Fig.
84 U); largo ala 1,9 mm ................................................................. Simulium shewellianum
– Vesícula sensorial del palpo maxilar más larga que la mitad del largo del palpómero;
cibario con 1+1 prolongación submediana roma (Fig. 84 T); largo ala 2,1 mm .......
.................................................................................. Simulium dinellii, Simulium nuneztovari
23(21) Largo del ala 2,2-2,3 mm; escudo amarillo opaco con 1+1 cuñas plateadas conectadas
con el área plateada anterolateral (Fig. 84 E); área mediana deprimida del cibario rugosa
o con pequeñas espinas (Fig. 84 V); metanoto castaño oscuro ......... Simulium ochraceum
– Largo del ala 1,9 mm; escudo amarillo rojizo con 1+1 cuñas no bien conectadas
con el área plateada anterolateral (Fig. 84 D); área mediana deprimida del cibario
lisa, sin espinas, no rugosa y bordeada basalmente por un grueso borde (Fig. 84
W); metanoto amarillento .................................................................. Simulium antillarum
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 379

Machos
Desconocidos: Simulium anaimense, S. antonii, S. roquemayu, S. tolimaense.

1 Escudo, escudete, metanoto y abdomen gris oscuro; escudo con el área del cuarto
posterior negra (Fig. 86 C); placa ventral con una fuerte contracción lateral (Fig.
87 G) .................................................................................................... Simulium mayuchuspi
– Escudo y abdomen desde todo amarillo o reducido este pigmento al borde del
escudo y al tergito abdominal I (Figs. 86 A, D); placa ventral con o sin contracción
lateral (Fig. 87 B, H) .......................................................................................................... 2
2(1) Escudo naranja amarillento pasando al negro en diferente proporción desde
pequeñas manchas castaño oscuro a negruzco sobre las áreas medianas y
sublaterales del disco (Fig. 86 B); metanoto negruzco .............................................. 3
– Escudo amarillento; metanoto amarillo castaño a castaño oscuro ........................ 14
3(2) Gonostilo aproximadamente 1/2 a 2/3 del largo del gonocoxito, subcuadrado
con borde distal truncado (Figs. 86 M, O); placa ventral ancha con una relación de
largo /ancho = 3.0-4.0 (Figs. 87 E, I) ........................................................................... 4
– Gonostilo aproximadamente 3/4 o más del gonocoxito, subcilíndrico con borde
distal aguzado (Figs. 86 K, N); placa ventral no tan ancha como en el dilema
anterior con un índice de largo/ancho = 1,1-1,9 (Figs. 87 D, H) ........................... 5
4(3) Escudo negro, reducida el área amarilla lateralmente a los ángulos pronotales y al
borde posterior angosto; escudete castaño oscuro y abdomen negruzco; gonostilo
con 6-8 espolones terminales (Fig. 86 O) ..................................... Simulium wolffhuegeli
– Escudo amarillento con pigmentación negra sobre el disco (como en Simulium
romanai Fig. 86 D), en algunos especimenes; tergito abdominal I-III amarillento,
gonostilo con un espolón apical (Fig. 86 M) ................................... Simulium albanense
5(3) Costados laterales de la placa ventral con una fuerte concavidad (Fig. 87 H) ..... 6
– Costados laterales de la placa ventral con una suave concavidad (Figs. 87 A, C, D) ..... 7
6(5) Relación largo/ ancho del basitarso posterior = 4,8-5,1; escudo con frecuencia
mayormente negro como Figura 86 D .............................................. Simulium romanai
– Relación largo/ancho del basitarso posterior = 6,5-7,6; escudo naranja
amarillento ............................................................................................ Simulium lutzianum
7(5) Sector basal de R con pelos ............................................................................................. 8
– Sector basal de R glabro ................................................................................................... 9
8(7) Tergitos abdominales I-IV castaño amarillento, los siguientes tergitos negro ........
.............................................................................. Simulium rubiginosum, Simulium ignescens
– Tergitos abdominales todos negros ..................................................................................
.................................................. Simulium bicoloratum, S, cormonsi, Simulium jaimeramirezi
9(7) Largo ala 2,9-3,0 mm; tergitos abdominales I-III o IV castaño amarillento a
amarillo (Fig. 86 F); placa ventral con escasa concavidad lateral .......................... 10
– Largo ala 2,5 mm; tergitos abdominales I-II ó I-III amarillentos, los siguientes tergitos
negruzcos, placa ventral lateralmente convexa o con suave concavidad ................... 11
10(9) Tergitos abdominales I-IV amarillo verdoso, con pequeña área castaño oscuro
sobre el tergito IV (Fig. 86 F); relación largo/ancho del basitarso posterior = 3,5
................................................................................................................ Simulium bicornutum
380 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

– Tergitos abdominales I-III amarillento, con una mancha grande sobre tergito IV
castaño oscuro (Fig. 86 G); relación largo/ancho del basitarso posterior = 4,0 ...
.................................................................................................................Simulium furcillatum
11(9) Placa ventral convexa lateralmente (Fig. 87 B); gonostilo más corto que el
gonocoxito (Figs. 86 I - J) .............................................................................................. 12
– Placa ventral no convexa lateralmente (Figs. 87 A, D); gonostilo aproximadamente
del mismo largo que el gonocoxito (Figs. 86 H, L) .................................................. 13
12(11) Escudo con 1+1 banda negruzca lateral alongada y una mediana en forma de T en
el tercio anterior (Fig. 86 B) ................................................................. Simulium pautense
– Escudo con 1+1 banda castaña corta lateral ...................................... Simulium pifanoi
13(11) Relación largo/ancho del basitarso posterior = 4,3; largo del ala 2,7 mm; tergito
abdominal III amarillento con la placa tergal castaño a negro ......... Simulium tunja
– Relación largo/ancho del basitarso posterior = 7,5; largo del ala 2,5 mm; tergito
abdominal III todo amarillo con placa tergal castaño ............... Simulium arcabucense
14(2) Placa ventral con fuerte concavidad lateral (Fig. 87 F); tergitos abdominales
negros ......................................................................................................Simulium gabaldoni
– Placa ventral generalmente sin fuerte concavidad lateral (Fig. 87 O); tergitos
abdominales amarillos o castaño rojizo amarillentos ............................................... 15
15(14) Escudo con cortas bandas plateadas submedianas anteriores (Fig. 86 E); placa
ventral lateralmente convexa (Fig. 87 O) .................................................................... 16
– Escudo con ornamentación homogénea sin bandas; placa ventral generalmente
con una pequeña concavidad lateral (Figs. 87 J-N) .................................................. 17
16(15) Escudo naranja amarillento, con 1+1 bandas plateadas submedianas en el tercio
anterior ................................................................................................... Simulium ochraceum
– Escudo castaño rojizo amarillento con 1+1 bandas plateadas en la mitad anterior
o sobrepasandola ................................................................................. Simulium antillarum
17(15) Placa ventral con porción distal haciéndose angosta suavemente (Fig. 87N);
gonostilo más corto que el gonocoxito (Fig. 86 S) ................................................... 18
– Placa ventral con porción distal volviéndose abruptamente angosta después de la
mitad de su largo (Figs. 87 K-M); gonostilo aproximadamente del mismo largo o
más largo que el gonocoxito (Figs. 86 P-R) .......... grupo sp. PERFLAVUM......20
18(17) Abdomen castaño negruzco; largo ala 1,8 mm ........................ Simulium shewellianum
– Abdomen amarillo rojizo a amarillo verdoso; largo del ala 2,0-2,5 mm .............. 19
19(18) Abdomen amarillo rojizo; largo ala 2,5 mm ........................................ Simulium dinellii
– Abdomen amarillo verdoso; largo ala 2,0 mm ............................ Simulium nuneztovari
20(17) Metanoto y abdomen amarillentos; gonostilo más largo que el gonocoxito;
longitud ala 2,6-2,7 mm ...................................................................... Simulium perflavum
– Metanoto y abdomen amarillentos; gonostilo no más largo que el gonocoxito;
longitud ala 1,8-2,6 mm .................................................................................................. 21
21(20) Todos los tergitos abdominales castaño a castaño negruzco; largo ala 2,0-2,1 mm
................................................................................................................ Simulium trombetense
– Tergitos abdominales I o II amarillo o castaño amarillento, restantes tergitos
castaño negruzco; largo ala 1,8-2,6 mm ...................................................................... 22
22(21) Tergito abdominal I castaño o castaño amarillento, restantes castaño oscuro .......
............................................................................... Simulium suarezi, Simulium kabanayense
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 381

– Tergito abdominal I-II castaño amarillento, restantes castaño oscuro .....................


.............................................................................. Simulium maroniense, Simulium rorotaense

Pupas
Desconocida: Simulium antonii.

1 Branquia compuesta por 2-5 ramas; capullo con prominente prolongación antero
dorsal (Figs. 88 C, D, F) .................................................................................................... 2
– Branquia compuesta por 6-250 filamentos; capullo sin prominente prolongación
anterior-dorsal ..................................................................................................................... 4
2(1) Ramas relativamente finas (Fig. 88 F) .............................................. Simulium tolimaense
– Branquia compuesta por 2-4 ramas muy gruesas ....................................................... 3
3(2) Branquia compuesta por 2 ramas similares con una pequeña protuberancia sobre
cada una (Fig. 88 C) ........................................................................... Simulium bicornutum
– Branquia con 4 ramas, 1 gruesa inferior y 3 finas sobrepuestas a la cabeza (Fig. 88
D) ............................................................................................................Simulium furcillatum
4(1) Capullo con aberturas anteriores (Fig. 88 L); branquias relativamente gruesas en
número de 17-250 (Figs. 89 N, O ) ................................................................................ 5
– Capullo sin aberturas anteriores (Fig. 88 G); ramas de la branquia finas y en
número de 6-12 .................................................................................................................. 8
5(4) Branquias con 7 ramas basales gruesas, y finos y cortos en número superior a 100
y hasta 250 (Fig. 89 O) ...................................................................... Simulium trombetense
– Branquias con 14-23 ramas no arborescentes, sin filamentos finos ....................... 6
6(5) Ramas de las branquias relativamente gruesas en número de 20, emergiendo de un
corto y grueso tallo común (Fig. 89 N) .............................................. Simulium suarezi
– Ramas de las branquias no gruesas basalmente en número de 10-23 emergiendo
desde ramas primarias, secundarias y terciarias o más ramas subdivididas ........... 7
7(6) Ramas de las branquias cambiando gradualmente de tamaño desde arriba hacia abajo,
siendo las ramas dorsales más cortas que las ventrales (Fig. 89 L); en vista frontal los
filamentos forman una roseta basalmente; frontoclípeo y tórax con abundantes
tubérculos, y tricomas con porción basal robusta (Fig. 88 V) .......... Simulium maroniense
– Ramas branquiales todas del mismo largo (Fig. 89 M); en vista frontal los
filamentosa no forman una roseta, frontoclípeo y tórax con moderado número de
tubérculos, y tricomas con la porción basal no robusta .............. Simulium rorotaense
8(4) Branquias con 10-12 filamentos ..................................................................................... 9
– Branquias con 6-8 filamentos ........................................................................................ 11
9(8) Frontoclípeo sin tubérculos, excepto un pequeño grupo sobre la base de la frente; tricomas
del frontoclípeo con 3-6 ramas; branquias con 10 filamentos ............... Simulium ignescens
– Frontoclípeo con tubérculos regularmente distribuidos; tricomas del frontoclípeo
con 1-2 ramas (Fig. 88 Q); branquias con 10-12 filamentos (Fig. 89 K) ............. 10
10(9) Frontoclípeo y tórax con escasos tubérculos; branquias aproximadamente tan
largas como el largo del capullo; branquias con 10-12 filamentos (Fig. 89 K) ......
.............................................................................................................. Simulium kabanayense
– Frontoclípeo y tórax con abundantes tubérculos (Fig. 88 Q); branquias más cortas que
el largo del capullo (Fig. 88 G); branquias con 10 filamentos ................ Simulium tunja
382 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

11(8) Branquia con 6 ramas ...................................................................................................... 12


– Branquia con 8 ramas ...................................................................................................... 17
12(11) Capullo blanquecino con el del capullo (Fig. 88 H) ................................................. 13
– Capullo no blanquecino, sin margen anterior engrosado ni carena dorsal .......... 15
13(12) Tricomas cefálicos y torácicos con base en forma de rulo (Fig. 88 Y); ramas de las
branquias bien abiertas (Fig. 89 F) ...................................................Simulium anaimense
– Tricomas cefálicos y torácicos con base recta (similar a Fig. 88 P); ramas de las
branquias no muy abiertas ............................................................................................. 14
14(13) Borde anterior del capullo fuertemente oblicuo ..................... Simulium jaimeramirezi
– Borde anterior del capullo escasamente oblicuo ............................. Simulium cormonsi
15(12) Largo de branquia aproximadamente igual que el del capullo (Figs. 88 A, E) .......... 16
– Largo de branquia más corto que el del capullo ............................ Simulium gabaldoni
16(15) Capullo sin prolongación anterior ventral (Fig. 88 A); bifurcación de las ramas
terciarias próximas a la base (Fig. 89 A) ........................................ Simulium arcabucense
– Capullo con prolongación anterior ventral (Fig. 88 E); bifurcación de las ramas
terciarias mediana y ventral no próximas a la base (Fig. 89 B) .... Simulium pautense
17(11) Ramas terminales de las branquias emergiendo muy próximas a la base,
mostrando ramas primarias y secundarias muy cortas (Fig. 89 J) .............................
.................................................................................................................. Simulium perflavum
– Ramas terminales de las branquias emergiendo no próximo a la base, mostrando
ramas primarias y secundarias largas (Figs. 89 C, D) ............................................... 18
18(17) Tubérculos del frontoclípeo y tórax más largos que anchos y aguzados distalmente
(Fig. 88 Z) ............................................................................................... Simulium albanense
– Tubérculos del frontoclípeo y tórax más cortos que anchos y romos distalmente
.............................................................................................................................................. 19
19(18) Capullo generalmente blando y grueso; fibras del tejido no bien evidentes con
aspecto de fieltro o esponjoso, generalmente con borde anterior engrosado y con
una carena dorsal longitudinal (Figs. 88 B, I, J) ......................................................... 20
– Capullo compacto, fino, fibras del tejido evidentes, borde anterior puede estar
reforzado pero no engrosado y sin una carena dorsal (Fig. 88 K) ........................ 25
20(19) Tricomas del frontoclípeo y tórax con 35-40 ramas (Fig. 88 M); superficie de las
ramas de las branquias con pequeños tubérculos dando apariencia rugosa ...........
............................................................................................................... Simulium bicoloratum
– Tricomas del frontoclípeo y tórax con 3-8 ramas; ramas de la branquia sin
tubérculos y con apariencia lisa ..................................................................................... 21
21(20) Ramas de las branquias relativamente abiertas en la base (Fig. 89 H) ................. 22
– Ramas de las branquias relativamente cerradas en la base (Fig. 89 G) ................ 23
22(21) Frontoclípeo y tórax con escasos tubérculos ........ (en parte)......Simulium lutzianum
– Frontoclípeo y tórax con abundantes tubérculos (Fig. 88 T) ....... Simulium romanai
23(21) Frontoclípeo y tórax con escasos tubérculos (Fig. 88 R) ........... Simulium lutzianum
– Frontoclípeo y tórax con medianamente (Fig. 88 O) a abundantes tubérculos (Fig.
88 S) .................................................................................................................................... 24
24(23) Frontoclípeo y tórax con escasos tubérculos (Fig. 88 O); tricomas con 3-7 ramas
....................................................................................................................... Simulium pifanoi
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 383

– Frontoclípeo y tórax con abundantes tubérculos (Fig. 88 S); tricomas con 7-12
ramas .................................................................................................... Simulium mayuchuspi
25(19) Ramas terciarias emergen sobrepasando la mitad del largo de la branquia (Fig. 89 D)
.................................................................................................................Simulium roquemayu
– Ramas terciarias emergen debajo de 1/3 de la distancia a la base (Figs. 89 E, I)
.............................................................................................................................................. 26
26(25) Tricomas del frontoclípeo y tórax con 2-4 ramas relativamente largos (Fig. 88 X)
............................................................................................................. Simulium shewellianum
– Tricomas del frontoclípeo y tórax con variable número de ramas y no tan
alargadas (Fig. 88 W) ....................................................................................................... 27
27(26) Frontoclípeo y tórax con escasos tubérculos (Figs. 88 P, W) ................................. 28
– Frontoclípeo y tórax con abundantes tubérculos (Fig. 88 U) ................................. 29
28(27) Capullo prolongado dorsal y basalmente; largo branquia 2,7-3,5 mm; tubérculos
del frontoclípeo ausentes o escasos (Fig. 88 P) ..........................Simulium rubiginosum
– Capullo sin prolongaciones anteriores; largo branquia 2,0-2,2 mm; tubérculos más
abundantes que lo mostrado en el dilema opuesto (Fig. 88 W) ...... Simulium dinellii
29(27) Capullo con fibras abiertas .............................................................. Simulium nuneztovari
– Capullo compacto sin fibras abiertas (Fig. 88 K) ...................................................... 30
30(29) Tricomas torácicos con 4-9 ramas; ramificación de las ramas primarias y
secundarias no próximas a la base (Fig. 89 I) .............................. Simulium wolffhuegeli
– Tricomas torácicos con 2-6 ramas; ramificación de las ramas primarias y
secundarias próximas a la base ...................... Simulium ochraceum, Simulium antillarum

Larvas
Desconocidas: Simulium antonii, S. cormonsi, S. gabaldoni, S. jaimeramirezi, S. nuneztovari, S.
roquemayu, S. tolimaense.

1 Artejo mediano de la antena similar o menor que el proximal (Fig. 90 O); anillo
anal con 84-100 hileras de ganchos; apotoma cefálico oscurecido en el borde basal
y una banda mediana longitudinal (Fig. 90 H) ............................................................. 2
– Artejo mediano de la antena generalmente más largo que el proximal (Fig. 90 N);
anillo anal generalmente con un número de hileras por debajo de 84; apotoma
cefálico con diferente ornamentación ........................................................................... 5
2(1) Relación del largo puente postgenal/hipostomio = 0,1-0,2 (Fig. 90 K) ..................
.............................................................................. Simulium maroniense, Simulium rorotaense
– Relación del largo puente postgenal/hipostomio = 0,3-0,4 ..................................... 3
3(2) Anillo anal con alrededor de 84 hileras de ganchos; relación del largo puente
postgenal/hipostomio = 0,4 .......................................................... Simulium kabanayense
– Anillo anal con 96-110 hileras de ganchos; relación del largo puente postgenal/
hipostomio = 0,3 ................................................................................................................ 4
4(3) Relación de los artejos antenales proximal, mediano y distal = 1:0,9-1,0:1,1-1,3;
abanico cefálico con 45-50 rayos .................................................... Simulium trombetense
– Relación de los artejos antenales proximal, mediano y distal = 1:0,8:1,2; abanico
cefálico con 40 rayos ............................................................................... Simulium suarezi
384 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

5(1) Anillo anal sobrepasando 80 hileras de ganchos; longitud del cuerpo sobre 6,0
mm ........................................................................................................................................ 6
– Anillo anal por debajo de 80 hileras de ganchos; longitud del cuerpo generalmente
por debajo de 6,0 mm ..................................................................................................... 11
6(5) Relación del largo puente postgenal/hipostomio =0,3-0,4 (Fig. 90 J); anillo anal
con 83-85 hileras de ganchos .......................................................................................... 7
– Relación del largo puente postgenal/hipostomio = 0,2; anillo anal con número
variable de hileras de ganchos ......................................................................................... 8
7(6) Longitud del cuerpo 5,7 mm; abanico cefálico con 35 rayos .............................
.................................................................................................................Simulium furcillatum
– Longitud del cuerpo 6,0-6,7 mm; abanico cefálico con 45-50 rayos ........................
......................................................................... Simulium rubiginosum, Simulium bicoloratum
8(6) Anillo anal con 86-88 hileras, con 17-20 ganchos cada una ......... Simulium ignescens
– Anillo anal con 77-85 hileras, con 12-16 ganchos cada una ..................................... 9
9(8) Abanico cefálico con alrededor de 30-32 rayos; papilas rectales con 8-11
divertículos en cada lóbulo; esclerito anal sin pelos .......................... Simulium pifanoi
– Abanico cefálico con alrededor de 37-40 rayos; papilas rectales con 10-16
divertículos en cada lóbulo (Fig. 90 T); esclerito anal con pelos ........................... 10
10(9) Artejo mediano de la antena más largo que el distal (Fig. 90 N); relación de los
artejos antenales: proximal, mediano y distal = 1:1,7:1,4-1,6; longitud cuerpo 6,5
mm .......................................................................................................... Simulium perflavum
– Artejo mediano de la antena más corto que el distal (Fig. 90 M); relación de los
artejos antenales: proximal, mediano y distal = 1:1,7:2,5; longitud del cuerpo 5,8
mm ........................................................................................................ Simulium bicornutum
11(5) Relación del largo puente postgenal/hipostomio 0,2-0,3; apotoma cefálico
oscurecido subbasalmente con área submediana más clara (Fig. 90 F) ...................
................................................................................................................ Simulium wolffhuegeli
– Relación del largo puente postgenal/hipostomio 0,10-0,25; apotoma cefálico
oscurecido medio basalmente con una ventana subbasal más clara (Figs. 90 B, D, E)
.............................................................................................................................................. 12
12(11) Anillo anal con 70-79 hileras de ganchos ................................................................... 13
– Anillo anal con por debajo de 69 hileras de ganchos .............................................. 17
13(12) Abanico cefálico con 42-48 rayos ....................................................... Simulium pautense
– Abanico cefálico con 27-39 rayos ................................................................................. 14
14(13) Abanico cefálico con 27-35 rayos ................................................................................. 15
– Abanico cefálico con 36-39 rayos ................................................................................. 16
15(14) Longitud cuerpo 4,8-5,2 mm; hendidura postgenal subtriangular (Fig. 90 I);
apotoma cefálico sin áreas oscurecidas y escasos pelos (Fig. 90 C) ..........................
................................................................................................................Simulium arcabucense
– Longitud cuerpo 6,0 mm; hendidura postgenal en forma de domo (Fig. 90 J);
apotoma cefálico levemente oscurecido sobre el borde basal y con abundantes
pelos (Fig. 90 D) .................................................................................... Simulium albanense
16(14) Longitud cuerpo alrededor de 6,0 mm ................................................ Simulium dinellii
– Longitud cuerpo alrededor de 4,6 mm ....... Simulium ochraceum, Simulium antillarum
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 385

17(12) Abanico cefálico con alrededor de 47 rayos; anillo anal con 52 hileras de ganchos
.............................................................................................................................................. 18
– Abanico cefálico con alrededor de 35 rayos; anillo anal con 55-74 hileras de
ganchos .................................................................................................. Simulium lutzianum
18(17) Esclerito anal con pelos y escamas (Fig. 90 R) ..................................... Simulium tunja
– Esclerito anal con solamente pelos, sin escamas ....................................................... 19
19(18) Largo del cuerpo 6,0 mm ..................................................................... Simulium romanai
– Largo del cuerpo 4,3-4,7 mm ........................................................................................ 20
20(19) Tegumento del cuerpo con tricomas foliáceos (Fig. 90 P); relación largo puente
postgenal/hipostomio 0,2 ..................................................................Simulium anaimense
– Tegumento del cuerpo sin tricomas foliáceos; relación largo puente postgenal/
hipostomio 0,1 ............................................ Simulium mayuchuspi, Simulium shewellianum

BICOLORATUM species group

Simulium (Ectemnaspis) rubiginosum (ENDERLEIN) (Figs. 84 I, X; 86 K; 87 C; 88 P; 89 E; 90 A.)


Chirostilbia rubiginosa ENDERLEIN, 1934b: 287.
Simulium rubiginosum: WYGODZINSKY, 1971: 19.
Simulium (Ectemnaspis) rubiginosum: COSCARÓN, 1984: 294-295; 1987: 28; 1990: 121; 1991: 248-249;
CROSSKEY & HOWARD, 1997: 30; 2004: 21.
Simulium ignescens: WYGODZINSKY, 1951b: 223 (part.); 1953: 325; COSCARÓN & WYGODZINSKY,
1960: 1124.
Female: Wing length 2.4-2.9 mm. Scutum from orange to mostly black, but always
bordered anterolaterally and posteromedially by yellowish brown or grayish brown band as
in S. romanai; scutellum yellowish, metanotum from yellowish brown to velvety black,
pleurae yellowish orange with episternum and sternites black pruinose. Abdomen with
tergites I-IV yellow to yellowish-brown, V-IX dark brown. Legs yellowish with dark portion
from light brown to totally black. Sensory vesicle of maxillary palp about half palpomere
length (Fig. 84 X). Basal portion of cibarium with sclerotized border and without teeth (Fig.
84 I). Wings with hair on basal sector of R. Paraproct 2/3 longer than wide.
Male: Wing length 2.4-2.7 mm. Thorax and abdomen with similar coloration as in
female but darker. Abdomen with posterior border of tergite I black, V-VIII velvety black.
Hind basitarsus with sides approximately parallel, ratio of length/width = 4.0. Gonostylus
slightly shorter than gonocoxite, subcylindrical, with parallel sides on basal 2/3, becoming
thin and curved on distal one third, with apical spur (Fig. 86 K). Ventral plate about as
long as wide, without or with slight lateral invagination in frontal view (Fig. 87 C).
Pupa: Cocoon slipper shaped, moderately compact weave with threads evident,
slightly projected dorsally and basally; length along base 3.6-4.3 mm, on dorsum 3.1-3.5 mm,
gills 2.7-3.5 mm. Frontoclypeus and thorax with few tubercles arranged homogeneously and
enlarged trichomes with 2-6 branches (Fig. 88 P). Gill with 8 filaments, grayish dark, not
bifurcated at base, open at several levels, with tendency to meet at apex (Fig. 89 E).
Larva: Maximum length 6.0 mm. Cephalic apotome pale to brownish, scarcely
darkened on basal half (Fig. 90 A). Antennae shorter than cephalic fan stem; ratio of
386 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

A B C D E

I
J

M F G
H

O N L

R
P
b
U W
S a

V X
T
c
Z

Figure 84. Female. A-H: Scutum and scutellum; A: S. (Ectemnaspis) pautense; B: S. (E.) bicoloratum; C:
S. (E.) mayuchuspi; D: S. (E.) antillarum; E: S. (E.) ochraceum, abdomen; F: S. (E.) bicoloratum; G: S. (E.)
wolffhuegeli; H: S. (E.) shewellianum. I-W Cibarium; I: S. (E.) rubiginosum; J: S. (E.) arcabucense; K: S. (E.)
albanense; L: S. (E.) gabaldoni; M-N: S. (E.) lutzianum, showing different disposition of tubercles and
spines ; O: S. (E.) romanai; P: S. (E.) wolffhuegeli; Q: S. (E.) kabanayense; R: S. (E.) perflavum; S: S. (E.)
suarezi; T: S. (E.) dinellii; U: S. (E.) shewellianum; V: S. (E.) ochraceum; W: S. (E.) antillarum. X-Y, a-c:
Sensory vesicle of palpus; X: S. (E.) rubiginosum; Y: S. (E.) ignescens; Z: S. (E.) dinellii; a: S. (E.)
shewellianum; b: S. (E.) maroniense; c: S. (E.) rorotaense.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 387

proximal, medial and distal segments = 1:1.3-1.4:1.5-1.6. Cephalic fan with about 45 rays.
Mandible without latero-mandibular process; with 1-3 marginal teeth, the first about 2
times larger than following teeth. Anal ring with 85 rows with about 15 hooks each. Rectal
papillae generally with 4-9 diverticula each, but some specimens with only 0-2 diverticula.
Distribution: Argentina: Buenos Aires, Jujuy, Mendoza, Salta, San Juan, Tucumán;
Bolivia: Cochabamba; Peru: Cuzco.
Bionomics: This species breeds in creeks of crystalline water, with fast flow (not
torrential) in mountain areas. Larvae and pupae occur on rocks and leaves. Female feeding
habits are unknown.
Discussion: The closest species is S. ignescens. Differences are explained under that
species.

Simulium (Ectemnaspis) ignescens ROUBAUD (Fig. 84 Y.)


Simulium (Ectemnaspis) ignescens ROUBAUD, 1906b: 107; WYGODZINSKY, 1951b: 223-229; 1971: 14-19;
LEON & WYGODZINSKY, 1953: 31; RAMÍREZ PÉREZ, 1971 : 315; 1983: 87.
Eusimulium ignescens: SOURCOUF & GONZALEZ RINCONES, 1911: 281; PINTO, 1931: 732.
Simulium (Ectemnaspis) ignescens COSCARÓN, 1984: 295-296; 1990: 121; CROSSKEY & HOWARD, 1997: 29;
2004: 21; COSCARÓN ARIAS, 2003: 195, 200, 205, 208, 215.
S. ignescens is similar to S. rubiginosum, and we include here only the differences.
Female: Sensory vesicle of maxillary palp shorter than half of palpomere length,
and with more abundant tubercles than in S. rubiginosum (Fig. 84 Y).
Male: Hind basitarsus wider at middle.
Pupa: Gill with 10 filamentous branches. Frontoclypeus with sparse tubercles
arranged on both sides near facial trichomes.
Larva: Distal antennal segment not longer than medial segment. Anal ring with 86-
88 rows with 17-20 hooks each. Rectal papillae with 15-20 diverticula on each.
Distribution: Bolivia: Chapare; Colombia: Cauca, Cundinamarca, Valle; Ecuador:
Chimborazo, Cotopaxi, Imbabura, Napo, Pastaza, Pichincha, Tungurahua; Peru: Cuzco;
Venezuela: Merida.
Bionomics: Simulium ignescens breeds in fast-flowing, small creeks of mountain areas
(1100-3050 m), with clear water, on submerged tree branches, aquatic vegetation and rocks.
Females bite humans in Ecuador and mules in Bolivia (LEÓN & WYGODZINSKY, 1953).
Discussion: The species is similar to S. rubiginosum, which can be differentiated by
the 8 pupal gill branches. Females can be differentiated by the length of the palpal sensory
vesicle, which is longer in S. rubiginosum.

Simulium (Ectemnaspis) roquemayu COSCARÓN (Figs. 85 F; 89 D.)


Simulium (Ectemnaspis) roquemayu COSCARÓN, 1984: 296-298; 1990: 122; 1991: 251; CROSSKEY &
HOWARD, 1997: 29; 2004: 21.
Female: Wing length 2.5 mm. Scutum and scutellum yellowish, metanotum black,
legs blackish; abdomen with tergites I-III yellowish, IV blackish-brown medially, V-IX
greenish-gray. Fronto-ocular triangle about as wide as high. Sensory vesicle of maxillary
388 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

palp about half palpomere length. Basal portion of cibarium scarcely sclerotized, more
intensely sclerotized laterally. Wings with hair on basal sector of R; Sc with about 22 setae.
Claws with subbasal tooth. eighth sternite well sclerotized medially, laterally with about 25
setae on each side. Paraproct subtriangular in shape, about as long as wide with rounded
distal border, abundant microtrichiae and without basal lobe (Fig. 85 F).
Male: Unknown.
Pupa: Cocoon slipper shaped with thick spongy weave, slightly elevated dorsally as
longitudinal carina. Length along base 2.6 mm, on dorsum 3.0 mm, gills 3.3 mm. Gills,
head, thorax and abdomen light brown. Frontoclypeus and thorax with numerous smooth
tubercles; large trichomes with 6-7 branches and curved base. Gills with 8 filamentous
branches, with tertiary branches bifurcated at more than half total gill length (Fig. 89 D).
Larva: Unknown.
Distribution: Bolivia: Cochabamba.
Bionomics: Unknown.
Discussion: Simulium antonii has coloration similar to that of S. roquemayu but can be
differentiated by the absence of hair on the basal sector of R. Also, S. bicornutum and S.
furcillataum have similar females, but the pupal gills with 2 thick branches readily
distinguish them. Simulium pifanoi shows similarities in scutum ornamentation, but
abdominal tergite III has a dark brown tergal plate, tergite V is velvety black and VI-IX are
blackish. The pupal gill branches are tightly arranged, forming a bundle.

Simulium (Ectemnaspis) pifanoi RAMÍREZ PÉREZ (Figs. 85 G; 86 J; 88 O; 89 C; 90 T.)


Simulium pifanoi RAMÍREZ PÉREZ, 1971: 346; 1983: 62.
Simulium (Ectemnaspis) pifanoi: COSCARÓN, 1984: 301; 1987: 28; 1990: 122-123; CROSSKEY & HOWARD,
1997: 29; 2004: 21.
Female: Wing length 2.4-2.6 mm. Scutum orange, with yellowish pronotal area and
scutellum; metanotum black. Abdomen with tergites I-II yellow; III-IV yellowish with
dark brown tergal plate; V velvety black; VI-IX shiny blackish. Legs black. Frons
convergent. Fronto-ocular triangle about as high as wide. Basal sector of R without hair.
Gonapophysis with internal border well sclerotized. Eighth sternite with about 30 setae on
each side; paraproct slightly longer than wide at base (Fig. 85 G).
Male: Wing length 2.5-2.6 mm. Coloration as in female but abdominal segments III-
IV dark-brown and scutum with dark stripes laterally. Gonostylus 3/4 of gonocoxite
length (Fig. 86 J); ventral plate without lateral constriction.
Pupa: Cocoon with basal anterior projection; weave with thin threads; length along
base 3.0-3.3 mm, on dorsum 2.6-2.8 mm, gills 3.2-3.5 mm. Frontoclypeus and thorax with
sparse tubercles and large trichomes with 3-7 branches (Fig. 88 O). Gill with 8 branches
with arrangement similar to that of S. roquemayu but arranged more open and with lower
bifurcation on tertiary branches (Fig. 89 C).
Larva: Maximum length 5.0-5.5 mm. Cephalic apotome slightly darkened on basal half.
Ratio of antennal segments: proximal, medial and distal = 1:1.3-1.4:1.6-1.7. Mandible with 1
latero-mandibular process. Postgenal bridge about 1/8 of hypostomium height. Cephalic
fan with 30-32 rays. Lateral sclerite of proleg with 26-30 teeth arranged in 7-8 groups. Anal
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 389

I
G

E K

H
J

L
N

Q
C B
C

Sc R
bR
A
P
Figure 85. Female. A: Basal section of wing of S. (Ectemnaspis) albanense (C = costa, Sc = subcosta,
R = radius, bR = basal sector of R). B-C: Claw; B: S. (E.) mayuchuspi; C: lutzianum. D-Q: Paraproct;
D: S. (E.) bicoloratum; E: S. (E.) furcillatum; F: S. (E.) roquemayu; G: S. (E.) pifanoi; H: S. (E.) pautense;
I: S. (E.) tunja; J: S. (E.) wolffhuegeli; K: S. (E.) perflavum; L: S. (E.) kabanayense; M: S. (E.) rorotaense; N:
S. (E.) suarezi; O: S. (E.) dinellii; P: S. (E.) shewellianum; Q: S. (E.) ochraceum.
390 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

sclerite without hair. Anal ring with 77-84 rows with 12-14 hooks each. Rectal papillae with
8-11 diverticula on each lobe (Fig. 90 T). Body integument without hair.
Distribution: Colombia: Cauca, Cundinamarca; Venezuela: Mérida.
Bionomics: Unknown.
Discussion: Simulium roquemayu is very similar and the differences were described
under that species. Simulium arcabucense is without hair on the basal sector of R and has a
similarly shaped paraproct but can be differentiated by the 6 branches of the pupal gill
and in some specimens by the dark brown area on the thoracic disc.

Simulium (Ectemnaspis) arcabucense COSCARÓN (Figs. 86 H; 87 A; 88 A; 89 A; 90 C.)


Simulium (Ectemnaspis) arcabucense COSCARÓN, 1990: 123-125; CROSSKEY & HOWARD, 1997: 29; 2004: 21.
Female: Wing length 2.7 mm. Scutum and scutellum yellowish; sometimes the first
with sublateral stripes and median anterior area dark brown. Metanotum black, pleurae
with silver pruinosity. Legs black with lighter portions on tibiae. Abdomen with tergites I-
II light yellow, darkened laterally; III-IV yellow with tergal plate brownish on posterior
border; V-IX blackish. Basal sector of R without hair. Eighth sternite with about 25 setae;
paraproct about as long as wide.
Male: Wing length 2.5 mm. Coloration as in female, sometimes with 1+1 lateral and
median frontal black stripes. Ratio of length/width of hind basitarsus 7.5. Gonostylus about
3/4 of gonocoxite length (Fig. 86 H). Ventral plate with slight lateral concavity (Fig. 87 A).
Pupa: Cocoon with thicker anterior rim, spongy weave and thin threads (Fig. 88 A);
length along base 2.9-3.0 mm, on dorsum 2.5-2.7 mm, gills 3.0-3.6 mm. Frontoclypeus
and thorax with numerous large tubercles arranged in irregular rows and wrinkled apically.
Cephalic and thoracic trichomes large, with thick base and 6-13 branches. Gill with 6
branches, emerging near base (Fig. 89 A).
Larva: Maximum length 4.8-5.2 mm. Cephalic apotome light, without special
ornamentation (Fig. 90 C). Ratio of antennal segments: proximal, medial and distal =
1:1.0-1.1:1.8-2.0. Postgenal bridge about 1/3-1/4 of hypostomium height. Mandible with
1 latero-mandibular process. Cephalic fan with about 27 rays. Lateral sclerite of thoracic
proleg with about 25 teeth in 8-9 groups. Anal sclerite without hair. Anal ring with about
70 rows with 12-13 hooks each. Body integument without hairs.
Distribution: Colombia: Boyacá.
Bionomics: Unknown.
Discussion: The most similar species is S. pautense, which can be differentiated by
the shorter gonostylus, thinner pupal cocoon, frontoclypeal and thoracic tubercles
uniformly arranged and apically smooth, gill branches bifurcating at a greater distance
from the base, darker cephalic apotome with negative spots and greater number of rays.
Simulium gabaldoni also has similar adult coloration, but the cibarium has the median area
well sclerotized with tubercles and lateral spines, the pupal tubercles are less elevated, and
the larval cephalic apotome has distinct spots. Simulium pifanoi, with similar adult
ornamentation, can be differentiated by the pupal gill with 8 branches. Simulium species A
and B LEWIS & LEE-PORTER (1964: 97-98) can be differentiated by the medially elevated
cibarium (sp. A) and the abdomen not darkened laterally and the longer paraproct (sp. B).
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 391

B A C D E

0.05 mm

H I
F G J

L K N
M

O P S
Q R

Figure 86. Male. A-E: Scutum and scutellum; A: S. (Ectemnaspis) furcillatum; B: S. (E.) pautense; C: S.
(E.) mayuchuspi; D: S. (E.) romanai; E: S. (E.) ochraceum. F-G: Abdomen; F: S. (E.) bicornutum; G: S.
(E.) furcillatum. H-S: Gonocoxite and gonostylus; H: S. (E.) arcabucense; I: S. (E.) pautense; J: S. (E.)
pifanoi; K: S. (E.) rubiginosum; L: S. (E.) tunja; M: S. (E.) albanense; N: S. (E.) romanai; O: S. (E.)
wolffhuegeli; P: S. (E.) kabanayense; Q: S. (E.) perflavum; R: S. (E.) rorotaense; S: S. (E.) shewellianum.
392 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Simulium (Ectemnaspis) pautense COSCARÓN & TAKAOKA


(Figs. 84 A; 85 H; 86 B, I; 87 B; 88 E; 89 B; 90 B.)
Simulium (Ectemnaspis) pautense COSCARÓN & TAKAOKA, 1989: 191; COSCARÓN, 1990: 125, CROSSKEY &
HOWARD, 1997: 29; 2004: 21; COSCARÓN ARIAS, 2003: 195, 200, 206, 208, 215, 219.
Female: Wing length 2.4-2.6 mm. Scutum orange, greenish orange laterally;
scutellum greenish yellow (Fig. 84 A); metanotum velvety black with 1+1 small grayish
patches. Pleura mostly pale yellow. Legs dark brown. Abdominal tergites I-IV yellow, with
tergal plates III and IV pale brown, tergal plates V-IX blackish brown; waxy, bright except
plate V velvety, bright. Basal portion of cibarium with reinforced border. Basal sector of
R bare. Eighth sternite with darker middle area and about 27 setae on each side. Paraproct
about as long as wide at base (Fig. 85 H).
Male: Wing length 2.3-2.5 mm. Scutum pale but yellow orange on top and with T-
shaped black stripe on anterior third and pair of black, broad longitudinal stripes
sublaterally (Fig. 86 B). Ratio of length/width of hind basitarsus = 5. Gonostylus about
4/5 of gonocoxite length (Fig. 86 I), ratio of length/wide = 2.0. Ventral plate without
lateral concavity (Fig. 87 B).
Pupa: Cocoon with small basal anterior projection (Fig. 88 E); length along base 3.0-
3.3 mm, on dorsum 2.4-2.8 mm, gills 2.7-3.3 mm. Frontoclypeus and thorax with rounded
and smooth tubercles; trichomes curved basally and with 3-6 branches. Gill with 6
branches (Fig. 89 B).
Larva: Maximum length 5.5-5.8 mm. Posterior half of cephalic apotome darkened,
with negative spots (Fig. 90 B). Postgenal cleft deep and narrow; postgenal bridge about
1/6 of hypostomium height. Antennae shorter than stem of cephalic fan; ratio of
proximal, medial and distal segments = 1:1.0-1.2:1.6-2.0. Cephalic fan with 42-48 rays.
Mandible with 1 latero-mandibular process. Lateral sclerite of thoracic proleg with 23-25
teeth. Anal sclerite with hairs. Anal ring with 73-76 rows with 14-15 hooks each. Rectal
papillae with 12 diverticula on each lobe.
Distribution: Ecuador: Azuay, Cuenca, Pichincha.
Bionomics: Unknown.
Discussion: Simulium arcabucense is similar but has a longer gonostylus (ratio of length/
width = 2.7), the trichomes of the pupal frontoclypeus and thorax with a straight and wide
base, the tubercle borders wrinkled and the tertiary gill branches bifurcated nearer the base.
Simulium pifanoi can be differentiated by the 8 gill branches and longer gonostylus. Simulium
gabaldoni has fewer tubercles on the pupal frontoclypeus and thorax and the basal portion of
the cibarium has tubercles and spines, and the legs are entirely black.

Simulium (Ectemnaspis) antonii WYGODZINSKY


Simulium antonii WYGODZINSKY, 1953b: 325.
Simulium (Ectemnaspis) antonii: COSCARÓN, 1984: 302; 1987: 29; 1991: 249; CROSSKEY & HOWARD,
1997: 29; 2004: 21.
Female: Wing length 3.0 mm. Scutum pale orange, pale yellow border laterally and
posteriorly, sometimes with dark brown sublateral stripes. Scutellum orange; metanotum
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 393

A C D

P
E
B

G H

K
J L

O M N

Figure 87. Male. Ventral plate. A: S. (Ectemnaspis) arcabucense. B: S. (E.) pautense. C: S. (E.)
rubiginosum. D: S. (E.) tunja. E: S. (E.) albanense. F: S. (E.) gabaldoni. G: S. (E.) mayuchuspi. H: S. (E.)
romanai. I: S. (E.) wolffhuegeli. J: S. (E.) perflavum. K: S. (E.) kabanayense. L: S. (E.) rorotaense. M: S. (E.)
dinellii. N: S. (E.) shewellianum. O: S. (E.) ochraceum. P: S. (E.) antillarum.
394 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

black; pleurae orange with episternum on ventral half blackish with silver pruinosity. Legs
dark brown to black. Abdomen with tergites I-IV yellowish, sometimes IV darkened, V-IX
dark brown to blackish; I-V matte, the following tergites shiny. Basal sector of R bare.
Paraproct 1/3 longer than wide.
Male, Pupa and Larva: Unknown.
Distribution: Bolivia: Cochabamba.
Bionomics: Unknown.
Discussion: Without knowledge of the pupa and male, it is difficult to separate S.
antonii from other species of the subgenus Without hairs on the basal sector of R and with
similar scutum and abdomen coloration, the closest species are S. pautense, S. pifanoi, S.
arcabucense and S. furcillatum, which are smaller (wing length 2.4-2.7 mm). In S. pautense, the
paraproct is as long as wide, and in S. furcillatum the scutum is brownish and the
metanotum grayish brown. Simulium tunja has the paraproct about half as wide as long and
no dark brown sublateral stripes on the scutum.

Simulium (Ectemnaspis) tunja COSCARÓN (Figs. 85 I; 86 L; 87 D; 88 G, Q; 90 R.)


Simulium (Ectemnaspis) tunja COSCARÓN, 1990: 126-128; CROSSKEY & HOWARD, 1997: 30; 2004: 21.
Female: Wing length 2.8-2.9 mm. Scutum yellowish with median and sublateral area
from brownish orange to black, scutum rim and scutellum yellow, and metanotum black.
Abdomen with tergites I-II yellowish, III (IV) yellow with tergal plates brownish to dark
brown IV-V generally velvety black, VI blackish, waxy bright. Basal sector of R bare.
Eighth sternite with about 40 hairs on each side; paraproct about 2/3 as wide as long, with
thick apex (Fig. 85 I).
Male: Wing length 2.7 mm. Scutum blackish; abdomen as in female but more
darkened, with segment III tergal plate largest and darker, VI-VII with 1+1 silvery spots.
Ratio of length/width of hind basitarsus = 4.3. Gonostylus slightly shorter than
gonocoxite (Fig. 86 L). Ventral plate with slight lateral depression (Fig. 87 D).
Pupa: Cocoon with anterior basal projection (Fig. 88 G), weave spongy; length along
base 3.2-3.7 mm, on dorsum 2.9-3.1 mm, gills 2.1-2.4 mm. Frontoclypeus and thorax with
numerous smooth tubercles (Fig. 88 Q); trichomes single or bifid. Gills with 10 branches,
primary dorsal and median branches bifurcated near base.
Larva: Maximum length 5.0-5.2 mm. Cephalic apotome scarcely pigmented, without
positive spots. Postgenal bridge about 1/6 of hypostomium length. Antennae about same
length as cephalic stem apex; ratio of proximal, medial and distal segments = 1:1.2-1.5:1.2-
1.5. Cephalic fan with 29-32 rays. Anal sclerite with hairs and scales (Fig: 90 R). Anal ring
with 67-70 rows with 12-13 hooks each. Rectal papillae with 16-18 diverticula on each lobe.
Distribution: Colombia: Boyacá, Cundinamarca.
Bionomics: Unknown.
Discussion: Simulium kabanayense has similar coloration but can be differentiated by
the presence of hair on the basal sector of R, small teeth on the cibarium, smaller
tubercles on the frontoclypeus and thoracic trichomes with 3-6 branches. Simulium
ignescens, with 10 branches, has a thinner cocoon, the frontoclypeus and thorax without
tubercles and hair on the basal sector of R. Simulium bicoloratum, also with 10 branches, has
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 395

B D C

A
H
J
F
E
G
K
L

P
M

N
O

U Y

S
µm
50

V Z
T

0.1 mm

R W X

Figure 88. Pupa. A-L: General aspect; A: S. (Ectemnaspis) arcabucense; B: S. (E.) bicoloratum; C: S. (E.)
bicornutum; D: S. (E.) furcillatum; E: S. (E.) pautense; F: S. (E.) tolimaense; G: S. (E.) tunja; H: S. (E.) anaimense;
I: S. (E.) lutzianum; J: S. (E.) romanai; K: S. (E.) wolffhuegeli; L: S. (E.) suarezi. M-X: Frontoclypeus; M: S.
(E.) bicoloratum; N: S. (E.) pautense; O: S. (E.) pifanoi; P: S. (E.) rubiginosum; Q: S. (E.) tunja; R: S. (E.)
lutzianum; S: S. (E.) mayuchuspi; T: S. (E.) romanai; U: S. (E.) wolffhuegeli; V: S. (E.) maroniense; W: S. (E.)
dinellii; X: S. (E.) shewellianum. Y: Trichome of S. (E.) anaimense. Z: Tubercles of S. (E.) albanense.
396 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

B
E
A

H
C

L
K

J M

Figure 89. Pupa. Gill. A: S. (Ectemnaspis) arcabucense. B: S. (E.) pautense. C: S. (E.) pifanoi. D: S. (E.)
roquemayu. E: S. (E.) rubiginosum. F: S. (E.) anaimense. G: S. (E.) lutzianum. H: S. (E.) romanai. I: S. (E.)
wolffhuegeli. J: S. (E.) perflavum. K: S. (E.) kabanayense. L: S. (E.) maroniense. M: S. (E.) rorotaense. N: S.
(E.) suarezi. O: S. (E.) trombetense (based on HAMADA & ADLER, 1988). P: S. (E.) shewellianum.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 397

the frontoclypeal and thoracic trichomes with more than 20 branches and hair on the
basal sector of R. Simulium antonii has similar coloration but without the darker median
and sublateral stripes sometimes present in S. tunja, but the unknown male, pupa and larva
of S. antonii are inconvenient in evaluating the possible synonymy of these species.

Simulium (Ectemnaspis) tolimaense COSCARÓN (Fig. 88 F.)


Simulium (Ectemnaspis) tolimaense COSCARÓN, 1984: 301 n. nom. to Simulium (Ectemnaspis) simplex
WYGODZINSKY & COSCARÓN, 1982: 11 (preoc. by Simulium simplex GIBBINS, 1936); COSCARÓN,
1990: 128; CROSSKEY & HOWARD, 1997: 30; 2004: 21.
Female (Pharate): Scutum very similar to that of males of S. bicornutum and S.
furcillatum; fronto-ocular triangle about as long as wide. Cibarium as in S. furcillatum. Claw
with subbasal tooth well developed
Male (Pharate): Pigment pattern not fully developed, mesonotum yellowish brown.
Genitalia not discernible.
Pupa: Cocoon with anterior subrectangular flap-like projection dorsally and also
projected basally (Fig. 88 F); walls of cocoon thick, individual threads not perceptible; length
along base 3.4-3.5 mm, on dorsum 3.2-3.5 mm, gills 1.8-2.0 mm. Frontoclypeus and thorax
with relatively abundant tubercles; trichomes with 5-8 branches, with curved base. Gills with 5
branches, surface minutely wrinkled. First branch perpendicular to stem of gill. Secondary
branch bifurcated; lower branch subdivided subapically, with resulting branch about as long as
common stem. Upper branch also bifurcated but the two branches longer than common stem.
Larva: Unknown.
Distribution: Colombia: Tolima.
Bionomics: The species has been collected in the Central Cordillera at 3100 m, on
the slope of the hillside. No more information is available.
Discussion: The scutum ornamentation is typical of the species of the Simulium
(Ectemnaspis) subgenus. The unique gill number and branching permit identification.

Simulium (Ectemnaspis) bicornutum WYGODZINSKY & COSCARÓN (Figs. 86 F; 88 C; 90 M.)


Simulium (Ectemnaspis) bicornutum WYGODZINSKY & COSCARÓN, 1982: 2; COSCARÓN, 1984: 300; 1990:
128; CROSSKEY & HOWARD, 1997: 29; 2004: 21.
Female (Pharate): Scutum and scutellum yellow; color pattern of scutum similar to
that of S. bicoloratum. Abdominal tergites I-IV grayish yellow, V-IX dark grayish brown.
Fronto-ocular triangle approximately as deep as wide. Eighth sternite with about 30 setae
on each side; gonapophysis subtriangular, glabrous, with few bands of heavier sclerotiza-
tion; paraproct about as long as wide at base, rounded apically.
Male: Wing length 3.0 mm. Scutum orange, sublateral pattern dark brown to black.
Metanotum black. Abdomen with segment I brown, II-III pale greenish yellow, IV brown
dorsally, V-IX velvety black; VI with 1+1 large silvery spots laterally (Fig: 86 F). Basal
sector of R glabrous. Ratio of length/width of hind basitarsus = 3.5. Gonostylus as long
as gonocoxite, elongate, subconical, with apical portion somewhat curved. Basal plate
subtriangular, its sides almost straight.
398 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Pupa: Cocoon with a broad median projection dorsally. Walls of cocoon thick;
length of cocoon along base 3.7 mm, on dorsum 4.0 mm, gills 1.8 mm. Frontoclypeus and
thorax with few or without tubercles; trichomes of head with 4-8 branches, of thorax with
2-6 branches. Gill with 2 thick branches, roughly half-moon shaped in lateral view; arms
pointed apically, each with small tooth-like process on inner surface (Fig. 88 C).
Larva: Maximum length 5.8 mm. Cephalic apotome rather uniformly pigmented,
somewhat darker at base. Ratio of antennal segments: proximal, medial and distal =
1:1.7:2.5 (Fig: 90 M). Cephalic fan with about 40 rays. Postgenal cleft narrow and deep;
postgenal bridge about 1/5 of hypostomium length. Ninth segment without ventral
tubercles. Anal sclerite with abundant and minute setae. Anal ring with about 85 rows with
16 hooks each. Rectal papillae with about 16 diverticula on each lobe.
Distribution: Colombia: Cundinamarca.
Bionomics: The species has been collected in the Paramo region at and altitude of
3300-3700 m. The immatures live on trailing blades of grass in torrential, small creeks 1-
3 m wide, with clear water.
Discussion: The unique pupal gill allows this species to be differentiated from other
Neotropical black flies.

Simulium (Ectemnaspis) furcillatum WYGODZINSKY & COSCARÓN (Figs. 85 E; 86 A, G; 88 D.)


Simulium (Ectemnaspis) furcillatum WYGODZINSKY & COSCARÓN, 1982: 6; COSCARÓN, 1984: 300; 1990:
128; CROSSKEY & HOWARD, 1997: 29; 2004: 21.
Female: Wing length 2.4 mm. Scutum orange brown, lateral borders and lateral
incision posterior to pronotal incisions, greenish yellow. Scutellum greenish yellow.
Metanotum grayish brown with 1+1 anterior grayish spots with certain illumination.
Abdominal segments I-IV with terga grayish yellow, fourth tergal plate light brown.
Cibarium smooth with slight median depression and large, heavily pigmented cornuae.
Wing without hairs on basal sector of R. Claws with small but distinct subbasal tooth.
Eighth sternite with inner border conspicuously sclerotized. Paraproct subtriangular,
about as long as wide at base (Fig. 85 E).
Male: Wing length 2.9 mm. Scutum and scutellum pale greenish yellow; scutum
with wide median dark marking covering anterior 3/4 and 1 pair of lateral wedge-
shaped spots beginning at anterior sixth of dark central marking (Fig. 86 A).
Metanotum dark grayish brown. Abdominal segment I brown, II-III with tergal plates
pale grayish yellow; pleuron and sternum grayish white; V-IX dark brown, VIII with
1+1 lateral silvery spots (Fig. 86 G). Ratio of length/width of hind basitarsus about
4.0. Gonocoxite about 2.5 times as long as wide with well-developed apical spine.
Ventral plate subtriangular.
Pupa: Cocoon translucent, with flap-like projection dorsally at middle. Fabric of
cocoon uniform, parchment like, individual threads difficult to discern. Cocoon length
ventrally 3.2 mm, on dorsum 3.7 mm, gills 2.1 mm. Frontoclypeus and thorax without
tubercles, except group between facial and frontal trichomes; trichomes with 3-6
branches. Gill consisting of 2 rigid arms forming U shape, lower arm of gill pointed,
apical half of upper arm fork-like, with 3 tine-like processes (Fig. 88 D).
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 399

A B E C

D O
F G
H

I J M
0.1 mm

0.1 mm
K
0.1 mm

L
0.05 mm

P N

S T
Figure 90. Larva. A-H: Cephalic apotome; A: S. (Ectemnaspis) rubiginosum; B: S. (E.) pautense; C: S.
(E.) arcabucense; D: S. (E.) albanense; E: S. (E.) romanai; F: S. (E.) wolffhuegeli; G: S. (E.) perflavum; H: S.
(E.) rorotaense. I-L: Hypostomium and postgenal cleft; I: S. (E.) arcabucense; J: S. (E.) albanense; K: S.
(E.) rorotaense; L: S. (E.) shewellianum. M-O: Antennae; M: S. (E.) bicornutum; N: S. (E.) perflavum; O:
S. (E.) rorotaense. P: Trichomes of body integument of S. (E.) anaimense. Q: Apical portion of
mandible of S. (E.) mayuchuspi. R: Anal sclerite of S. (E.) tunja. S-T: Diverticula of rectal papillae
lobes; S: S. (E.) perflavum; T: S. (E.) pifanoi.
400 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Larva: Maximum length 5.7 mm. Cephalic apotome rather homogeneously pigment-
ed, somewhat darker at base. Ratio of antennal segments: proximal, medial and distal =
1:1.3:1.8. Cephalic fan with about 35 rays. Postgenal cleft deep, narrowly subtriangular;
postgenal bridge about half of hypostomium length. Lateral sclerite of proleg with about
24 teeth. Body integument glabrous. Anal ring with 84 rows with 14-16 hooks each. Rectal
papillae with about 16 diverticula each.
Distribution: Colombia: Boyacá, Cundinamarca.
Bionomics: This species breeds on blades of grasses trailing in rapid, clean streams in the
Paramos of the Eastern Cordillera at 3200-3400 m and at water temperatures of about 10 °C.
Discussion: The most similar species is S. bicornutum, which has a pupal gill with the
two arms undivided, unlike in S. furcillatum, which has the upper arm divided into 3 tine-
like appendages. Adult coloration is similar between these two species.

Simulium (Ectemnaspis) bicoloratum MALLOCH (Figs. 84 B, F; 85 D; 88 B, M.)


Simulium bicoloratum MALLOCH, 1912: 649; VARGAS & DIAZ NAJERA, 1953b: 140; WYGODZINSKY,
1953b: 327-330; 1958: 133.
Simulium (Ectemnaspis) bicoloratum WYGODZINSKY, 1971: 19-31, 34; COSCARÓN, 1984: 298-299; 1987:
28; 1990: 128; 1991: 249-251; CROSSKEY & HOWARD, 1997: 29; 2004: 21; COSCARÓN ARIAS,
2003: 195, 200, 205, 209, 215, 218.
Odagmia bicolorata: ENDERLEIN, 1925: 208; 1930: 97.
Simulium gaudeatum KNAB, 1914b: 81; 1914c: 123.
Ectemnaspis macca ENDERLEIN, 1934b: 281 (preoc.); 1936: 122.
Simulium molli VARGAS, 1943a: 142; 1945a: 165.
Dasypelmoza macca ENDERLEIN, 1934b: 276. (n. syn.).
Female: Wing length 2.8-3.2 mm. Frons and clypeus silver gray. Scutum yellow with
extensive area blackish as in figure 84 B; central area can change from brown to orange.
Scutellum yellow; metanotum velvety black with 1+1 silvery spots anteromedially.
Anepisternum and katepisternum black, remaining pleural area brownish. Abdomen with
segments I-II golden yellow, fringe of first segment blackish; dorsum of III-IV velvety
black, dorsum VI entirely yellow to partly or entirely dark brown or black; VII-IX brown
to black; terga III-V dull, VI-IX polished (Fig. 84 F). Cibarium with large heavily
pigmented cornuae, space between cornuae smooth. Basal sector of R with setae. Claws
with medium-sized subbasal tooth. Eighth sternite with about 15-20 setae on each side;
gonapophysis subtriangular, glabrous, rather conspicuously sclerotized on median half.
Paraproct slightly longer than wide at base, subconical (Fig. 85 D).
Male: Wing length 2.8-3.2 mm. Color of scutum much as in female, but light-
colored area of center of posterior portion shorter, not as long as posterior black area.
Scutellum yellow, darkened at sides and in many cases narrowly along base. Metanotum as
in female. Abdomen velvety black with 1+1 large silvery spots on tergite VI. Hind
basitarsus 4 times as long. Ventral plate subtriangular, strongly constricted at sides,
producing abruptly narrowed apical portion, slightly incised medially.
Pupa: Cocoon very thick, spongy, threads poorly visible; anterior margin
conspicuously thickened, dorsal surface with 2 or 3 longitudinal ridges along whole
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 401

length of cocoon (Fig. 88 B); length along base 3.5-4.0 mm, on dorsum 2.7-3.0 mm,
gills 2.0 mm. Frontoclypeus and thorax with numerous large tubercles with small
protuberances, arranged in irregular fashion. Trichomes large, heavily sclerotized,
almost black with 30-40 branches (Fig. 88 M). Gill with 8 branches emerging from 3
short primary branches. Surface of branches and filaments with minute tubercles of 2
sizes, giving wrinkled appearance.
Larva: Maximum length 6.5-7.0 mm. Cephalic apotome yellowish with positive
spots. Antennae as long as stem of cephalic fan; ratio of proximal, medial and distal
segments = 1:1.2-1.5:1.1-1.5. Mandible with 1 latero-mandibular process. Cephalic fan
with 40-50 rays. Hypostomium with intermediate teeth scarcely prominent, median and
lateral teeth about same height. Postgenal cleft pointed, subtriangular; postgenal bridge
slightly shorter than hypostomium. Body integument without hair. Anal sclerite with small
setae. Anal ring with 80-85 rows of about 17 hooks each. Rectal papillae with 15-20
elongate diverticula on each lobe.
Distribution: Bolivia; Colombia: Cauca, Cundinamarca, Tolima; Ecuador: Pichin-
cha; Peru: Lima; Venezuela: Mérida.
Bionomics: This species has been collected on aquatic vegetation, grasses and under
stones in mountainous areas at 2000-3600 m in fast-flowing creeks 3-5 m wide and with
water temperatures of about 9 °C. It also has been found in irrigation ditches. Female
feeding habits are unknown.
Discussion: Simulium bicoloratum varies in color. WYGODZINSKY (1971) considered
this condition to justify its position in Simulium (Ectemnaspis). The closest species are S.
cormonsi, S. jaimeramirezi, S. romanai and S. lutzianum. Differentiation from similar species is
explained in the description of each similar species. Dasypelmoza macca, based on the
original description, fits into the range of variation of S. bicoloratum.

Simulium (Ectemnaspis) cormonsi WYGODZINSKY


Simulium cormonsi WYGODZINSKY, 1971: 31-32; RAMÍREZ PÉREZ, 1983: 43.
Simulium (Ectemnaspis) cormonsi; COSCARÓN, 1984: 299; 1987: 28; 1990: 129; CROSSKEY & HOWARD,
1997: 29; 2004: 21.
Female and Male: Not distinguishable by color or morphology from S. bicolora-
tum. Female: fronto-ocular triangle wider than high; paraproct longer and narrower.
Male: gonostylus slightly narrowed and ventral plate more constricted laterally, with
apical half narrower.
Pupa: Cocoon as in S. bicoloratum. Trichomes as in S. bicoloratum. Gill length 3.0 mm.
Gill with 6 branches, emerging from 3 short primary branches.
Larva: Unknown.
Distribution: Venezuela: Mérida.
Bionomics: The immature stages of this species are found on vegetation and stones
at altitudes of 3000-3400 m in torrential streams 3-5 m wide, with water temperatures of
about 13-15 °C.
Discussion: Differences from S. bicoloratum are in the number of gill branches and
the slight morphological differences in shape, as explained in the description.
402 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Simulium (Ectemnaspis) jaimeramirezi WYGODZINSKY


Simulium jaimeramirezi WYGODZINSKY 1971: 32-34; RAMÍREZ PÉREZ, 1983: 48.
Simulium (Simulium) jaimeramirezi: RAMÍREZ PÉREZ & VULCANO, 1973: 395.
Simulium (Ectemnaspis) jaimeramirezi COSCARÓN, 1984: 299; 1987: 28; 1990: 129; CROSSKEY & HOWARD,
1997: 29; 2004: 21.
Female and Male: Color and structure, including genitalia, similar to those of S.
bicoloratum.
Pupa: Cocoon with structure as in S. bicoloratum, dorsally with only median
longitudinal ridge; length at along 4.4 mm, on dorsum 2.9-3.2 mm; gills 1.9-2.8 mm.
Frontoclypeus and thorax with tubercles less numerous than in S. bicoloratum.
Larva: Unknown.
Distribution: Venezuela: Mérida.
Bionomics: This species was collected in torrential streams 3-5 m wide, at 3050-
3400 m, on the undersurfaces of stones at 13-15 °C.
Discussion: Simulium jaimeramirezi is very similar to S. cormonsi and S. bicoloratum, but
can be differentiated by the weak coloration and small frontoclypeal and thoracic
trichomes. It also can be separated from S. bicoloratum by the number of gill branches. It
would be useful to have more material and conduct cytological or molecular studies
comparing this species with S. cormonsi.

Simulium (Ectemnaspis) anaimense COSCARÓN & MUÑOZ DE HOYOS


(Figs. 88 H, Y; 89 F; 90 P.)
Simulium (Ectemnaspis) anaimense COSCARÓN & MUÑOZ DE HOYOS, 1995: 587-592.
Simulium (Psilopelmia) anaimense: CROSSKEY & HOWARD, 1997: 60; 2004: 54.
Female (Pharate): Scutum yellow. Cibarium with tubercles in space between
cornuae. Sensory vesicle of maxillary palp about half length of palpomere. Claw
subbasally with medium-sized, pointed projection.
Male (Pharate): Scutum and scutellum yellowish; metanotum blackish.
Pupa: Cocoon whitish, thick, reinforced at the anterior rim and on median dorsal
ridge (Fig. 88 H); texture felt like, threads only visible at high magnification. Length along
base 3.4 mm, on dorsum 2.6 mm, gill 2.3 mm. Frontoclypeal and thoracic tubercles
superficially rugose, moderate in number, but abundant on base of frontoclypeus and
insertion of gill; trichomes with curled base and 4-13 branches (Fig. 88 Y). Gill whitish,
thick, projected in several planes, dotted homogeneously and with 6 branches (Fig. 89 F).
Larva: Maximum length 4.0-5.5 mm. Cephalic apotome brown, dark basally, with
pattern similar to that of S. pautense. Antenna surpassing length of cephalic fan stem; ratio of
proximal, medial and distal segments = 1:1.0-1.1:1.5-1.6. Mandibles with 1 latero-mandibu-
lar process. Postgenal cleft deep, dome shaped; postgenal bridge about 1/6 hypostomium
length. Cephalic fan with 28-34 rays. Body integument with palmiform and simple trichomes
intermixed (Fig. 90 P). Thoracic proleg sclerite with about 16 teeth in 6-7 groups. Ninth
segment with 2 ventral papillae. Anal sclerite with abundant unbranched hairs. Anal ring with
64-70 rows with 11-14 hooks each. Rectal papillae with 12-18 diverticula on each lobe.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 403

Distribution: Colombia: Tolima on the Cordillera Central.


Bionomics: This species breeds in medium-sized torrential streams with clear, cold
water of 12 °C. Larvae and pupae were collected from trailing roots in swift flow.
Discussion: Simulium anaimense is similar in cocoon shape to several species of the S.
bicoloratum and S. romanai groups, but the unique, relatively thick and short gill branches
distinguish it. Also, the tubercles on the cibarium are similar to those of the S. romanai
group. It is also similar to S. albanense in relation to the integumentary setae of the larva.

ROMANAI species group

Simulium (Ectemnaspis) romanai WYGODZINSKY (Figs. 84 O; 86 D, N; 87 H; 88 T; 89 H; 90 E.)


Simulium romanai WYGODZINSKY, 1951b: 229.
Simulium (Ectemnaspis) romanai: COSCARÓN, 1984: 302-303; 1987: 29; 1990: 129-131; 1991: 252;
COSCARÓN ARIAS, 2003: 195, 200, 205, 209, 215, 219.
Simulium (Psilopelmia) romanai: CROSSKEY & HOWARD, 1997: 60; 2004: 54.
Simulium adolfolutzi WYGODZINSKY, 1951b: 234.
Female: Wing length 1.9-2.8 mm. Scutum yellowish to brown blackish on central area.
Scutellum orange; metanotum black. Abdomen with tergite I yellowish, II-IV brown with
center of tergite III dark brown or all tergites black, V-IX shiny dark brownish gray. Legs
yellowish brown with distal portion of femur, tibia and tarsomeres dark brown to mostly
brown. Cibarium with well-sclerotized rim, depressed at center, medially with rounded
protuberances and laterally with abundant small acuminate teeth-like spines (Fig. 84 O).
Basal sector of R with hair. Ratio of length/width of posterior basitarsus = 6.6. Eighth
sternite with about 25 setae, gonapophyses subtriangular, membranous, with divergent
internal border and slightly sclerotized; paraproct 2/3 longer than wide with rounded apex,
and genital fork with median stem length about same as 2 lateral branches together.
Male: Wing length 2.1-2.4 mm. Coloration as in female but often darker as in figure
86 D. Abdomen velvety black with 1+1 lateral silvery spots on tergites VI-VII. Ratio of
length/width of hind basitarsus = 4.8-5.1. Gonostylus subcylindrical with sides subparal-
lel on basal 2/3 and becoming thin toward apex, with 1 apical spur (Fig. 86 N). Ventral
plate wider than long, with strong lateral depression, in lateral view appearing as 2
superimposed pieces (Fig. 87 H).
Pupa: Cocoon thick, weave spongy, slightly projected basally and frequently with
dorsal longitudinal ridge, especially on anterior portion. Frontoclypeus and thorax with
abundant tubercles (Fig. 88 T); trichomes with 4-12 branches. Gill with 8 branches
relatively open from base (Fig. 89 H).
Larva: Maximum length 6 mm. Cephalic apotome with darkened median basal area,
head spots negative (Fig. 90 E). Antenna not surpassing apex of cephalic fan stem; ratio
of proximal, medial and distal segments = 1:1.0-1.4:0.9-1.5. Mandible with 1 latero-
mandibular process; cephalic fan with about 37 rays. Postgenal cleft very deep, acuminate
anteriorly; postgenal bridge very short. Body integument without hairs, anal sclerite
without scales or hairs. Lateral thoracic proleg sclerite with about 21 teeth. Anal sclerite
with about 68 rows with 12 hooks each. Rectal papillae with 5-9 diverticula on each lobe.
404 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Distribution: Argentina: Catamarca, Córdoba, Jujuy, La Rioja, Salta, San Juan,


Tucumán; Bolivia: Potosí; Colombia: Cauca, Cundinamarca, Valle; Ecuador: Los Rios,
Pastaza, Pichincha; Peru: Junin; Venezuela: Mérida.
Bionomics: This species breeds in creeks with clear, fast flow over stones and
aquatic vegetation. Female feeding habits are unknown.
Discussion: The most similar species is S. lutzianum because that species shows
variation in coloration, creating confusion. Simulium lutzianum can be differentiated in the
female by abdominal tergites II-III, which are frequently yellowish, and the cibarium with
several rows of rounded tubercles; in the male by the lack of black on the scutum disc and
thinner hind legs; in the pupa by the sparse tubercles on the frontoclypeus and thorax; and
in the larva by the lighter cephalic apotome and hair on the body integument. Cytological
studies by MORENO RAMÍREZ (Magister thesis, 1997) of S. lutzianum from Colombia and S.
romanai from Colombia and Argentina reveal different cytotypes. Simulium bicoloratum is
similar in pupal appearance and color of the adult, but the smooth basal portion of the
cibarium permits it to be differentiated.

Simulium (Ectemnaspis) lutzianum PINTO (Figs. 84 M, N; 85 C; 88 I, R; 89 G.)


Simulium lutzianum PINTO, 1931: 748; RAMÍREZ PÉREZ, 1971: 311; SHELLEY & ARZUBE, 1985: 330.
Simulium (Ectemnaspis) lutzianum: COSCARÓN, 1984: 303; 1987: 20; 1990: 131-132; 1991: 253-254.;
COSCARÓN ARIAS, 2003: 195, 200, 205, 209, 215, 218.
Ectemnaspis lutziana PY-DANIEL & MOREIRA SAMPAIO, 1995: 118.
Simulium (Psilopelmia) lutzianum: SHELLEY et al., 1997: 34; CROSSKEY & HOWARD, 1997: 60; 2004: 54.
Simulium lewisi RAMÍREZ PÉREZ, 1971: 349.
Simulium (Ectemnaspis) lewisi: COSCARÓN, 1984: 304; 1990: 132-133; SHELLEY et al., 1989: 93.
Simulium iguazuense COSCARÓN, 1976: 147; 1984: 304; 1987: 29.
Simulium (Psilopelmia) alirioi RAMÍREZ PÉREZ & VULCANO, 1973: 383 n. syn.
Female: Wing length 1.8-2.2 mm. Scutum and scutellum yellowish orange, metano-
tum blackish. Pleurae dark brown, with silver pruinosity. Legs dark brown except basal 2/
3 of fore tibiae, and mid and hind basitarsus, which are whitish with a reduced dark area
at apex of femora and tibiae. Abdomen with tergites I-II (III) yellowish to light brown,
(III) IV-IX brownish, dark gray to blackish, III-IV opaque, V-IX shiny. Tergal plates IV-IX
evident. Fronto-ocular area wider than high. Cibarium with sclerotized rim; central area
with 5-6 rows of rounded tubercles and acute small teeth-like spines sublaterally (Figs. 84
M-N); some specimens similar to S. romanai, wrinkled without definite tubercles. Basal
sector of R with hair. Ratio of length/width of hind basitarsus = 8. Claw with subbasal
tooth (Fig. 85 C). Genitalia similar to those of S. romanai.
Male: Wing length 1.6-2.0 mm. Coloration similar to that of female. Abdomen with
segments I-III yellowish or only I brownish, rest of tergites black, VI-VII with 1+1 lateral
silver spots. Ratio of length/width of hind basitarsus = 6.5-7.6. Genitalia as in S. romanai.
Pupa: Cocoon thick, weave spongy, with very thin fibers; anterior border reinforced
and dorsal medium longitudinal ridge (Fig. 88 I); length along base 2.2-3.4 mm, on
dorsum 1.8-2.8 mm, gills 1.8-3.3 mm. Frontoclypeal and thoracic tubercles sparse with
rounded borders (Fig. 88 R). Cephalic trichomes with 2-3 branches, thoracic with 5-7
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 405

branches. Gill with 8 branches from very open to meeting basally. Bifurcation of
secondary and tertiary branches at variable distances from base (Fig. 89 G).
Larva: Maximum length 3.7-4.8 mm. Cephalic apotome without special ornamenta-
tion, slightly darkened on posterior border, with short single hairs. Antennae surpassing
apex of cephalic fan stem; ratio of proximal, medial and distal segments = 1:0.9-1.2:1.3-
1.6. Mandible with 1 latero-mandibular process. Cephalic fan with about 35 branches.
Postgenal cleft nearly reaching hypostomium base. Body integument with numerous small
hairs. Thoracic proleg lateral sclerite with about 20 teeth. Anal ring with 55-74 rows with
11-13 hooks each. Rectal papillae with 8-12 diverticula on each lobe.
Distribution: Argentina: Misiones; Bolivia: Cochabamba; Brazil: Goiás, Paraná,
Roraima, Santa Catarina; Colombia: Cauca, Cundinamarca, Valle; Ecuador: Chimborazo,
El Oro, Esmeraldas, Guayas, Imbabura, Los Rios, Manabí, Napo, Pastaza, Pichincha,
Tungurahua, Zamora-Chinchipe; Panama: Panama; Peru: Cuzco, Huanuco, Junin; Vene-
zuela: Amazonas, Aragua, Barinas, Bolivar, Carabobo, Cojedes, Distrito Federal, Lara,
Mérida, Miranda, Monagas, Táchira, Trujillo, Yaracuy, Zulia.
Bionomics: Simulium lutzianum breeds in torrential creeks of mountainous areas, on
aquatic vegetation and stones. Female feeding habits are unknown.
Discussion: Simulium lutzianum shows great variation in morphology, including the
arrangement of the cibarial tubercles and size of the teeth, extent of yellow on the female
and male abdomen and the length of the pupal gill and distance of bifurcation of the
secondary branches. SHELLEY et al. (1997: 36) synonymized S. lewisi based on molecular
and cytological studies in Ecuador (SAWYER, 1991), considering that the color variation is
not linked to cytospecies and that cytoforms A, B and C belong to the S. lutzianum species
complex. Additional non morphological studies with material from other regions will be
necessary to elucidate the extent of S. lutzianum variation. Similarity with S. romanai is
explained in that species description. Simulium bicoloratum is very similar in pupal shape and
adult coloration, but has a smooth cibarium between the cornuae. Because the description
of S. alirioi is included in the range of variation of S. lutzianum, it is synonymized here.

Simulium (Ectemnaspis) gabaldoni RAMÍREZ PÉREZ (Figs. 84 L; 87 F.)


Simulium gabaldoni RAMÍREZ PÉREZ, 1971: 352; 1983: 46.
Simulium (Ectemnaspis) gabaldoni: COSCARÓN, 1984: 307-308; 1987: 29; 1990: 134.
Simulium (Psilopelmia) gabaldoni: CROSSKEY & HOWARD, 1997: 60; 2004: 54.
Female: Wing length 2.0-2.1 mm. Scutum yellowish-orange, scutellum yellow-
lemon, metanotum dark brown. Legs brown with darkened areas. Abdomen with tergite I
yellow-lemon, II-III yellow with posterior border brown, IV-IX blackish. Cibarium with
several rows of tubercles on median area and acute teeth at sides to cornuae (Fig. 84 L).
Basal sector of R with 1 row of hair. Claws with subbasal tooth. Paraproct about as long
as wide at base, very similar to that of S. arcabucense.
Male: Wing length 1.8-1.9 mm. Color as in female but darker; legs with more
extensive blackish area. Abdomen all black except 1+1 lateral silvery spots on segments
VI-VII. Gonostylus subcylindrical, slightly shorter than gonocoxite. Ventral plate about as
long as wide, with lateral constrictions (Fig. 87 F).
406 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Pupa: Cocoon thick, threads not evident, with slight anterior projection; length
along base 2.3-2.6 mm, on dorsum 2.0-2.5 mm, gills 2.6-3.2 mm. Frontoclypeus and
thorax with sparse tubercles; trichomes with 3-6 branches. Gills with 6 branches,
bifurcated near base.
Larva: Unknown.
Distribution: Colombia: Valle; Venezuela: Monagas and Sucre.
Bionomics: In Colombia, this species was collected at an altitude of 1400 m. No
other information is available.
Discussion: The coloration and number of gill branches of this species are similar
to the condition in S. arcabucense and S. pautense, but these species have a smooth cibarium
and no hairs on the basal sector of R. Simulium lutzianum is very similar, but has a pupa
with 8 gill branches.

Simulium (Ectemnaspis) albanense COSCARÓN (Figs. 84 K; 85 A; 86 M; 87 E; 88 Z; 90 D, J.)


Simulium (Ectemnaspis) albanense COSCARÓN, 1990: 135-136.
Simulium (Psilopelmia) albanense: CROSSKEY & HOWARD, 1997: 60; 2004: 54.
Female: Wing length 2.8-3.0 mm. Frons and clypeus brownish with silvery
pruinosity. Scutum and scutellum yellowish; scutum in some specimens with median and
sublateral areas blackish brown. Pleurae yellowish to light or dark brown. Metanotum
black. Legs blackish. Abdomen with segments I-III yellow, III with tergal plate brown, IV-
V velvety black, VI-IX shiny brown black. Fronto-ocular triangle wider than high.
Cibarium with rounded tubercles on median area and with sparse spines laterally (Fig. 84
K). Basal portion of R without hair (Fig. 85 A). Eighth sternite with about 16 setae on
each side. Gonapophysis and paraproct very similar to those of S. romanai.
Male: Wing length 2.3-2.5 mm. Coloration as in female, with darkened scutum area
larger, and frequently abdominal tergite III yellowish. Gonostylus about 1/3 gonocoxite
length, subquadrate, with strong apical spur (Fig. 86 M); ventral plate as in figure 87 E.
Pupa: Cocoon weave thin with delicate threads, anterior border slightly reinforced;
length along base 3.0-3.2 mm, on dorsum 2.7-3.0 mm, gills 2.0-2.5 mm. Frontoclypeus
and thorax with numerous elongate tubercles (Fig. 88 Z); trichomes with 4-8 branches.
Gill with 8 branches grouped as bundle; bifurcation of tertiary branches at about half
total gill length.
Larva: Maximum length 6.0 mm. Cephalic apotome homogeneously darkened, with
sublateral areas slightly lighter and with numerous multibranched hairs (Fig. 90 D).
Postgenal cleft deep, dome shaped (Fig. 90 J); postgenal bridge about 1/8 hypostomium
length. Cephalic fan with about 30 rays. Ratio of antennal segments: proximal, medial and
distal = 1:1.6:2.3. Mandible with 1 latero-mandibular process. Body integument with hairs
single or bifid. Lateral sclerite of thoracic proleg with about 21 teeth. Anal sclerite with
abundant hair of 1-4 branches or spatulate shape. Anal ring with 64-77 rows of 12-13
hooks each. Rectal papillae with 3 diverticula on each lobe.
Distribution: Colombia: Bogotá, Cundinamarca.
Bionomics: Material of this species was collected at altitudes of 2400-3050 m. No
more information is available.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 407

Discussion: The adults are very similar in coloration to other species of this group,
but can be differentiated by the short subquadrate gonostylus, elongate tubercles of the
pupal frontoclypeus and thorax, and pupal gill filaments arranged in a bundle.

Simulium (Ectemnaspis) wolffhuegeli (ENDERLEIN)


(Figs. 84 G, P; 86 O; 87 I; 88 K, U; 89 I; 90 F.)
Wilhelmia wolffhuegeli ENDERLEIN, 1922: 70.
Morops wolffhuegeli ENDERLEIN, 1930: 94.
Simulium wolffhuegeli EDWARDS, 1931: 127; WYGODZINSKY, 1950: 88-96; COSCARÓN & WYGODZINSKY,
1960: 1124; COSCARÓN, 1968: 66.
Psilopelmia wolffhuegeli ENDERLEIN, 1934b: 284.
Tricodagmia haarupiana ENDERLEIN, 1936: 127.
Simulium (Ectemnaspis) wolffhuegeli: COSCARÓN, 1984: 308-310; 1987: 29; 1990: 136; 1991: 255-257.
Simulium (Psilopelmia) wolffhuegeli: CROSSKEY & HOWARD, 1997: 60; 2004: 54.
Female: Wing length 1.9-2.1 mm. Scutum yellowish-orange with yellow hair,
scutellum grayish yellow; metanotum dark brown; pleurae yellow with grayish pruinosity.
Legs grayish yellow with apex of femora, tibiae and tarsomeres dark brown. Abdomen
yellowish brown with great variation in color, mostly with tergites I-IV yellowish, tergal
plate of tergites III-V darker, VI-IX shiny brownish gray (Fig. 84 G). Cibarium with
several rows of rounded tubercles medially and numerous spines laterally (Fig. 84 P).
Basal sector of R with hair. Claws with short subbasal tooth. Paraproct slightly longer than
wide, gonapophysis subovoidal with internal border sclerotized.
Male: Wing length 1.8-2.0 mm. Color generally blackish. Scutum black with pronotal
and posterior border yellowish gray and covered by abundant silky, grayish pilosity.
Scutellum and metanotum black. Abdomen blackish. Legs black, yellowish gray on clear
areas. Gonostylus subrectangular, slightly longer than wide, with 7-15 spurs on apical
border (Fig. 86 O). Ventral plate wider than long, with small projection mediodistally,
without lateral constriction (Fig. 87 I).
Pupa: Cocoon weave uniform, with threads evident and surface wrinkled, anterior
border straight (Fig. 88 K). Length along base 2.6-3.0 mm, on dorsum 2.0-2.4 mm, gills
1.5-1.9 mm. Frontoclypeus and thorax with numerous tubercles. Cephalic trichomes with
1-3 branches; thoracic with 4-9 branches (Fig. 88 U). Gills with 8 branches (Fig. 89 I).
Larva: Maximum length 5.0 mm. Cephalic apotome with variable ornamentation,
typically darkened basally and posteriolaterally, sometimes with negative spots (Fig. 90 F).
Antenna not surpassing apex of cephalic fan stem; ratio of antennal segments: proximal,
medial and distal = 1:1.1-1.4:1.1. Postgenal cleft deep, dome shaped, nearly reaching
hypostomium. Body integument with short hairs. Lateral sclerite of thoracic proleg with
about 22 teeth. Cephalic fan with 25-31 rays. Anal ring with 65-70 rows with 14 hooks
each. Rectal papillae generally with 0-1 diverticula on each lobe. Anal sclerite without
scales or hairs, or the latter very scarce.
Distribution: Argentina: Buenos Aires, Catamarca, Chubut, Córdoba, Jujuy, La
Pampa, La Rioja, Mendoza, Neuquén, Río Negro, Salta, San Juan, San Luis, Santa Cruz,
Tucumán; Bolivia: Cochabamba.
408 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Bionomics: This species breeds in rivers, slower creeks and irrigation canals with clear,
but not necessarily crystalline, water and aquatic vegetation, from sea level to 2000 m. Females
are haematophagous and dangerous to horses and cows; they generally don’t bite humans.
Discussion: Females are very similar to some other species of the S. romanai group,
but in S. wolffhuegeli the males have a black thorax, short gonostylus with 7-15 spurs, and
ventral plate much wider than high.

Simulium (Ectemnaspis) mayuchuspi COSCARÓN (Figs. 84 C; 85 B; 86 C; 87 G; 88 S; 90 Q.)


Simulium (Ectemnaspis) mayuchuspi COSCARÓN, 1990: 136-139; COSCARÓN ARIAS, 2003: 198, 205, 209, 215.
Simulium (Psilopelmia) mayuchuspi: CROSSKEY & HOWARD, 1997: 60; 2004: 54.

Female: Wing length 2.5-2.6 mm. General coloration blackish. Scutum blackish gray
with 1+1 light grayish subtriangular anterior submedian spots and lateral border light
grayish (Fig. 84 C); with certain lighting, 1+1 submedian blackish stripes can be seen
joining on posterior 1/3, and aforementioned anterior grayish spots change to black;
scutellum brown with posterior border yellowish. Hairs of mesonotum from silvery to
golden, depending on illumination. Metanotum brown with dark grayish pruinosity.
Pleurae and legs brownish black, with light brown on anterior half of fore tibia, anterior
1/5 of femora, anterior 3/4 of hind basitarsus and anterior half of second hind
tarsomere; leg hairs yellowish-gray. Abdomen dark brown to black with tergites II-IV
velvety black and V-IX shiny brown; ventrally brownish gray. Fronto-ocular triangle not
deep. Cibarium with well-sclerotized rim; rounded tubercles medially and abundant acute
teeth laterally, as in S. romanai and S. lutzianum. Basal sector of R with hairs. Claws without
subbasal tooth (Fig. 85 B). Eighth sternite with 25-30 setae on each side. Gonapophyses
subtrianglular, with internal borders slightly divergent and little sclerotized.
Male: Wing length 2.5-2.6 mm. Scutum black; with posterior illumination 1+1
grayish anterior submedian triangular spots can be seen continued anterolaterally by short
silvery gray bands, and irregular transverse spots on posterior third (Fig. 86 C); with
anterior light, scutum gray opaque with median and posterior 1/3 black. Scutellum,
metanotum and pleurae as in female. Abdomen black with 1+1 large lateral silvery spots
on segments II, III, VI, VII. Ratio of length/width of hind basitarsus = 5.2. Gonostylus
about 3 times as long as wide, curved on half distal and with apical spur similar to that of
S. arcabucense. Ventral plate with strong lateral constriction (Fig. 87 G).
Pupa: Cocoon thick with spongy weave; threads very thin not obvious, slightly thicker on
anterior border, and dorsal ridge not well defined, similar to that of S. romanai. Frontoclypeus
and thorax with numerous thin tubercles (Fig. 88 S). Cephalic trichomes with 2-7 branches and
thoracic with 7-12 branches. Gill with 8 branches, with branching close to base.
Larva: Cephalic apotome light brown, slightly ornamented with basal median area
darker. Ratio of antennal segments: proximal, medial and distal = 1:1.3:1.5. Cephalic fan
with about 36 rays. Mandible with 2 latero-mandibular processes (Fig. 90 Q). Postgenal
cleft about 1/7 hypostomium length. Body integument without hairs. Anal sclerite with or
without very sparse hairs but no scales. Anal ring with 65 rows with 13-17 hooks each.
Rectal papillae with 5-10 diverticula on each lobe.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 409

Distribution: Ecuador: Pichincha; Peru: Junín.


Bionomics: Simulium mayuchuspi breeds in small creeks with fast flowing, crystalline
waters, and aquatic vegetation. Female feeding habits are unknown.
Discussion: The presence of black females allows this species to be differentiated
from other Simulium (Ectemnaspis) species.

PERFLAVUM species group

Simulium (Ectemnaspis) perflavum ROUBAUD (Figs. 84 R; 85 K; 86 Q; 87 J; 89 J; 90 G, N, S.)


Simulium perflavum ROUBAUD, 1906a: 518; LUTZ, 1910: 230; PINTO, 1931: 478; D’ANDRETTA, 1956: 33-
38; RAMÍREZ PÉREZ, 1971: 301; 1983: 62; COSCARÓN & WYGODZINSKY, 1972: 223; PY-DANIEL,
1989: 254; HAMADA, 1998: 523; HAMADA & ADLER, 1998b: 207; 1999: 273-288; 2001: 119;
HAMADA & MC CREADY, 1999: 71; HAMADA & GRILLET, 2001: 35; ALENCAR et al., 2001: 561.
Simulium (Ectemnaspis) perflavum: COSCARÓN, 1984: 310; 1987: 29; 1990: 140-141; 1991: 257; STRIEDER
et al., 1992: 56.
Simulium (Eusimulium) perflavum: LUTZ, 1909: 131.
Simulium (Odagmia) perflavum: RUBTZOV, 1940: 532.
Eusimulium perflavum: SURCOUF & GONZALEZ RINCONES, 1911: 283; LANE & PORTO, 1939: 169-172;
LANE & VULCANO, 1943: 432.
Ectemnaspis perflava: PY-DANIEL & MOREIRA SAMPAIO, 1995: 118; STRIEDER , 2004: 129-130.
Simulium (Psilopelmia) perflavum: CROSSKEY & HOWARD, 1997: 59; 2004: 54.
Eusimulium antunesi LANE & PORTO, 1940: 185; LANE & VULCANO, 1943: 436.
Simulium mauense NUNES DE MELLO, 1974: 35.
Simulium nilesi RAMBAJAN, 1979: 63-77.
Female: Wing length 2.6-2.8 mm. Scutum and scutellum yellowish orange, metano-
tum, pleurae, sternites and abdomen yellowish brown. Abdominal pleurae grayish. Legs
yellowish with darker portion of cibarium covered by abundant acute teeth-like spines,
and projected medially with small median invagination (Fig. 84 R). Basal sector of R with
hair. Eighth sternite with about 25 setae; gonapophyses with internal margins divergent,
slightly produced distally and internal rim poorly sclerotized. Paraproct about 2 times
longer than wide at base, narrower on distal 3/4 (Fig. 85 K).
Male: Wing length 2.5-2.7 mm. Coloration similar to that of female but slightly
darkened. Ratio of length/width of hind basitarsus = 6.0-6.4. Gonostylus as long as
gonocoxite, subcylindrical, with apical spur (Fig. 86 Q). Ventral plate about 2 times longer
than width, with small lateral constrictions (Fig. 87 J).
Pupa: Cocoon with thick and compact weave, slightly projected basally; length along
base = 2.1-2.5 mm, on dorsum 2.6-3.0 mm, gills 2.0-2.5 mm. Frontoclypeus and thorax with
numerous tubercles; trichomes with 1 or 2 (3-4) branches. Gill with 8 branches arising very
close to base with very short basal trunk and without evident primary branches (Fig. 89 J); in
some areas (e.g., Corrientes, Argentina) branches darker and thicker.
Larva: Maximum length 6.5 mm. Cephalic apotome slightly darkened medially with 1
pair each of basal and sublateral negative spots (Fig. 90 G). Postgenal bridge shorter than half
of hypostomium length. Antennae not surpassing stem apex of cephalic fan; ratio of proximal,
410 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

media1 and distal segments = 1:1.7:1.4-1.6 (Fig. 90 N). Mandible with 1-2 latero-mandibular
processes. Anal ring with 78-87 rows with 13-14 hooks each. Anal sclerite with hair relatively
numerous, no scales. Rectal papillae with 10-15 diverticula on each lobe (Fig. 90 S).
Distribution: Argentina: Corrientes, Formosa, Misiones; Brazil: Acre, Amazonas,
Amapá, Bahia, Espírito Santo, Goiás, Mato Grosso, Minas Gerais, Pará, Paraná, Pernam-
buco, Rio de Janeiro, Rio Grande do Sul, Rondônia, Roraima, Santa Catarina, São Paulo;
Paraguay: Caaguazu, Cordillera; Venezuela: Amazonas, Bolivar, Carabobo.
Bionomics: Simulium perflavum is a tropical species breeding in small flows that are
not fast flowing, and that are frequently in open areas with high insolation. The water is
clear to black or muddy yellowish-red in areas that suffer deforestation, road construction,
and other degradation. Females are not anthropophilic.
Discussion: The yellowish coloration of the female, the male genitalia and the gill
branch arrangement differentiate this species from other Simulium (Ectemnaspis) species.
Cytologically analyzed populations from 10 states in Brazil and 1 in Venezuela represent a
single species (HAMADA & ADLER, 1999).

Simulium (Ectemnaspis) kabanayense RAMÍREZ PÉREZ & VULCANO


(Figs. 84 Q; 85 L; 86 P; 87 K; 89 K.)
Simulium (Psilopelmia) kabanayense RAMÍREZ PÉREZ & VULCANO, 1973: 390; CROSSKEY & HOWARD,
1997: 59; 2004: 53.
Simulium kavanayense (sic.): RAMÍREZ PÉREZ, 1983: 48.
Simulium kabanayense: HAMADA & ADLER, 1998b: 221; HAMADA & GRILLET, 2001: 34.
Simulium (Ectemnaspis) kabanayense: COSCARÓN, 1984: 312; 1987: 29; 1990: 141.
Female: Wing length 2.1-2.3 mm. Scutum orange with silvery lateral rim; scutellum
yellowish; pleurae yellowish brown; metanotum dark brown. Legs yellowish brown with
darker portion blackish. Abdomen with tergites I-II yellowish, III-IX blackish; tergal plate
III brown, V-IX shiny. Fronto-ocular triangle about as wide as long. Cibarium with area
between cornuae protruded, covered by acute teeth-like spines scarce at middle (Fig. 84
Q). Paraproct about as long as wide (Fig. 85 L).
Male: Wing length 2.0 mm. Coloration as in female, but abdominal tergite I
yellowish brown, the following blackish. Gonostylus relatively robust, about as long as
gonocoxite (Fig. 86 P); ventral plate with deep lateral constrictions (Fig. 87 K).
Pupa: Cocoon slightly projected anteriorly at base; weave compact with obvious
threads. Length along base 2.5 mm, on dorsum 2.3 mm, gills 2.0-2.5 mm. Frontoclypeal
and thoracic tubercles sparse with smooth surface; cephalic trichomes bifid, thoracic with
2-6 branches. Gill with 10 (12) branches, with branching close to base (Fig. 89 K).
Larva: (based on HAMADA & ADLER, 1998). Cephalic apotome without special
ornamentation; hypostomium with median tooth at same level as lateral teeth; postgenal
cleft pointed; cephalic fan with 40-49 rays; anal ring with 83-85 rows of 12-18 hooks each.
Distribution: Brazil: Roraima; Venezuela: Bolivar.
Bionomics: Unknown.
Discussion: Females of S. kabanayense are very similar to those of S. maroniense, S.
rorotaense and S. suarezi, but can be differentiated by the length of the paraproct. The male
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 411

of S. kabanayense can be distinguished by the yellowish brown abdominal tergites I-II, and
the pupa by the gill of 14-23 branches.

Simulium (Ectemnaspis) maroniense FLOCH & ABONNENC (Figs. 84 B; 88 V; 89 L.)


Simulium maroniense FLOCH & ABONNENC, 1946a: 9; RAMÍREZ PÉREZ, 1983: 53; SHELLEY et al., 1984:
153; HAMADA & ADLER, 1998b: 207; 1999: 277; 2001: 119; HAMADA & GRILLET, 2001: 35.
Simulium (Ectemnaspis) maroniense: COSCARÓN, 1984: 313; 1987: 29; 1990: 141.
Simulium (Psilopelmia) maroniense: SHELLEY et al., 2004: 455-458; CROSSKEY & HOWARD, 2004: 53.
Female and Male: Morphologically and ornamentally similar to that of S. rorotaense,
but female gonapophysis slightly longer, fronto-ocular triangle lower, and sensory vesicle
of maxillary palp about 1/3 of palpomere length (Fig. 84 B); male ventral plate with
deeper constrictions laterally.
Pupa: Cocoon similar to that of S. rorotaense. Frontoclypeal and thoracic tubercles
more abundant and larger than in S. rorotaense (Fig. 88 V); trichomes elongate with 3-6
branches. Gill with 18-20 filamentous branches that increase in length from dorsal to
ventral position (Fig. 89 L) and also are darker than in S. rorotaense.
Larva: (based on HAMADA & ADLER, 1998). Antenna with medial segment shorter
than distal.
Distribution: Brazil: Amapá, Roraima; French Guiana; Venezuela: Amazonas,
Bolivar.
Bionomics: Unknown.
Discussion: This species was synonymized by SHELLEY et al. (1984: 153) but was
revalidated by COSCARÓN (1990: 141) and HAMADA & ADLER (1998: 207), who confirmed
the validity of this species. Simulium suarezi is very similar; differences are explained in the
discussion of that species. HAMADA & ADLER (1999) found four cytotypes of S. maroniense,
showing the complexity of this species.

Simulium (Ectemnaspis) rorotaense FLOCH & ABONNENC


(Figs. 84 c; 85 M; 86 R; 87 L; 89 M; 90 H, K.)
Simulium rorotaense FLOCH & ABONNENC, 1946a: 4; PY-DANIEL, 1981: 175; RAMÍREZ PÉREZ, 1983: 66;
HAMADA & ADLER, 1998b: 221; 2001: 119 HAMADA & GRILLET, 2001: 35.
Simulium (Ectemnsapis) rorotaense: COSCARÓN, 1984: 313; 1987: 29; 1990: 141-143.
Simulium (Psilopelmia) rorotaense: SHELLEY et al., 2004: 455-458; CROSSKEY & HOWARD, 1997: 59; 2004: 54.
Simulium fulvinotum CERQUERA & NUNES DE MELLO, 1968: 205.
? Simulium wuayaraka ORTIZ, 1957: 163.
? Simulium (Psilopelmia) ignacioi RAMÍREZ PÉREZ & VULCANO, 1973: 387; RAMÍREZ PÉREZ, 1983: 8;
HAMADA & ADLER, 1998b: 207.
Female: Wing length 2.0-2.5 mm. Scutum, scutellum and metanotum yellowish-
orange, scutum with lateral borders nacreous and hair yellowish. Legs yellowish, darkened
mostly on apices of segments. Abdominal segments I, II (III) light brown-yellow, IV-IX
blackish. Sensory vesicle of maxillary palp longer than half of palpomere length (Fig. 84
c). Cibarium slightly elevated medially, very similar to that of S. suarezi and S. kabanayense.
412 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Basal portion of R with hair. Eighth sternite with 19-22 setae on each side; gonapophysis
slightly sclerotized on internal rim. Paraproct nearly 2 times longer than wide, acute
distally, with large microtrichiae on posterior border (Fig. 85 M).
Male: Wing length 2.0-2.2 mm. Coloration similar to that of female. Gonostylus
subcylindrical, slightly curved on distal third and about same length as gonocoxite (Fig. 86 R).
Ventral plate slightly shorter than width at base, with strong lateral depressions (Fig. 87 L).
Pupa: Cocoon weave not compact, with open areas especially near aperture Length
along base 2.7 mm. Frontoclypeus and thorax slightly sclerotized, tubercles not abundant,
trichomes with 2-3 branches. Gills with 18-20 thin filamentous branches, bifurcated at
variable distances from base (Fig. 89 M).
Larva: Maximum length 5.0 mm. Cephalic apotome with darkened median basal
area, without special ornamentation, and with several negative spots near basal border
(Fig. 90 H). Ratio of antennal segments: proximal, media1 and distal = 1:0.7:1.4. Postgenal
bridge about 1/5 of hypostomium height (Fig. 90 K). Mandible with 1 latero-mandibular
process. Cephalic fan with about 45 rays. Rectal papillae with 13-15 elongate diverticula on
each lobe. Body integument glabrous.
Distribution: Brazil: Amazonas; Venezuela: Amazonas, Bolivar.
Bionomics: This species breeds in clear, acidic creeks of fast flow and about 25 °C.
Larvae are found on aquatic vegetation. Females bite humans and cattle.
Discussion: This species is very similar to S. maroniense and S. suarezi .It can be
differentiated by the pupal gill, which viewed frontally looks like an open rosette; branch
bifurcation is closer to the base. Simulium wuayaraka was synonymized by RAMÍREZ PÉREZ
(1983) and SHELLEY et al. (1984); the female coloration and paraproct are similar, but the
cibarium morphology, according to the original drawings, is closer to that of S.
shewellianum and S. ochraceum. It would be useful to confirm the identity of this species with
more material from different life stages. Simulium ignacioi was synonymized by RAMÍREZ
PÉREZ (1983: 8) and accepted by SHELLEY et al. (1984) and according to the original
description, it is very similar to S. rorotaense, especially in the pupal gill, but the description
is limited. Cytological studies by HAMADA & ADLER (1998) indicate that it is a good species,
although morphological characters were not found to separate it from S. rorotaense.

Simulium (Ectemnaspis) suarezi RAMÍREZ PÉREZ, RASSI & RAMÍREZ


(Figs. 84 S; 85 N; 88 L; 89 N.)
Simulium (Psilopelmia) suarezi RAMÍREZ PÉREZ et al., 1977: 173; RAMÍREZ PÉREZ, 1983: 120; HAMADA &
ADLER, 1998b: 221; HAMADA & GRILLET, 2001: 35; CROSSKEY & HOWARD, 1997: 60; 2004: 54.
Simulium (Ectemnaspis) suarezi: COSCARÓN, 1984: 314-315; 1987: 29; 1990: 143.
Ectemnaspis suarezi PY- DANIEL & MOREIRA SAMPAIO, 1995: 118, 120.
Female: Wing length 2.3 mm. Scutum yellowish with silvery lateral borders.
Scutellum yellow. Metanotum dark brown. Pleurae and legs brownish yellow. Abdomen
with segment I and tergal plate II-III brownish yellow, IV-IX dark brown, V-IX shiny and
waxy. Fronto-ocular triangle wider than high. Cibarium with median elevation bearing
spines and central depression (Fig. 84 S). Claws with subbasal tooth well developed.
Paraproct nearly 2 times longer than wide at base, subtriangular ,acute apically (Fig. 85 N).
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 413

Male: Limited information is available. RAMÍREZ PÉREZ (1983: 120; Fig. 83) shows
gonostylus relatively thin, distally curved and longer than gonocoxite; ventral plate longer
than wide basally and with strong lateral constriction.
Pupa: Cocoon with non compact weave, with holes on anterior border and extended
covering laterally on gills (Fig. 88 L). Frontoclypeus and thorax with numerous wrinkled
tubercles; trichomes with several to 10 branches. Gill with 20 thick filamentous branches
emerging from basal trunk and, when viewed frontally, very open (Fig. 89 N).
Larva: (based on HAMADA & ADLER, 1998): Antenna with medial segment shorter
than distal. Anal ring with 98-101 rows with 16 hooks each. Cephalic fan with about 40
primary rays.
Distribution: Venezuela: Amazonas.
Bionomics: Unknown.
Discussion: The most similar species are S. maroniense and S. rorotaense. The pupal gill
filaments are thinner, and not open, with longer primary and secondary branches.

Simulium (Ectemnaspis) trombetense HAMADA, PY-DANIEL & ADLER (Fig. 89 O.)


Simulium (Ectemnaspis) trombetense HAMADA et al., 1999 (in HAMADA & ADLER, 1998b: 210); HAMADA &
ADLER, 2001: 119; HAMADA & GRILLET, 2001: 35; CROSSKEY & HOWARD, 2004: 54.
(Based on original description of HAMADA & ADLER, 1998).
Female: Wing length 2.2 mm. General coloration orange. Frons and clypeus with
silver blue pruinosity. Scutum orange with silvery lateral borders, hair golden-brown.
Pleurae brownish orange to brown with silver pruinosity. Scutellum and metanotum
orange, the latter with some brownish orange. Legs yellowish, dark brown on tibiae, fore
basitarsus and tarsomeres. Abdomen with tergites I-II orange, II with silver pruinosity,
III-VI greenish orange to gray, VI-VIII orange, shiny. Fronto-ocular triangle very flat.
Cibarium with 1+1 anterior submedian elevation covered by well-developed teeth. Basal
sector of R with setae, claws with strong subbasal tooth. Paraproct, longer than wide,
apically acute, similar to that of S. suarezi.
Male: Wing length 2.0-2.1 mm. Coloration as in female. Gonostylus 0.7 times as long
as gonocoxite, with strong curvature apically. Ventral plate with lateral constrictions, shape
very similar to that of S. kabanayense.
Pupa: Cocoon dark brown; weave parchment like with open loops anteriorly, threads
distinct; length along base 3.2 mm, on dorsum 3.4-3.7 mm, frontoclypeus and thorax with
relatively abundant rounded tubercles; large trichomes with 6-9 branches. Gills with 160-
250 thin short filaments arising from 7 thick trunks (Fig. 89 O).
Larva: Maximum length 6.0-8.0 mm. Cephalic apotome with brown anteromedian,
anterolateral and posterior spots. Postgenal cleft deep, tapered apically, postgenal bridge
about 1/4 of hypostomium length. Hypostomium with median tooth as long as lateral
teeth. Mandible with 1 latero mandibular process, thin, surpassing inferior margin of
mandible. Antenna as long as cephalic fan stem; ratio of proximal, media1 and distal
segment = 1:0.9-1.0:1.1-1.3. Cephalic fan with 45-50 rays. Anal ring with 89-113 rows with
14-16 hooks each. Rectal papillae with 11-13 elongate diverticula on each lobe.
Distribution: Brazil: Amapá, Amazonas, Pará, Roraima; Venezuela: Bolivar.
414 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Bionomics: This species breeds in forested areas of the Guiana Shield in fast
currents. Larvae are found on bedrock, fallen leaves and aquatic vegetation. Female
feeding habits are unknown.
Discussion: Adults of the S. perflavum species group are very difficult to separate.
Simulium perflavum can be separated by the absence of dark patches on the middle legs. The
pupal gill with over 100 branches is a very useful character to separate S. trombetense from
other closely related species.

DINELLII species group

Simulium (Ectemnaspis) dinellii JOAN (Figs. 84 T, Z; 85 O; 87 M; 88 W.)


Melusina dinellii JOAN, 1912: 363.
Simulium dinellii KNAB, 1913: 155; 1914b: 81; EDWARDS, 1933: 210; WYGODZINSKY, 1950: 77-87;
BARRETO, 1969: 31.
Simulium (Simulium) dinellii: ORFILA, 1935: 1532.
Eusimulium dinellii: PATERSON & SHANNON, 1927: 738; PINTO, 1931: 677; LANE & VULCANO, 1943: 434.
Eusimulium ochraceum: LANE & PORTO, 1939: 171 (nec WALKER); LANE & VULCANO, 1943: 434.
Dasypelmoza miniata ENDERLEIN, 1934b: 275.
Trichodagmia miniata ENDERLEIN, 1934a: 194; 1936: 129.
Simulium martinezi VARGAS, 1943a: 137; VARGAS, 1945a: 158; SMART, 1945: 508.
Simulium (Ectemnaspis) dinellii: COSCARÓN, 1984: 317-318; 1987: 29; 1990: 143; 1991: 258-260;
COSCARÓN ARIAS, 2003: 195, 198, 205, 215.
Simulium (Psilopelmia) dinellii: CROSSKEY & HOWARD, 1997: 58; 2004: 52.
? Simulium flavipictum KNAB, 1914b: 83.
Ectemnaspis dinellii: STRIEDER, 2004: 129.
Female: Wing length 1.6-2.7 mm. Scutum yellowish orange, a little lighter on anterolater-
al and posteromedian borders; scutellum light yellow, metanotum orange. Pleurae yellowish
orange, mesosternum with bluish pruinosity. Legs yellowish orange, with dark brown areas.
Abdomen with segments I-IV yellowish green, V-VII brownish green, VI-VII slightly lighter,
VIII-IX brownish yellow. Cibarium with median rounded invagination flanked by 1+1
submedian blunt projections covered by short spines (Fig. 84 T). Fronto-ocular triangle about
as long as wide. Sensory vesicle of maxillary palp about 1/4 palpomere length (Fig. 84 Z). Basal
sector of R with hair. Eighth sternite with about 27 setae; paraproct 1.5 longer than wide at
base (Fig. 85 O). Gonapophysis with internal border lightly sclerotized.
Male: Wing length 2.5-2.6 mm. Coloration as in female, with abdomen more orange
reddish, and 1+1 silvery spots on segments II, VI, VII and IX. Ratio of length/width of
hind basitarsus = 6. Gonostylus subcylindrical with distal portion subconical, curved
inwardly, and about 2/3 of gonocoxite length. Ventral plate about as long as wide at base,
with slight lateral constrictions (Fig. 87 M).
Pupa: Cocoon without anterior projection, thin weave, threads sparse and obvious;
length along base 3.1-3.7 mm, on dorsum 2.9-3.2 mm, gills 2.0-2.2 mm. Frontoclypeus
and thorax with medially sparse tubercles; trichomes with 3-5 branches (Fig. 88 W). Gills
with 8 branches, not bifurcating close to base, curved, resembling basket.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 415

Larva: Maximum length 6.0 mm. Cephalic apotome slightly darker basally, and
prolonged by 1+1 submedian stripes, with 1+1 light spots basally and on subapical portion of
submedian stripes. Postgenal cleft nearly reaching hypostomium base. Antenna about same
length as cephalic fan stem; ratio of proximal, medial and distal segments = 1:0.9:1.0. Cephalic
fan with 35-43 rays. Lateral sclerite of proleg with about 21 teeth arranged in 10 groups. Anal
ring with 76-82 rows, with 12-16 hooks each. Rectal papillae with 3-4 diverticula on each lobe.
Distribution: Argentina: Catamarca, Jujuy, La Rioja, Salta, Santiago del Estero,
Tucumán; Bolivia: Cochabamba, La Paz; Brazil: Minas Gerais, Rio Grande do Sul;
Colombia; Ecuador: Esmeraldas, Pichincha; Peru: Apurimac; Venezuela: Aragua.
Bionomics: Simulium dinellii breeds in small, clear watercourses, sometimes in warm
flows with high levels of Fe, Su and Cl. Females are anthropophilic and are considered a
dangerous species, biting on the face and arms.
Discussion: The most similar species is S. shewellianum, but it has dark brown legs,
and the cibarium has a more sclerotized rim with 1+1 submedian acute apical projections
without spines. Females of S. lutzianum, S. gabaldoni, S. albanense and most species of the S.
romanai group are similar, but they have a more flattened basal portion of the cibarium,
with tubercles on the median area and numerous spines between the tubercles and
cornuae. The males are mostly blackish. Simulium flavipictum has coloration and external
morphology similar to that of S. dinellii, although it is a little larger. Simulium flavipictum is
known from a female from Peru (Charape River). We synonymize S. flavipictum with
doubt, because of the unknown cibarium and genitalia.

Simulium (Ectemnaspis) shewellianum COSCARÓN


(Figs. 84 H, U; 85 P; 86 S; 87 N; 88 X; 89 P; 90 L.)
Simulium (Ectemnaspis) shewellianum COSCARÓN, 1984: 318-319; 1987: 30; 1990: 144; MUÑOZ &
COSCARÓN, 1999: 183; COSCARÓN ARIAS, 2003: 200, 205, 209, 215.
Simulium (Psilopelmia) shewellianum CROSSKEY & HOWARD, 1997: 58; 2004: 52.
Simulium shewellianum: PEPINELLI et al., 2003: 653.
Female: Wing length 1.9 mm. Scutum yellowish with silvery lateral borders, pleurae
brownish yellowish, with silvery pruinosity under certain illumination. Scutellum and
metanotum yellowish. Legs mostly dark brown with whitish area on mid and hind
basitarsus. Abdomen I-IV yellow, V-IX brown, tergite V velvety and VI-IX shiny.
Cibarium with well sclerotized rim, with 1+1 submedian acute projections and median
depressed area wrinkled (Fig. 84 U). Fronto-ocular triangle longer than wide. Sensory
vesicle of maxillary palp longer than half of palpomere (Fig. 84 H). Basal sector of R with
hair. Eighth sternite with about 20 setae on each side; gonapophysis more sclerotized
along inner margin. Paraproct slightly longer than wide with rounded apex (Fig. 85 P).
Male: Wing length 1.8 mm. Coloration similar to that of female. Ratio of length/
width of hind basitarsus = 6.6 Gonostylus very similar to that of S. dinellii with
pronounced curvature apically (Fig. 86 S), slightly shorter than gonocoxite. Ventral plate
with slight lateral constrictions (Fig. 87 N).
Pupa: Cocoon with weave compact, resistant, threads obvious; length along base and
dorsum 2.8 mm, gills 2.3 mm. Frontoclypeus and thorax with moderate quantity of
416 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

tubercles (Fig. 88 X); cephalic trichomes with 2-3 branches, thorax with 3-4 branches.
Gills with 8 branches, not bifurcated close to base (Fig. 89 P).
Larva: Maximum length 4.3-4.5 mm. Cephalic apotome with few ornamentations,
with median and sublateral areas slightly lighter and with simple hair. Postgenal bridge
absent or very narrow (Fig. 90 L). Cephalic fan with 30-34 rays. Ratio of antennal
segments proximal, medial and distal = 1:1.1:1.3. Mandible with 1 latero-mandibular
process. Lateral sclerite of proleg with 21 teeth arranged in 7 groups. Anal ring with 72
rows with 11-13 hooks each. Rectal papillae with 6-7 diverticula on each lobe. Eighth
segment with 1 pair of ventral papillae. Integument without hair.
Distribution: Brazil: São Paulo; Colombia; Ecuador: Los Rios, Pastaza, Pichincha.
Bionomics: This species breeds in small creeks with clear water, on aquatic
vegetation, at an altitude of about 400 m. Female feeding habits are unknown.
Discussion: Simulium lutzianum has similar adult coloration, pupal gill branches and
trichomes, but it is possible to distinguish it by the cibarial morphology. Simulium ochraceum
also shows some similarity but has 1+1 anterior oblique submedian silvery-whitish stripes
on the scutum. Similarity with S. dinellii was treated in the discussion of that species.

Simulium (Ectemnaspis) ochraceum WALKER (Figs. 84 E, V; 87 O.)


Simulium ochraceum WALKER, 1861: 332; FAIRCHILD, 1990: 706; VARGAS, 1942a: 238; 1946: 173.
Simulium (Simulium) ochraceum: DALMAT, 1955: 225.
Simulium (Eusimulium) ochraceum: HOFFMAN, 1931: 207 (in part).
Eusimulium (Eusimulium) ochraceum: BEQUAERT, 1934: 208.
Eusimulium ochraceum: LANE & VULCANO, 1943: 437.
Psilopelmia ochraceum: RUBTZOV & GARCÍA AVILA, 1972: 15.
Simulium (Psilopelmia) ochraceum: CROSSKEY & HOWARD, 1997: 58; 2004: 52.; SHELLEY et al., 2002: 164.
Simulium (Ectemnaspis) ochraceum: COSCARÓN, 1984: 320; 1987: 30; 1990: 147; COSCARÓN ARIAS, 2003:
195, 200, 205, 215, 218.
Simulium bipunctatum MALLOCH, 1912: 650.
Simulium (Psilopelmia) bipunctatum: SHELLEY et al., 1989: 130; 1997: 28.
Simulium (Ectemnaspis) bipunctatum: COSCARÓN, 1984: 321; 1987: 29; 1990: 144.
Simulium wolcotti FOX, 1953: 138.
? Simulium scutellatum LANE & PORTO, 1940: 192; LANE & VULCANO, 1943: 434 n. syn.
Simulium (Ectemnaspis) scutellatum: COSCARÓN, 1990: 145.
Female: Wing length 1.9-2.3 mm. Scutum yellow, to yellowish red with 1+1
anterolateral silvery spots continued by short submedian stripes not reaching to half of
scutum length (Fig. 84 E) but in accordance with DALMAT material from Guatemala, it
reaches the prescutellar border. Scutellum yellow, pleurae yellowish brown, showing
silvery tones on basal half under certain illumination. Metanotum dark brown, silvery
posteriorly. Legs yellowish, darkened on apex of femora and tibiae of hind and mid leg,
tibiae and tarsomeres dark brown. Abdomen with segments I-IV yellowish to brownish
yellow as also lateral borders of V; VI-IX dark grayish brown; V tergite with velvety
appearance, VI-IX shiny, waxy. Frons slightly divergent; fronto-ocular triangle about as
long as wide. Cibarium with basal portion well sclerotized, with 1+1 submedian apically
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 417

acute processes, with rounded median area, and rim between cornuae wrinkled or with
small denticle-like spines (Fig. 84 V). Sensory vesicle of maxillary palp surpassing half of
palpomere length. Basal sector of R with hair. Claws with well-developed subbasal tooth.
Eighth sternite with about 15 setae on each side. Gonapophysis with slight sclerotization
on internal rim. Paraproct about as long as wide at base, similar to Simulium shewellianum.
Male: Wing length 1.9-2.0 mm. Coloration as in female but darker ochre and silvery
submedian stripes shorter. Gonostylus elongated, subcylindrical, curved distally with
apical spur and slightly shorter than gonocoxite length. Ventral plate mostly shorter than
wide at base (Fig. 87 O) and with small lateral constrictions.
Pupa: Cocoon compact, slightly thicker dorsally, threads evident. Length along base
2.7-3.8 mm, on dorsum 2.5-3.3 mm, gills 1.7-2.5 mm. Frontoclypeus and thorax with since
numerous to moderate number of tubercles; trichomes with 2-4 branches. Gill with 8
branches arising close to base.
Larva: Maximum length 4.4-5.0 mm. Cephalic apotome with median dark spot wider
basally. Postgenal cleft reaching hypostomium base, postgenal bridge generally absent.
Cephalic fan with 28-36 rays. Antenna about as long as or a little longer than length of
cephalic fan stem; ratio of proximal, medial and distal segments = 1:1.1:1.3. Anal sclerite
with or without sparse hairs. Anal ring with 76-78 rows of 14-15 hooks each. Rectal papillae
with about 8 elongate diverticula on each lobe. Ninth segment with 2 ventral tubercles.
Distribution: Brazil: Amazonas, Roraima; Colombia: Cundinamarca; Costa Rica;
Cuba; Dominican Republic; Ecuador: Esmeraldas, Pichincha; Guatemala Amatitlan,
Chimaltenango, Escuintla, Suchitepequez; Mexico: Chiapas, Oaxaca, Veracruz; Panama:
Chiriqui, Coclé, Colón; Peru; Puerto Rico; Venezuela: Aragua, Carabobo, Monagas.
Bionomics: Simulium ochraceum breeds in small non torrential creeks and water courses
on rocks, decayed leaves and aquatic vegetation. Females bite humans and cattle. It is a
vector of Onchocerca volvulus in Mexico and Guatemala.
Discussion: Simulium ochraceum is very similar to other members of the S. dinellii
species group. Simulium antillarum (synonymized by SHELLEY et al., 1987), with 1+1
submedian silvery spots can be differentiated by the scutum silvery stripes more elongated
reaching or surpassing the half of scutum length, metanotum yellowish, the cibarium with
a medially depressed area without spines and a thicker basal rim medially, and male with
ventral plate more elongated and thinner gonostylus. Simulium shewellianum can be
differentiated by the absence of 1+1 submedian silvery stripes on the scutum, and the
cibarium medially wrinkled. Simulium ochraceum shows variation in the ventral plate, such as
lack of lateral constrictions, especially in Mesoamerican specimens that also have an
absence of or very scarce tubercles on the frontoclypeus and thorax. Additional variation
includes 1+1 submedian silvery stripes reaching the silver posterior area, and larval
cephalic apotome darkened medially and basally (DALMAT, 1955). Three cytotypes are
known (HIRAI & HUEMOTO, 1983) in Guatemala and Mexico. Probably the morphological
variation corresponds to specific cytological or molecular differences. In accordance with
the great variation, S. ochraceum can be considered a species complex.
Simulium scutellatum from Colombia, known only from females, has similar coloration
and cibarium morphology. Here it is synonymized with doubts. Simulium wolcotti, known
from adults, falls morphologically into the range of variation for S. ochraceum.
418 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Simulium (Ectemnaspis) antillarum JENNINGS (Figs. 84 D, W; 87 P.)


Simulium antillarum JENNINGS, 1915: 200.
Simulium (Psilopelmia) antillarum: DÍAZ NÁJERA, 1961: 79; STONE, 1969: 312-313; COSCARÓN, 1984:
319; 1987: 29; 1990: 146.
Simulium (Psilopelmia) pseudoantillarum RAMÍREZ PÉREZ & VULCANO, 1973: 379.
Simulium pseudoantillarum: RAMÍREZ PÉREZ, 1983: 63.
Psilopelmia antillarum: RUBTSOV & GARCIA AVILA, 1972: 11-15.
Female and Male: Scutum yellowish with 1+1 submedian silvery spots, reaching or
surpassing the half of scutum length (Fig. 84 D), abdomen with segments I-IV yellowish
and the following brownish gray,. Cibarium with smooth median area between submedian
processes, without spines or wrinkled sclerotized area, bordered basally by thick rim (Fig.
84 W). Male ventral plate longer than wide, with small lateral constrictions (Fig. 87 P).
Pupa: Slight differences from S. ochraceum: frontoclypeus and thorax with moderate
number of tubercles, more abundant and arranged in small groups, and trichomes with 2-
6 branches. Gill branching near base (DÍAZ NÁJERA, 1961) or primary and secondary
branches bifurcated at further distance, as in S. shewellianum.
Larva: Similar to that of S. ochraceum; cephalic apotome can show some lighter lateral areas.
Distribution: Colombia: Amazonas, Cundinamarca, Valle; Cuba; Dominica; Ja-
maica; Mexico: Chiapas; Ecuador: Galápagos; Peru: Cuzco, Huanuco; Puerto Rico;
Venezuela: Monagas, Sucre.
Bionomics: This species breeds in small fast-flowing creeks. Larvae are found on
fallen leaves, roots, aquatic vegetation and stones. Females are anthropophilic.
Discussion: The typical cibarium is smooth medially, which separates it from S.
ochraceum and S. shewellianum; this cibarium is very close to that of S. dinellii, but it has the
1+1 submedian processes thinner and more elongated, with a narrow basal rim. Also, in S.
dinellii and S. shewellianum the scutum has no submedian silvery stripes. Simulium
pseudoantillarum is in accordance with material from Peru, Colombia and Venezuela, but
material from Mexico (DÍAZ NÁJERA 1961) has a pupal gill with branching close to the
base. Material from Peru and Galápagos, which we now are considering S. antillarum, has
a few differences: a yellow ochreous scutum with thicker silvery submedian stripes. (Fig.
84 D). Surely, here we have another species complex.

Simulium (Ectemnaspis) nuneztovari RAMÍREZ PÉREZ, RASSI & RAMÍREZ


Simulium nuneztovari RAMÍREZ PÉREZ et al., 1977: 169; RAMÍREZ PÉREZ, 1983: 59.
Simulium (Ectemnaspis) nuneztovari: COSCARÓN, 1984: 316; 1987: 29; 1990: 145.
Simulium (Psilopelmia) nuneztovari: CROSSKEY & HOWARD, 1997: 58; 2004: 52.
(Information based on RAMÍREZ PÉREZ et al ., 1977 and RAMÍREZ PÉREZ, 1983).
Female: Wing length 2.1 mm. Scutum orange with decumbent hair dark golden; no
silvery border. Scutellum yellow-lemon with black hair. Metanotum and pleurae yellowish
with nacreous or silvery pruinosity. Legs yellowish with tibiae and tarsomeres dark brown.
Abdomen yellowish lemon. Fronto-ocular triangle wider than high. Cibarium with 1+1
short blunt submedian projections, covered with some small teeth, as is rim from
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 419

submedian process to cornuae, similar to that of S. antillarum. Paraproct about 2 times


longer than wide at base (as in S. bicoloratum).
Male: Wing length 2.0 mm. Coloration as in female, abdomen with inferior border
of VII-VIII brownish. Gonostylus subcylindrical, shorter than gonocoxite; ventral plate
with small lateral constrictions as in S. dinellii.
Pupa: Cocoon with weave relatively open, especially on sides; length along base 3.0
mm, on dorsum 2.3 mm. Frontoclypeus and thorax with numerous tubercles; trichomes
with 3-8 branches. Gill with 8 branches, with primary and very short secondary branches
Larva: Unknown.
Distribution: Venezuela: Amazonas.
Bionomics: Unknown.
Discussion: The closest species are S. dinellii and S. antillarum. The first can be
differentiated by the cibarial submedian process, which is thinner and more elongated,
and the abdomen with greenish brown to yellowish brown on segments VI-IX. Simulium
antillarum has a similar cibarium but has 1+1 silvery submedian stripes on the scutum.
Probably by examining more material, it can be established that this species is a variant of
the aforementioned species.

Species inquirenda

Simulium varians LUTZ


Simulium (Eusimulium) varians LUTZ, 1909: 131.
Simulium varians: LUTZ, 1910: 234; PINTO, 1931: 672; COSCARÓN, 1984: 322; 1990: 148.
Discussion: Simulium varians was described based on 3 females characterized by
being dark brown with orange spots from Brazil: São Paulo, Rio Grande, Franca. This
material was not found in Instituto Oswaldo Cruz. During a search in the type area,
material with this coloration was not collected. It will be necessary to repeat the search for
additional material. At present, S. varians is considered species inquirenda.

Simulium (Psilopelmia) ENDERLEIN (Figs. 91-100, 149.)


Type-species: Psilopelmia rufidorsum ENDERLEIN, 1934b = Simulium escomeli ROUBAUD, 1909.
Lanea VARGAS, MARTÍNEZ PALACIOS & DÍAZ NÁJERA, 1946: 107.
(Type-species Simulium haematopotum: MALLOCH, 1914).
Female: Medium-sized species, wing length 1.5-3.1 mm; general coloration from
yellowish brown to grayish brown; scutum yellowish to brown blackish, generally with
1+1 silvery to gray bluish submedian longitudinal vittae or 1 median brown reddish;
scutellum from yellowish to brown yellowish, metanotum from yellowish brown to
grayish brown; abdominal dorsum generally with median, 1+1 submedian and frequently
1+1 lateral subquadrate darker rows of spots; fronto-ocular triangle generally higher than
wide; sensory palpal vesicle less than half palpomere length.
Basal portion of cibarium with 1+1 short blunt submedian elevations, covered by
short denticles or spiculae, flanking median depressed area, frequently well sclerotized
420 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

halfmoon shaped; basal section of R generally unhaired. Sc with or without hair; claws
generally without or with very short subbasal tooth; paraprocts with basal lobe and distally
generally projected as anterior thin process; gonapophysis slightly divergent inner
sclerotized margin, without hair or with sparse microtrichiae; genital fork with sclerotized
stem and well-developed anterior branches.
Male: Wing 1.9-2.7 mm; coloration as in female but slightly darker; scutum
frequently with 1+1 silvery cuneiform stripes reduced to anterior half anterior or reaching
light posterior area; abdomen with variants yellowish, gray to blackish; posterior basitarsus
ratio length/width from 3.8-7.8; gonostylus shorter than gonocoxite, subtriangular to
subtrapezoidal, frequently wide distally, projected on internal side, and with apical spur;
ventral plate in ventral position wider than high, laterally with or without slight concavity;
endoparameres with strong hooks.
Pupa: Cocoon resistant, weaves with threads evident; slipper shaped, generally with
anterior reinforced border and mild anterior basal projection; gill branches generally in
number of 8 but can be 4-6 or 20, all forward directed; frontoclypeus and thoracic
trichomes mostly single or bifid; abdomen terminal spur small.
Larva: Maximum length 6.0 mm; integument without trichomes, spines or scales;
cephalic apotome scarcely ornamented, sometimes slightly darkened as median basal bell
shape with 1 median and 1+1 lateral subbasal light windows; antenna generally with medial and
distal segments similar to or longer than proximal; postgenal cleft moderately deep; postgenal
bridge length about as long as or longer than hypostomium height; anal sclerite with 63-105
rows of 10-17 hooks each; diverticula of rectal papillae generally 0-4 but can reach 9-12.
Simulium (Psilopelmia) is a relatively homogeneous taxa composed of 24 species. The
Neotropical species and one inquirenda are subdivided into two species groups: ESCOMELI
with 21 species and BLANCASI with three species. Simulium panamense, S. callidum, and S.
pseudocallidum have some unique characters and resemble S. (Ectemnaspis) species. The subgenus
Psilopelmia has different arrangements according to CROSSKEY & HOWARD (1997, 2004),
including five species groups: DINELLII, PERFLAVUM, ROMANAI, ESCOMELI and
QUADRIVITTATUM. The first three are considered to belong to S. (Ectemnaspis) by
COSCARÓN (1987: 19); both taxa have a close relationships that some day might lead to
including all of the groups in the same subgenus. We maintain S. (Escomeli) as a separate species
group from S. (Psilopelmia), being reinforced by the results of cladistic studies that separate
them. To S.(Psilopelmia) the S. (Blancasi) species group is also added.

KEYS TO SIMULIUM (PSILOPELMIA) SPECIES

Females
1 Scutum uniformly dull gray, without stripes ..................................................................
......................................................................................... BLANCASI species group......2
– Scutum with 1 median or 1+1 longitudinal stripes ......................................................
......................................................................................... ESCOMELI species group......4
2(1) Abdominal tergites III-V velvety black, their center anteriorly (Fig. 98 A) with or
without transverse dumbbell-shaped light-colored spots; setae of scutum sparse,
not conspicuous under low magnification ......................................... Simulium blancasi
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 421

– Abdominal tergites III-V brown or piceous, always without light-colored spots


anteriorly (Fig. 99 A); setae of scutum very abundant, conspicuous under low
magnification ....................................................................................................................... 3
3(2) Abdominal tergites III-V entirely dark velvety brown (Fig. 99 A); calcipala
comparatively large, broadly rounded apically (Fig. 99 D) ............ Simulium tenuipes
– Abdominal tergites III-V dark anteriorly, distinctly lighter posteriorly, entire
posterior half light colored on V; calcipala much reduced, subtriangular (Fig. 100 A)
......................................................................................................................... Simulium penai
4(1) Scutum grayish with 1 median longitudinal reddish brown band (Fig. 91 A) or
unicolorous; cibarium with well-defined small teeth on basal portion (Fig. 91 J) ...... 5
– Scutum with 1+1 light submedian bands generally from anterior to posterior
border (Figs. 91 B-E); cibarium with teeth, spiculate or smooth on basal portion
(Figs. 91 L-Q) ...................................................................................................................... 7
5(4) Metanotum blackish; wing length 1.9-2.4 mm ........................ Simulium mediovittatum
– Metanotum grayish brown; wing length 1.8-2.2 mm .................................................. 6
6(5) Wing length 2.0-2.2 mm; scutellum grayish brown; abdominal tergite II grayish
(Fig. 91 G) .................................................................................................... Simulium dugesi
– Wing length 1.8 mm; scutellum yellow to yellowish brown; abdominal tergite II
grayish yellow .............................................................................................. Simulium ochoai
7(4) Abdominal tergites without rows of darker subquadrate spots; distal portion of
paraproct elongated, narrowing slightly (Figs. 92 F, G); claws with well developed
subbasal tooth (Figs. 91 V, W) ......................................................................................... 8
– Abdominal tergites with 1 median, 1+1 submedian and sometimes 1+1 sublateral
rows of darker subquadrate spots at least from segments III to VI (Figs. 91 H, I);
distal portion of paraproct relatively short (Figs. 92 B-E, J) or elongated,
narrowing abruptly and projected as narrow appendix (Figs. 92 H-N); claws
without subbasal tooth (Figs. 91 T-U) ......................................................................... 10
8(7) Cibarium with small median depression and abundant denticles submedially (Fig.
91 N); paraproct distally elongated, as long as basal width and acuminate distally
(Fig. 92 F) ............................................................................................. Simulium panamense
– Cibarium with thick rim, with 1+1 slight sublateral protuberances sometimes
covered by thin spiculae (Fig. 91 O); distal process of paraproct absent and blunt
apically (Fig. 92 G) ............................................................................................................. 9
9(8) Basal sector of R with hair .................................................................. Simulium callidum
– Basal sector of R glabrous ......................................................... Simulium pseudocallidum
10(7) Abdominal tergites with 1 median and 1+1 submedian rows of subquadrate
darker spots (Fig. 91 H); paraproct distal process shorter than half of basal width
(Figs. 92 B, E, J) ................................................................................................................ 11
– Abdominal tergites with 1 median, 1+1 submedian and 1+1 sublateral rows of
subquadrate darker spots (Fig. 91 I); distal process of paraproct length about equal
to width at base (Figs. 92 C, D) or very elongated (Figs. 92 H, I, K-N) ................ 13
11(10) Scutellum grayish brown; sensory vesicle of maxillary palp shorter than half
length of palpomere (Fig. 91 S); paraproct blunt apically (Fig. 92 E) ......................
........................................................................................................... Simulium haematopotum
422 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

– Scutellum yellowish brown; sensory vesicle of maxillary palp generally longer


than half of length of palpomere; paraproct slightly acuminate apically (Figs. 92
B-J) ...................................................................................................................................... 12
12(11) Wing length 2.1-2.7 mm .................................................................... Simulium bivittatum
– Wing length 2.4-3.0 mm .................................................................... Simulium dandrettai
13(10) Paraproct apically blunt and distal process about as long as half of basal width
(Fig. 92 D); scutum yellowish orange with very thin 1+1 submedian silvery stripe
(Fig. 91 B) .......................................................................................................................... 14
– Paraproct apically acuminate and distal process as long as basal width (Figs. 92 H,
I, K-N); scutum yellowish gray with brown to blackish medial sublateral stripe
(Figs. 91 E, F) (except Simulium downsi) ....................................................................... 15
14(13) Wing length 1.5-1.6 mm; sensory vesicle of maxillary palp about as long as half
of palpomere (Fig. 91 R) ................................................................ Simulium iracouboense
– Wing length 1.8-2.1 mm; sensory vesicle of maxillary palp shorter than half of
palpomere ..................................................................................................Simulium samboli
15(13) Basal sector of R with hairs; scutum yellowish with light yellowish brown median
and sublateral stripes ................................................................................ Simulium downsi
– Basal sector of R glabrous; scutum yellowish gray with dark brown to black
median and sublateral stripes ........................................................................................ 16
16(15) Scutum generally yellowish with light brown yellowish to grayish yellow stripes, or
with dark brown to blackish on median and sublateral stripes (Fig. 91 E) .............
..................................................................................................................... Simulium escomeli
– Scutum grayish yellow with dark brown to blackish on median and sublateral
stripes (Fig. 91 F) ............................................................................................................. 17
17(16) Distal process of paraproct shorter than half width of base (similar to Simulium
dandretta ) (Fig. 92 J) ..........................................................................Simulium zempoalense
– Distal process of paraproct length, as wide as width of base (Figs. 92 K-N)
.............................................................................................................................................. 18
18(17) Scutum with median black vittae wider than submedian silvery vittae; sublateral
black vittae not reaching anterior rim (Fig. 91 F) ..................... Simulium mangabeirai
– Scutum with median black vittae about same width as submedian silvery vittae;
sublateral black vittae reaching or very close to anterior rim .....................................
........................... Simulium longithallum, Simulium gonzalezherrejoni, Simulium trivittatum,
Simulium bobpetersoni, Simulium veracruzanum

Males
1 Scutum uniformly dull gray without silvery wedges ....................................................
......................................................................................... BLANCASI species group......2
– Scutum with 1+1 silvery wedges (Figs. 93 A-G) ...........................................................
......................................................................................... ESCOMELI species group......4
2(1) Abdominal tergites III-V velvety black, their anterior border with distinct light-
colored spot at center (Fig. 98 P); setae of scutum sparse, not conspicuous under
low magnification; genitalia with ventral plate constricted laterally near base (Fig.
98 W) ......................................................................................................... Simulium blancasi
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 423

– Abdominal tergites III-V dark, sometimes with lighter regions posteriorly but not
on anterior border; setae of scutum very abundant, conspicuous under low
magnification; genitalia with ventral plate not constricted laterally near base (Fig.
99 J) ....................................................................................................................................... 3
3(2) Length of hind basitarsus more than 4 times its width; calcipala much reduced,
subtriangular (Fig. 100 A) .......................................................................... Simulium penai
– Length of hind basitarsus less than 4 times its width; calcipala not as reduced,
broadly rounded apically (as in female, Fig. 99 D) .......................... Simulium tenuipes
4(1) Scutum and abdomen yellowish, abdomen can be grayish brown but at least with
some yellowish orange portion; scutum with submedian silvery wedges only on
anterior area (Fig. 93 B) .................................................................................................... 5
– Scutum and abdomen gray to blackish; scutum with silvery wedges with variable
extension (Figs. 93 A, C-G) ........................................................................................... 10
5(4) Body of ventral plate subovoidal (Fig. 94 F) ............................................................... 6
– Body of ventral plate of variable shape but not subovoidal .................................... 7
6(5) Basal sector of R with hair .................................................................. Simulium callidum
– Basal sector of R glabrous ......................................................... Simulium pseudocallidum
7(5) Ventral plate with strong lateral constrictions on 1/2 of body, continued by
narrow subovoidal mediodistal portion (Fig. 94 E) ..................... Simulium panamense
– Ventral plate variable, but without strong lateral constrictions on body (Figs. 94 C, H)
................................................................................................................................................ 8
8(7) Gonostylus subtrapezoidal with strong expansion on subterminal external side
(Fig. 93 N); ventral plate of body reduced, subtriangular (Fig. 94 H) and narrower
than position of branches ....................................................................... Simulium downsi
– Gonostylus without lateral or subterminal expansion (Figs. 93 K, L); ventral plate
body more developed, not subtriangular and wider than position of branches (Fig.
94 C) ..................................................................................................................................... 9
9(8) Gonostylus subtriangular (Fig. 93 J); ratio of length/width of posterior basitarsus
= 7.8 ........................................................................................................... Simulium samboni
– Gonostylus subtrapezoidal (Fig. 93 K); ratio of length/width of posterior
basitarsus = 6.6 ................................................................................. Simulium iracouboense
10(4) Scutum with 1+1 short silvery stripes not reaching posterior declivity (Fig. 93 A)
.............................................................................................................................................. 11
– Scutum with 1+1 long silvery stripes reaching posterior declivity (Fig. 93 G) ......... 19
11(10) Scutum with 1+1 submedian silvery blue areas triangular to oval shaped,
contiguous anteriorly with same-colored lateral areas (Fig. 93 A) ........................ 12
– Scutum with 1+1 silvery blue areas not contiguous with same-colored lateral areas
(Fig. 93 D) .......................................................................................................................... 18
12(11) Silvery submedian wedges strongly tapered posteriorly, (Fig. 93 A) surpassing or
not half of scutum; gonostylus about as long as width of external straight side
(Fig. 93 I); ventral plate subtriangular with rounded apex (Fig. 94 A) ................. 13
– Silvery submedian wedges generally surpassing half of scutum; gonostylus expanded
over external distal subterminal side (Figs. 93 P, Q, S); ventral plate variable, from
subtriangular with or without lateral constrictions to subovoidal ............................... 15
424 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

13(12) Submedian wedges of scutum not reaching posterior declivity (Fig. 93 A) ....... 14
– Submedian wedges of scutum strongly tapered posteriorly and reaching posterior
declivity ........................................................................................... Simulium mediovittatum
14(13) Wing length 1.9 mm .................................................................................. Simulium dugesi
– Wing length 1.6 mm .................................................................................. Simulium ochoai
15(12) Gonostylus less than 2 times as long as wide at base, distally expanded, wider than
at base (Fig. 93 O); ventral plate without lateral constrictions, with rounded
borders (Fig. 94 I); ratio of length/width at base = 1.2 ............ Simulium dandrettai
– Gonostylus generally over 2 times longer than width at base, not expanded
distally; ventral plate usually without rounded borders; ratio of length/width over
1.2 ........................................................................................................................................ 16
16(15) Gonostylus subrectangular, with subparallel sides (Fig. 93 I); ventral plate
relatively short without lateral constrictions (Fig. 93 B) ............. Simulium bivittatum
– Gonostylus subtrapezoidal, with strong curvature distally and slightly acuminate
distally (Fig. 93 N); ventral plate from short and wide (Fig. 94 J) to more elongated
with lateral constrictions (Fig. 94 G) ........................................................................... 17
17(16) Ventral plate relatively short with base projected basally (Fig. 94 J), ratio of
length/width = 2.3; wing length 1.9-2.1 mm .............................. Simulium bobpetersoni
– Ventral plate relatively long, with base not projected basally and laterally with
slight constrictions (Fig. 94 G); ratio of length/width = 1.3-1.4 wing length 2.0-
3.0 mm ....................................................................................................... Simulium escomeli
18(11) Ventral plate subtriangular without lateral constrictions (Fig. 94 D); gonostylus
strongly enlarged distally, with long curved spur (Fig. 93 L); wing length 1.8-2.1
mm ................................................................................................... Simulium haematopotum
– Ventral plate with strong lateral constrictions and distal portion after constriction
narrow, about 1/3 width of body (Fig. 94 K); gonostylus not very enlarged distally
and with moderate-sized spur (Fig. 93 P); wing length 2.6-3.0 mm .........................
............................................................................................................ Simulium veracruzanum
19(10) Gonostylus about 2/3 as wide as long (Fig. 93 R); ventral plate without lateral
constrictions; distal border of body subcircular ........................ Simulium zempoalense
– Gonostylus about 1/2 as long as wide (Fig. 93 Q); ventral plate with or without
constrictions and different than above (Figs. 94 L-N) ............................................. 20
20(19) Hind basitarsus wide, ratio of length/width = 3.8; wing length 2.5-2.8 mm;
scutum with 1+1 narrow light wedges not reaching posterior edge (Fig. 93 F) ....
.............................................................................................................. Simulium mangabeirai
– Hind basitarsus narrower with ratio of length/width >5.0; scutum with 1+1 wide
light wedges reaching posterior edge (Fig. 93 G) ...................................................... 21
21(20) Abdominal segments I-II yellowish; wing length 2.5 mm .......... Simulium trivittatum
– Abdominal segments I-II not yellowish; wing length 2.0-2.5 mm ........................ 22
22(20) 1+1 blackish lateral stripes of scutum not reaching postpronotal lobes (Fig. 93 G);
wing length 2.5 mm .................................................................. Simulium gonzalezherrejoni
– 1+1 blackish lateral stripes of scutum reaching postpronotal lobes; wing length
2.0 mm ................................................................................................ Simulium longithallum
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 425

Pupae
1 Gill with 20 branches (Fig. 96 D) .................................................... Simulium panamense
– Gill with 4-8 branches ....................................................................................................... 2
2(1) Gill with 4 branches (Fig. 96 M) ..................................................................................... 3
– Gill with 6-8 branches ....................................................................................................... 6
3(2) Cocoon well developed covering most of thorax and head (Fig. 95 M); ventral
anterior edge more advanced than dorsal ................................... Simulium mangabeirai
– Cocoon scarcely developed, not covering most of thorax and head; dorsal
anterior edge more advanced than ventral edge (Fig. 98 X) ......................................
......................................................................................... BLANCASI species group......4
4(3) Gill with short trunk followed by 2 distinct secondary branches bearing 2 + 2
filaments (Fig. 98 Y) ............................................................................... Simulium blancasi
– Gills with all 4 filaments seemingly arising from very short common trunk (Figs.
99 L, 100 C) ......................................................................................................................... 5
5(4) Frontoclypeus and exposed portion of thorax with abundant tubercles; surface of
cocoon rough, its color brown when examined in liquid ............... Simulium tenuipes
– Frontoclypeus and exposed portion of thorax with or without very sparse
tubercles (Figs. 100 D, E); surface of cocoon smooth, its color brown with leaden
sheen, parchment-like, when examined in liquid ................................. Simulium penai
6(2) Gill with 6 branches .......................................................................................................... 7
– Gill with 8 branches ........................................................................................................ 10
7(6) Gill with wrinkled surface; blunt apex, brown to blackish pigmentation (Figs. 96 I,
J); gill branches shorter than cocoon length (Figs. 95 I, J) ....................................... 8
– Gill with smooth surface, generally acuminate apex; yellowish pigmentation; gill
branches as long as or longer than cocoon length (Fig. 95 N) ................................ 9
8(7) Gill secondary dorsal outer and inner branches thicker than other branches (Fig.
96 I); frontoclypeal and thoracic tubercles smooth; cocoon without longitudinal
ridges (Fig. 95 I) .................................................................................. Simulium dandrettai
– Gill with only dorsal outer branch thicker than other branches (Fig. 96 J);
frontoclypeal and thoracic tubercles wrinkled; cocoon with several longitudinal
ridges (Fig. 95 J) ................................................................... Simulium sp. near dandrettai
9(7) Gill with branches off of primary and secondary branches very close to base (Fig.
96 Q); dorsal thoracic integument with sparse tubercles but with small rugosities
simulating tubercles (Fig. 95 O) .....................................................Simulium zempoalense
– Gill with branches off of primary ventral and secondary branches not close to
base (Fig. 96 P); dorsal thoracic integument smooth with sparse tubercles ...........
................................................................................................................. Simulium trivittatum
10(6) Gill with basal stem short, with primary and secondary branches emerging very
close to base, dorsal primary branch thicker than others (Figs. 96 E, F) ............ 11
– Gill with primary and secondary branches emerging from basal stem more distant
from base, and with primary branches about same diameter ................................. 12
11(10) Gill tertiary dorsal branch distinctly thicker and 1/3 longer than other branches
(Fig. 96 F) ...................................................................................... Simulium pseudocallidum
426 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

– Gill tertiary dorsal branch only slightly thicker and with similar length to other
branches (Fig. 96 E) ............................................................................... Simulium callidum
12(10) Gill dorsal secondary and tertiary branches thicker than other branches (Figs. 96
N, O) ................................................................................................................................... 13
– Gill dorsal secondary and tertiary branches with diameter similar to that of other
branches ............................................................................................................................. 14
13(12) Gill dorsal secondary and tertiary branches arising far from base (Fig. 96 N) ......
.......................................................................................................Simulium gonzalezherrejoni
– Gill dorsal secondary and tertiary branches arising near base (Fig. 96 O) ..............
.............................................................................................................. Simulium longithallum
14(12) Gill length as long as or longer than cocoon length (Fig. 95 L) ................................
............................................................................................................ Simulium veracruzanum
– Gill length shorter than cocoon length (Figs. 95 A-G) ........................................... 15
15(14) Cocoon weave sparse, with very open weave, without anterior rim (Fig. 95 F) .....
.............................................................................................................. Simulium iracouboense
– Cocoon weave compact, with threads very close, and with anterior rim (Figs. 95
A-E, G, H, K) ................................................................................................................... 16
16(15) Cephalic and thoracic trichomes longer than half of frontoclypeal basal width
(Fig. 95 R); cocoon anterior rim reinforced and with anterodorsal projection (Fig.
95 E) .......................................................................................................... Simulium samboni
– Cephalic and thoracic trichomes not longer than half of frontoclypeal basal width
(Figs. 95 Q, T, U); cocoon anterior rim usually not reinforced and without
anterodorsal projection ................................................................................................... 17
17(16) Cocoon with small dorsal and ventral projection (Fig. 95 G); frontoclypeus and
thorax with sparse tubercles (Fig. 95 T) ................................... Simulium haematopotum
– Cocoon without dorsal projection (Figs. 95 A, K); frontoclypeus and thorax with
abundant to sparse tubercles ......................................................................................... 18
18(17) Gill branches relatively closed basally, subparallel (Figs. 96 A, K) ........................ 19
– Gill branches generally very open basally, and with tendency to converge distally
(Figs. 96 B, C, G, H) ........................................................................................................ 20
19(18) Secondary dorsal branch of gill thicker than secondary ventral branch; tertiary
dorsal branch bifurcated at long distance from base (Fig. 96 K); frontoclypeus and
thorax with moderate number of tubercles (Fig. 95 U) ........... Simulium bobpetersoni
– Secondary dorsal branch of gill with diameter similar to that of secondary ventral
branch; tertiary dorsal branch bifurcated at short distance from base (Fig. 96 A);
frontoclypeus and thorax with abundant tubercles (Fig. 95 P) ........ Simulium dugesi
20(18) Median tertiary gill branch bifurcated at short distance from base (Fig. 96 B);
frontoclypeus and thorax with sparse tubercles not elevated (Fig. 95 Q) ...............
........................................................................................................................ Simulium ochoai
– Median tertiary gill branch bifurcated at long distance from base (Figs. 96 C, G,
H); frontoclypeus and thorax with abundant to sparse tubercles, in latter case,
tubercles elevated as high as wide at base .................................................................. 21
21(20) Dorsal tertiary gill branch bifurcated at same distance as or farther than median
tertiary branch (Fig. 96 H) ...................................................................... Simulium downsi
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 427

– Dorsal tertiary gill branch bifurcated lower than median tertiary branch (Figs. 96 C, G)
.............................................................................................................................................. 22
22(21) Frontoclypeus and thorax with abundant tubercles (Fig. 95 S); gill length 2.2-2.5
mm; cocoon length 3.0-3.5 mm ........................................................... Simulium escomeli
– Frontoclypeus and thorax with moderate to few tubercles; gill length 1.5-2.1 mm;
cocoon length 2.4-2.7 mm ............................................................................................. 23
23(22) Proportionally short gill branches 1.5-1.6 mm (Fig. 95 C): ratio of gill/cocoon
length = 1.7 .................................................................................... Simulium mediovittatum
– Proportionally long gill branches 2.0-2.1 mm (Fig. 95 D): ratio of gill/cocoon
length = 1.2 .......................................................................................... Simulium bivittatum

Larvae
1 Postgenal cleft not developed (Figs. 99 M, 100 G) ........ Simulium tenuipes, Simulium penai
– Postgenal cleft developed to different depths ............................................................. 2
2(1) Postgenal cleft slightly developed (Figs. 97 L, N); ratio of postgenal bridge/
hypostomium = 1.2-1.7 .................................................................................................... 3
– Postgenal cleft more developed (Figs. 97 J, K, M); ratio of postgenal bridge/
hypostomium = 0.5-1.1 .................................................................................................... 6
3(2) Ratio of postgenal bridge/hypostomium = 1.7; body length of mature larvae 6.8
mm; cephalic apotome with basal half darkened, with light window medially (Fig.
97 G) ................................................................................................... Simulium mangabeirai
– Ratio of postgenal bridge/hypostomium = 1.2-1.4; body length of mature larvae
4.7-6.3 mm; cephalic apotome basally scarcely darkened and without light window
medially (Fig. 97 H) ........................................................................................................... 4
4(3) Ratio of postgenal bridge/hypostomium = 1.4; anal ring with about 78 rows of
hooks ....................................................................................................Simulium zempoalense
– Ratio of postgenal bridge/hypostomium = 1.2; anal ring with 66-70 rows of
hooks ..................................................................................................................................... 5
5(4) Ratio of proximal, medial and distal antennal segments = 1:1.3:1.5; anal ring with
about 66 rows of hooks .................................................................... Simulium dandrettai
– Ratio of proximal, medial and distal antennal segments = 1:1.5:2.0; anal ring with
about 70 rows of hooks ................................................................. Simulium longithallum
6(2) Hypostomium shorter than postgenal bridge (Fig 98 g); ratio of postgenal bridge/
hypostomium length = 1.3 .................................................................... Simulium blancasi
– Hypostomium as long as or longer than postgenal bridge; ratio of postgenal
bridge/hypostomium length = 0.5-1.1 .......................................................................... 7
7(6) Hypostomium longer than postgenal bridge (Figs. 97 J, K); ratio of postgenal
bridge/hypostomium 0.5-0.9 ........................................................................................... 8
– Hypostomium as long as or shorter than hypostomial bridge (Fig 97 M), ratio of
postgenal bridge/hypostomium 1.0-1.1 ...................................................................... 14
8(7) Ratio of postgenal bridge/hypostomium = 0.5-0.6; cephalic apotome without
ornamentation ................................................................. Simulium dugesi, Simulium ochoai
– Ratio of postgenal bridge/hypostomium = 0.7-0.9; cephalic apotome with basal half
darkened and enclosing elongated median light window (Figs. 97 B, C, E) ................ 9
428 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

9(8) Body length 5.5-6.0 mm; antenna about as long as stem of cephalic fan; ratio of
postgenal bridge/hypostomium = 0.7 ................................................ Simulium escomeli
– Body length 4.3-5.4 mm; antenna longer than stem of cephalic fan; ratio of
postgenal bridge/hypostomium = 0.8-0.9 .................................................................. 10
10(9) Anal ring with 89 rows of hooks; distal antennal segment as long as medial
segment (Fig. 97 O) ............................................................................ Simulium panamense
– Anal ring with 63-77 rows of hooks; distal antennal segment longer than medial
segment (Fig. 97 P) .......................................................................................................... 11
11(10) Cephalic fan with 52-5 rays; body length 5.0-5.5 mm .......... Simulium mediovittatum
– Cephalic fan with 31-48 rays; body length 4.1-5.0 mm ........................................... 12
12(11) Cephalic fan with 31-35 rays; anal ring with 74-91 rows ........ Simulium haematopotum
– Cephalic fan with 38-48 rays; anal ring with 63-74 rows ........................................ 13
13(12) Anal sclerite with hairs and scales; anal ring with 65-74 rows; rectal papillae with 1
diverticulum on each lobe ....................................................... Simulium gonzalezherrejoni
– Anal sclerite without hairs and scales (Fig. 97 Q); anal ring with 63 rows of hooks;
rectal papillae with 2-3 diverticula on each lobe (Fig. 97 S) ......... Simulium iracouboense
14(7) Anal ring with 100-115 rows with 14-20 hooks each; anal sclerite with abundant
hairs and scales (Fig. 97 R); cephalic apotome without ornamentation (Fig. 97 D)
.............................................................................................................................................. 15
– Anal ring with 60-90 rows with 11-15 hooks each; anal sclerite usually without
scales; cephalic apotome with or without ornamentation ....................................... 16
15(14) Cephalic fan with 30-32 rays; lateral sclerite of thoracic proleg with about 50
teeth; maximum length 5.5-6.0 mm ................................................... Simulium callidum
– Cephalic fan with 40-42 rays; lateral sclerite of thoracic proleg with about 37
teeth; maximum length 4.8-5.0 mm ......................................... Simulium pseudocallidum
16(14) Anal ring with 60-62 rows with 11 hooks each .......................... Simulium bobpetersoni
– Anal ring with 68-90 rows with 13-15 hooks each ................................................... 17
17(16) Body length 5.5-6.0 mm; anal sclerite with abundant scales ....... Simulium veracruzanum
– Body length 3.9-5.5 mm; anal sclerite without scales ............................................... 18
18(17) Cephalic apotome darkened subbasally with median longitudinal elongated and
1+1 transverse subbasal light windows (Figs. 97 A, F); body length 5.1-5.4 mm
................................................... Simulium downsi, Simulium samboni, Simulium trivittatum
– Cephalic apotome scarcely pigmented, only slightly darkened basally; body length
3.7-4.5 mm ............................................................................................ Simulium bivittatum

CLAVES DE ESPECIES DE SIMULIUM (PSILOPELMIA)


Hembras
1 Escudo uniformemente gris opaco ................................ grupo BLANCASI sp.......2
– Escudo con una banda mediana o 1+1 bandas longitudinales ..................................
................................................................................................ grupo ESCOMELI sp.......4
2(1) Tergitos abdominales III-V negro aterciopelado con o sin manchas claras
centrales anteriores transversas (Fig. 98 A); cerdas del escudo esparcidas, no
conspicuas con poco aumento ............................................................. Simulium blancasi
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 429

– Tergitos abdominales III-V castaño o piceo, sin manchas claras centrales


anteriores transversas (Fig. 99 A); cerdas del escudo muy abundantes, conspicuas
con bajo aumento ............................................................................................................... 3
3(2) Tergitos abdominales III-V totalmente castaño oscuro aterciopelados (Fig. 99 A);
calcipala relativamente grande, redondeada apicalmente (Fig. 99 D) .......................
..................................................................................................................... Simulium tenuipes
– Tergitos abdominlaes III-V oscuros anteriormente, notoriamente más claros
posteriormente, ocupando la mitad posterior en el V; calcipala muy reducida,
subtriangular (Fig. 100 A) .......................................................................... Simulium penai
4(1) Escudo grisáceo con una banda mediana longitudinal castaño rojizo (Fig. 91 A) o
unicolor; cibario con dientes pequeños bien definidos en porción basal (Fig. 91 J)
................................................................................................................................................ 5
– Escudo con 1+1 bandas longitudinales submedianas generalmente cruzando
desde el borde anterior al posterior (Figs. 91 B-E); cibario con dientes, espículas, o
liso en la porción basal (Figs. 91 L-Q) .......................................................................... 7
5(4) Metanoto negruzco; longitud ala 1,9-2,4 mm ......................... Simulium mediovittatum
– Metanoto castaño grisáceo; longitud ala 1,8-2,2 mm ................................................. 6
6(5) Longitud alar 2,0-2,2 mm; escutelo castaño grisáceo; tergito abdominal II grisáceo
(Fig. 91 G) .................................................................................................... Simulium dugesi
– Longitud ala 1,8 mm; escutelo amarillo a castaño amarillento; tergito abdominal II
amarillo grisáceo ......................................................................................... Simulium ochoai
7(4) Tergitos abdominales sin hileras de manchas oscuras subcuadradas; porción distal
del paraprocto alargado, afindandose suavemente (Figs. 92 F, G); uñas con diente
subbasal bien desarrollado (Figs. 91 V, W) ................................................................... 8
– Tergitos abdominales con 1 hilera mediana de manchas oscuras subcuadradas,
1+1 submedianas y a veces 1+1 sublateral por lo menos en los segmentos III-VI
(Figs. 91 H, I); porción distal del paraprocto relativamente corto (Figs. 92 B, E, J)
o alargado, afinandose abruptamente y prolongado en un angosto apéndice (Figs.
92 H-N); uñas sin diente subbasal (Figs. 91 T-U) ..................................................... 10
8(7) Cibario con una pequeña depresión mediana y abundantes pequeños dientes
submedianamente (Fig. 91 N); paraprocto distalmente alargado, tan largo como el
ancho basal y acuminado distalmente (Fig. 92 F) ........................ Simulium panamense
– Cibario con un grueso reborde, con 1+1 suaves protuberancias a veces cubiertas
por finas espículas (Fig. 91 O); prolongación distal del paraprocto ausente y con
ápice romo (Fig. 92 G) ...................................................................................................... 9
9(8) Sector basal de R con pelos ................................................................. Simulium callidum
– Sector basal de R glabro ............................................................. Simulium pseudocallidum
10(7) Tergitos abdominales con 1 hilera mediana y 1+1 submediana de manchas
oscuras subcuadradas (Fig. 91 H); prolongación distal del paraprocto más corto
que la mitad del ancho en la base (Figs. 92 B, E, J) ................................................. 11
– Tergitos abdominales con 1 hilera mediana y 1+1 sublateral de manchas oscuras
subcuadradas (Fig. 91 I); prolongación distal del paraprocto con un largo
aproximado al ancho en la base (Figs. 92 C, D) o muy alargado (Figs. 92 H, I, K-N)
.............................................................................................................................................. 13
430 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

11(10) Escutelo castaño grisáceo; vesícula sensorial del palpo maxilar más corta que el
palpómero (Fig. 91 S); paraprocto con ápice romo (Fig. 92 E) .................................
........................................................................................................... Simulium haematopotum
– Escutelo castaño amarillento; vesícula sensorial del palpo maxilar generalmente
más larga que la mitad del largo del palpómero; paraprocto suavemente aguzado
(Figs. 92 B-J) ..................................................................................................................... 12
12(11) Longitud ala 2,1-2,7 mm ................................................................... Simulium bivittatum
– Longitud ala 2,4-3,0 mm ................................................................... Simulium dandrettai
13(10) Paraprocto romo apicalmente y prolongación distal aproximadamente tan larga
como la mitad del ancho en la base (Fig. 92 D); escudo amarillento-anaranjado
con 1+1 banda submediana plateada muy fina (Fig. 91 B) ..................................... 14
– Paraprocto aguzado apicalmente y prolongación distal tan larga como el ancho en
la base (Figs. 92 H, I, K-N); escudo gris amarillento con banda mediana y
sublaterales castaño a negro (Figs. 91 E, F) (excepto Simulium downsi) ................ 15
14(13) Longitud ala 1,5-1,6 mm; vesícula sensorial del palpo maxilar aproximadamente
tan larga como la mitad de la longitud del palpómero (Fig. 91 R) ...........................
.............................................................................................................. Simulium iracouboense
– Longitud ala 1,8-2,1 mm; vesícula sensorial del palpo maxilar más corta que la
mitad de la longitud del palpómero .................................................... Simulium samboni
15(13) Sector basal de R con pelos; escudo amarillento con bandas mediana y
sublaterales castaño amarillento claras ................................................. Simulium downsi
– Sector basal de R sin pelos; escudo gris amarillento con bandas medianas y
sublaterales castaño oscuro a negro ............................................................................. 16
16(15) Escudo generalmente amarillento con bandas medianas y sublaterales castaño
claro a amarillo grisáceo o bien de castaño oscuro a negruzco sobre bandas
mediana y sublaterales (Fig. 91 E) ....................................................... Simulium escomeli
– Escudo amarillo grisáceo con bandas medianas y sublaterales castaño oscuro a
negruzco (Fig. 91 F) ........................................................................................................ 17
17(16) Prolongación distal del paraprocto más corta que la mitad del ancho en la base
(similar a Simulium dandretta) (Fig. 91 J) ........................................ Simulium zempoalense
– Prolongación distal del paraprocto aproximadamente tan larga como el ancho en
la base (Figs. 92 K-N) ..................................................................................................... 18
18(17) Banda mediana del escudo negra más ancha que las bandas plateadas
submedianas, y bandas negras sublaterales no alcanzan el borde anterior (Fig. 91 F)
.............................................................................................................. Simulium mangabeirai
– Banda mediana negra del escudo aproximadamente del mismo ancho que las
bandas plateadas submedianas, y las bandas negras sublaterales alcanzan o llegan
muy cerca del borde anterior ....................................... Simulium longithallum, Simulium
gonzalezherrejoni, Simulium trivittatum, Simulium bobpetersoni, Simulium veracruzanum

Machos
1 Escudo uniformemente gris opaco sin cuñas plateadas ..............................................
................................................................................................ grupo BLANCASI sp.......2
– Escudo con 1+1 cuñas plateadas (Figs. 93 A-G) ........ grupo ESCOMELI sp.......4
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 431

2(1) Tergitos abdominales III-V negro aterciopelado, con borde anterior en el centro
con una mancha clara (Fig. 98 P); setas de escudo esparcidas y no conspicuas con
bajo aumento; genitalia con placa ventral contraida lateralmente cerca de la base
(Fig. 98 W) ................................................................................................ Simulium blancasi
– Tergitos abdominales III-V oscuros, a veces con áreas posteriores claras pero
nunca sobre el borde anterior; setas del escudo muy abundantes, conspícuas con
bajo aumento; genitalia con placa ventral no contraida lateralmente cerca de la
base (Fig. 99 J) .................................................................................................................... 3
3(2) Basitarso posterior 4 veces más largo que ancho; calcipala muy reducida
subtriangular (Fig. 100 A) .......................................................................... Simulium penai
– Basitarso posterior menos de 4 veces largo que ancho; calcipala no tan reducida y
redondeada apicalmente (como en la hembra, Fig. 99 D) ............. Simulium tenuipes
4(1) Escudo y abdomen amarillento pudiendo ser castaño grisáceo, pero por lo menos
con alguna porción amarillo-anaranjado; escudo con cuñas plateadas solo en área
anterior (Fig. 93 B) ............................................................................................................. 5
– Escudo y abdomen gris a negruzco; escudo con uñas plateadas de variable
extensión (Figs. 93 A, C-G) ........................................................................................... 10
5(4) Cuerpo de la placa ventral subovoidal (Fig. 94 F) ...................................................... 6
– Cuerpo de la placa ventral de forma variable, no subovoidal .................................. 7
6(5) Sector basal de R con pelos ................................................................. Simulium callidum
– Sector basal de R sin pelos ......................................................... Simulium pseudocallidum
7(5) Placa ventral con fuerte contracción lateral sobre la mitad del cuerpo, continuada
por una porción angosta medio distal (Fig. 94 E) ....................... Simulium panamense
– Placa ventral variable pero sin fuerte contracción lateral en el cuerpo (Figs. 94 C,
H) ........................................................................................................................................... 8
8(7) Gonostilo subtrapezoidal con fuerte expansión sobre el lado externo subterminal
(Fig. 93 N); placa ventral del cuerpo reducida,subtriangular (Fig. 94 H) y más
angosta que la posición de las ramas .................................................... Simulium downsi
– Gonostilo sin expansión lateral o subterminal (Figs. 93 K, L); cuerpo de la placa
ventral más desarrollado, no subtrioangular y más ancho que la posición de las
ramas (Fig. 94 C) ................................................................................................................ 9
9(8) Gonostilo subtriangular (Fig. 93 J); relación del largo/ancho del basitarso
posterior = 7,8 ......................................................................................... Simulium samboni
– Gonostilo subtrapezoidal (Fig. 93 K); relación del largo/ancho del basitarso
posterior = 6,6 .................................................................................. Simulium iracouboense
10(4) Escudo con 1+1 banda corta plateada no llegando al área posterior grisácea (Fig.
93 A) ................................................................................................................................... 11
– Escudo con 1+1 banda larga plateada arribando al área posterior grisácea (Fig.
93 G) ................................................................................................................................... 19
11(10) Escudo con 1+1 áreas azul plateado triangulares u ovales, contiguas
anteriormente con áreas laterales del mismo color (Fig. 93 A) ............................. 12
– Escudo con 1+1 áreas azul plateado no contiguas con áreas laterales del mismo
color (Fig. 93 D) ............................................................................................................... 18
432 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

12(11) Cuñas plateadas submedianas fuertemente afinadas posteriormente (Fig. 93 A),


sobrepasando o no la mitad del escudo; gonostilo aproximadamente tan largo
como ancho con el lado externo recto (Fig. 93 I); placa ventral subtriangular con
ápice redondeado (Fig. 94 A) ........................................................................................ 13
– Cuñas plateadas submedianas generalmente sobrepasando la mitad del escudo;
gonostilo expandido sobre el lado externo distal subterminal (Figs. 93 P, Q, S);
placa ventral variable desde subtriangular, con o sin contracciones laterales, a
subovoidal .......................................................................................................................... 15
13(12) Cuñas submedianas del escudo no llegando al área grisácea posterior (Fig. 93 A)
.............................................................................................................................................. 14
– Cuñas submedianas del escudo bien afinadas posteriormente y llegando al área
grisácea posterior .......................................................................... Simulium mediovittatum
14(13) Longitud del ala 1,9 mm ........................................................................... Simulium dugesi
– Longitud del ala 1,6 mm ........................................................................... Simulium ochoai
15(12) Gonostilo menos de 2 veces más larga que ancho en la base, expandido distalmente,
más ancho que en la base (Fig. 93 O); placa ventral sin contracciones laterales, con
bordes redondeados (Fig. 94 I); relación largo/ancho en la base = 1,2 .....................
................................................................................................................. Simulium dandrettai
– Gonostilo generalmente más de 2 veces largo que ancho en la base, no expandido
distalmente; placa ventral generalmente sin bordes redondeados; relación largo/
ancho en la base sobre 1,2 ............................................................................................. 16
16(15) Gonostilo subrectangular con los lados subparalelos (Fig. 93 I); placa ventral
relativamente corta, sin contracciones laterales (Fig. 93 B) ....... Simulium bivittatum
– Gonostilo subtrapezoidal, distalmente con fuerte curvatura y ápice suavemente
aguzado (Fig. 93 N); placa ventral relativamente corta y ancha (Fig. 94 J) hasta más
alargada con contracción lateral (Fig. 94 G) .............................................................. 17
17(16) Placa ventral relativamente corta con base prolongada basalmente (Fig. 94 J);
relación largo/ancho = 2,3; longitud ala 1,9-2,1 mm ............... Simulium bobpetersoni
– Placa ventral relativamente larga, con base no prolongada basalmente y
lateralmente con suave contracción (Fig. 94 G); relación largo/ancho = 1,3-1,4;
longitud ala 2,0-2,3 mm ......................................................................... Simulium escomeli
18(11) Placa ventral subtriangular sin contracción lateral (Fig. 94 D); gonostilo bien
alargado distalmente con un largo y curvo espolón (Fig. 93 L); longitud del ala 1,8-
2,1 mm ............................................................................................. Simulium haematopotum
– Placa ventral con fuerte contracción lateral, con porciónn posterior a la
contracción angosta, aproximadamente 1/3 del ancho del cuerpo (Fig. 94 K);
gonostilo no muy alargado distalmente y espolón con moderado tamaño (Fig. 93
P); longitud ala 2,6-3,0 mm ........................................................... Simulium veracruzanun
19(10) Gonostilo aproximadamente 2/3 el ancho con respecto al largo (Fig. 93 R); placa
ventral sin contracción lateral; borde distal del cuerpo subcircular ..........................
...............................................................................................................Simulium zempoalense
– Gonostilo aproximadamente 1/2 el ancho con respecto al largo (Fig. 93 Q); placa
ventral con o sin contracción y con diferente forma al dilema opuesto (Figs. 94 L-
N) ......................................................................................................................................... 20
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 433

20(19) Relación largo/ancho del basitarso posterior = 3,8; longitud ala 2,5-2,8 mm;
escudo con 1+1 cuñas claras no llegando al borde posterior (Fig. 93 F) ................
.............................................................................................................. Simulium mangabeirai
– Relación largo/ancho del basitarso posterior >5,0; escudo con 1+1 cuña clara
ancha llegando al borde posterior (Fig. 93 G) ........................................................... 21
21(20) Segmentos abdominales I-II amarillentos; longitud ala 2,5 mm ................................
................................................................................................................. Simulium trivittatum
– Segmentos abdominales I-II no amarillentos; longitud del ala 2,0-2,5 mm ........ 22
22(21) Escudo con 1+1 bandas laterales negruzcas no llegando a los lóbulos del
postpronoto (Fig. 93 G); longitud ala 2,5 mm .................... Simulium gonzalezherrejoni
– Escudo con 1+1 bandas laterales negruzcas llegando a los lóbulos del
postpronoto; longitud alas 2,0 mm .............................................. Simulium longithallum

Pupas
1 Branquia con 20 ramas (Fig. 96 D) ................................................. Simulium panamense
– Branquia con 4-8 ramas .................................................................................................... 2
2(1) Branquia con 4 ramas (Fig. 96 M) .................................................................................. 3
– Branquia con 6-8 ramas .................................................................................................... 6
3(2) Capullo bien desarrollado cubriendo la mayor parte del tórax y cabeza (Fig. 95 M);
borde ventral anterior más adelantado que el dorsal ............... Simulium mangabeirai
– Capullo escasamente desarrollado no cubriendo la mayor parte del tórax y cabeza;
borde dorsal anterior más adelantado que el ventral (Fig. 98 X) ..............................
................................................................................................ BLANCASI sp. grupo......4
4(3) Branquia con un corto tronco seguido por 2 ramas distinguibles con 2+2
filamentos (Fig. 98 Y) ............................................................................. Simulium blancasi
– Branquias con 4 filamentos aparentemente emergidos de un tallo muy corto
común (Figs. 99 L, 100 C) ................................................................................................ 5
5(4) Frontoclipeo y porción expuesta del tórax con abundantes tubérculos; superficie
del capullo rugosa y de color castaño cuando se lo examina en líquido .................
..................................................................................................................... Simulium tenuipes
– Frontoclipeo y porción expuesta del tórax con muy esparcidos tubérculos (Figs.
100 D, E); capullo liso y de color castaño con brillo plomizo como un pergamino
cuando se lo examina en líquido .............................................................. Simulium penai
6(2) Branquias con 6 ramas ...................................................................................................... 7
– Branquias con 8 ramas .................................................................................................... 10
7(6) Branquias con superficie rugosa; ápice romo con pigmentación castaño a
negruzca (Figs. 96 I, J); branquias más cortas que el largo del capullo (Figs. 95 I, J)
................................................................................................................................................ 8
– Branquias con superficie lisa; generalmente con ápice aguzado con pigmentación
amarillenta; branquias tan largas o más largas que el largo del capullo (Fig. 95 N)
................................................................................................................................................ 9
8(7) Ramas secundarias externa e interna de la branquia más gruesas que otras ramas
de la branquia (Fig. 96 I); tubérculos del frontoclípeo y tórax lisos; capullo sin
elevaciones longitudinales (Fig. 95 I) .............................................. Simulium dandrettai
434 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

– Rama dorsal externa únicamente más gruesa que otras ramas de la branquia (Fig.
96 J); tubérculos del frontoclípeo y tórax rugosos; capullo con elevaciones
longitudinales (Fig. 95 J) .......................................... Simulium sp. próximo a dandrettai
9(7) Ramas primarias y secundarias de las branquias emergiendo muy próximas a la base
(Fig. 96 O); tegumento dorsal del tórax con escasos tubérculos y elevaciones
circulares chatas rugosas simulando tubérculos (Fig. 95 Q) ...... Simulium zempoalense
– Ramas primarias y secundarias de las branquias no emergiendo próximas a la base
(Fig. 96 P); tegumento dorsal del tórax con tegumento liso y escasos tubérculos
................................................................................................................. Simulium trivittatum
10(6) Branquias con tallo basal corto, con ramas primarias y secundarias emergiendo
muy próximas a la base, con la rama primaria dorsal más gruesa que las otras (Figs.
96 E, F) .............................................................................................................................. 11
– Branquias con ramas primarias y secundarias emergiendo no tan cerca de la base
de un tallo basal más desarrollado y con las ramas primarias de aproximadamente
del mismo diámetro ......................................................................................................... 12
11(10) Terciaria rama dorsal de la branquia notoriamente más gruesa y 1/3 más larga que
las otras branquias (Fig. 96 F) ................................................... Simulium pseudocallidum
– Terciaria rama dorsal de la branquia solo un poco más gruesa y con similar
longitud que las otras ramas (Fig. 96 E) ........................................... Simulium callidum
12(10) Ramas dorsal secundaria y terciaria de la branquia más gruesas que las otras ramas
(Figs. 96 N, O) .................................................................................................................. 13
– Ramas dorsal secundaria y terciaria de la branquia del mismo diámetro que las
otras ramas ......................................................................................................................... 14
13(12) Ramas dorsal secundaria y terciaria emergiendo lejos de la base (Fig. 96 N) .........
.......................................................................................................Simulium gonzalezherrejoni
– Ramas dorsal secundaria y terciaria emergiendo próximas a la base (Fig. 96 O)
.............................................................................................................. Simulium longithallum
14(12) Longitud de las branquias tan largas o más largas que el largo del capullo (Fig. 95 L)
............................................................................................................ Simulium veracruzanum
– Longitud de las branquias más cortas que el largo del capullo (Figs. 95 A-G) ......... 15
15(14) Tejido del capullo esparcido con trama muy abierta, sin borde anterior (Fig. 95 F)
................................................................................................................ Simulium iracoubense
– Tejido del capullo compacto con trama cerrada, con borde anterior (Figs. 95 A-E,
G, H, K) ............................................................................................................................. 16
16(15) Tricomas cefálicos y torácicos más largos que la mitad del ancho basal del
frontoclípeo (Fig. 95 R); borde anterior del capullo reforzado y con una
prolongación dorsal anterior (Fig. 95 E) ............................................ Simulium samboni
– Tricomas cefálicos y torácicos no más largos que la mitad del ancho basal del
frontoclípeo (Figs. 95 Q, T, U) borde anterior del capulloo no reforzado y sin una
prolongación dorsal anterior .......................................................................................... 17
17(16) Capullo con pequeña prolongación dorsal y ventral (Fig. 95 G); frontoclípeo y
tórax con escasos tubérculos (Fig. 95 T) .................................. Simulium haematopotum
– Capullo sin prolongación dorsal (Figs. 95 A, K); frontoclípeo y tórax desde
abundantes a escasos tubérculos ................................................................................... 18
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 435

18(17) Ramas branquiales relativamente apretadas basalmente, con disposición


subparalela (Figs. 96 A, K) ............................................................................................. 19
– Ramas branquiales generalmente muy abiertas basalmente y con tendencia a
convergir distalmente (Figs. 96 B, C, G, H) ................................................................ 20
19(18) Rama secundaria dorsal de la branquia más gruesa que la rama secundaria ventral;
rama terciaria dorsal bifurcada a larga distancia de la base (Fig. 96 K); frontoclípeo
y tórax con moderado número de tubérculos (Fig. 95 U) ....... Simulium bobpetersoni
– Rama secundaria dorsal de la branquia con similar diámetro que la rama secundaria
ventral; rama terciaria dorsal bifurcada a corta distancia de la base (Fig. 96 A);
frontoclípeo y tórax con abundantes tubérculos (Fig. 95 P) ............... Simulium dugesi
20(18) Rama terciaria mediana de la branquia bifurcada desde corta distancia de la base
(Fig. 96 B); frontoclípeo y tórax con escasos tubérculos y manchas cuticulares
circulares rugosas no elevadas (Fig. 95 Q) ........................................... Simulium ochoai
– Rama terciaria mediana de la branquia bifurcada a larga distancia de la base (Figs. 96
C, G, H); frontoclípeo y tórax con tubérculos desde abaundantes a escasos, en el
último caso los tubérculos son elevados tan largos como anchos en la base ........ 21
21(20) Rama terciaria dorsal de la branquia bifurcada a la misma distancia o más lejos que
la rama terciaria mediana (Fig. 96 H) ................................................... Simulium downsi
– Rama terciaria dorsal de la branquia bifurcada por debajo de la rama terciaria
mediana (Figs. 96 C, G) .................................................................................................. 22
22(21) Frontoclípeo y tórax con abundantes tubérculos (Fig. 56 S); longitud de la branquia
2,2-2,5 mm; longitud capullo 3,0-3,5 mm ........................................... Simulium escomeli
– Frontoclípeo y tórax desde moderado número a escasos tubérculos; longitud de la
branquia 1,5-2,1 mm; longitud capullo 2,4-2,7 mm ................................................. 23
23(22) Branquias proporcionalmente cortas 1,5-1,6 mm; relación largo branquia/capullo
= 1,7 ................................................................................................. Simulium mediovittatum
– Branquias proporcionalmente largas 2,0-2,1 mm; relación largo branquia/capullo
= 1,2 ....................................................................................................... Simulium bivittatum

Larvas
1 Hendidura postgenal no desarrollada (Figs. 99 M, 100 G) .........................................
.......................................................................................... Simulium tenuipes, Simulium penai
– Hendidura postgenal desarrollada con diferente profundidad ................................ 2
2(1) Hendidura postgenal escasamente desarrollada (Figs. 97 L, N); relación del largo
puente hipostomial/hipostomio = 1,2-1,7 ................................................................... 3
– Hendidura postgenal más desarrollada (Figs. 97 J, K, M); relación del largo puente
hipostomial/ hipostomio = 0,5-1,1 ................................................................................ 6
3(2) Relación puente postgenal/hipostomio = 1,7; longitud del cuerpo de la larva
madura 6,8 mm; apotoma cefálico con la mitad basal oscurecida, con una ventana
mediana (Fig. 97 G) ......................................................................... Simulium mangabeirai
– Relación puente postgenal/hipostomio = 1,2-1,4; longitud del cuerpo de la larva
madura 4,7-6,3 mm; apotoma cefálico escasamente oscurecido y sin una ventana
mediana (Fig. 97 H) ........................................................................................................... 4
436 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

4(3) Relación puente hipostomial/hipostomio = 1,4; anillo anal con alrededor de 78


hileras de ganchos .............................................................................Simulium zempoalense
– Relación puente postgenal/hipostomio = 1,2; anillo anal con 66-70 hileras de
ganchos ................................................................................................................................. 5
5(4) Relación de los artejos antenales proximal, mediano y distal = 1:1,3:1,5; anillo anal
con alrededor de 66 hileras de ganchos ......................................... Simulium dandrettai
– Relación de los artejos antenales proximal, mediano y distal = 1:1,5:2,0; anillo anal
con alrededor de 70 hileras de ganchos ...................................... Simulium longithallum
6(2) Hipostomio más corto que el puente postgenal (Fig. 98 g); relación largo puente
postgenal/hipostomio = 1,3 ................................................................. Simulium blancasi
– Hipostomio tan largo o más largo que el puente postgenal; relación largo puente
postgenal/hipostomio = 0,5-1,1 ..................................................................................... 7
7(6) Hipostomio más largo que el puente postgenal (Figs. 97 J, K); relación largo puente
postgenal/hipostomio = 0,5-0,9 ...................................................................................... 8
– Hipostomio con similar largo o más corto que el puente postgenal (Fig. 97 M) );
relación largo puente postgenal/hipostomio = 1,0-1,1 ........................................... 14
8(7) Relación largo puente postgenal/hipostomio = 0,5-0,6; apotoma cefálico sin
ornamentación ................................................................ Simulium dugesi, Simulium ochoai
– Relación largo puente postgenal/hipostomio = 0,7-0,9; apotoma cefálico oscurecido
en mitad basal, encerrando una ventana clara mediana (Figs. 97 B, C, E) ................... 9
9(8) Longitud del cuerpo 5,5-6,0 mm, antena con similar largo que el tallo del abanico
cefálico; relación largo puente postgenal/hipostomio = 0,7 ......... Simulium escomeli
– Longitud del cuerpo 4,3-5,4 mm; antena más larga que el tallo del abanico
cefálico; relación largo puente postgenal/hipostomio = 0,8-0,9 ........................... 10
10(9) Anillo anal con 89 hileras de ganchos; artejo distal de la antena de similar longitud
que el mediano (Fig. 97 O) ............................................................... Simulium panamense
– Anillo anal con 63-77 hileras de ganchos; artejo distal de la antena más largo que
el mediano (Fig. 97 P) ..................................................................................................... 11
11(10) Abanico cefálico con 52-56 rayos; longitud del cuerpo 5,0-5,5 mm ........................
........................................................................................................... Simulium mediovattatum
– Abanico cefálico con 31-48 rayos; longitud del curpo 4,1-5.0 mm ...................... 12
12(11) Abanico cefálico con 31-35 rayos; anillo anal con 74-91 rayo ...................................
........................................................................................................... Simulium haematopotum
– Abanico cefálico con 38-48 rayos; anillo anal con 63-74 rayos ............................. 13
13(12) Esclerito anal con pelos y escamas; anillo anal con 65-74 hileras; papilas rectales
con 1 divertículo en cada lóbulo ............................................ Simulium gonzalezherrejoni
– Esclerito anal sin pelos ni escamas (Fig. 97 Q); anillo anal con 63 hileras de
ganchos;papilas rectales con 2-3 divertículos en cada lóbulo (Fig. 97 S) .................
.............................................................................................................. Simulium iracouboense
14(7) Anillo anal con 100-115 hileras con 14-20 ganchos cada una; esclerito anal con
abundantes pelos y escamas (Fig. 97 R); apotoma cefálico sin ornamentación (Fig.
97 D) ................................................................................................................................... 15
– Anillo anal con 60-90 hileras con 11-15 ganchos cada una; esclerito anal
generalmente sin escamas; apotoma cefálico con o sin ornamentación .............. 16
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 437

15(14) Apotoma cefálico con 30-32 rayos; esclerito lateral de la propata torácica con
alrededor de 50 dientes; longitud máxima 5,5-6,0 mm .................. Simulium callidum
– Apotoma cefálico con 40-42 rayos; esclerito lateral de propata torácica con
alrededor de 37 dientes; longituud máxima 4,8-5,0 mm ..... Simulium pseudocallidum
16(14) Anillo anal con 60-62 hileras con 11 ganchos cada una ........... Simulium bobpetersoni
– Anillo anal con 68-90 hileras con 13-15 ganchos cada una .................................... 17
17(16) Longitud del cuerpo 5,5-6,0 mm; esclerito anal con abundantes escamas ..............
............................................................................................................Simulium veracruzanum
– Longitud del cuerpo 3,9-5,5 mm; esclerito anal sin escamas ................................. 18
18(17) Apotoma cefálico oscurecido subbasalmente con una ventana clara mediana
longitudinal alargada y 1+1 transversa subbasal (Figs. 97 A, F); longitud del
cuerpo 5,1-5,4 mm ................ Simulium downsi, Simulium samboni, Simulium trivittatum
– Apotoma cefálico escasamente pigmentado, solamente un poco más oscurecido
basalmente; longitud del cuerpo 3,7-4,5 mm ................................ Simulium bivittatum

ESCOMELI species group

Simulium (Psilopelmia) dugesi VARGAS, MARTÍNEZ PALACIOS & DÍAZ NÁJERA


(Figs. 91 A, G, J; 92 A; 93 A, H; 94 A; 95 A, P; 96 A; 97 J.)
Simulium (Lanea) dugesi VARGAS et al., 1946: 107; DALMAT, 1955: 59.
Simulium (Psilopelmia) dugesi VARGAS & DÍAZ NÁJERA, 1957b: 182; COSCARÓN, 1987: 31; COSCARÓN et
al., 1996: 48-50; CROSSKEY & HOWARD, 1997: 58; 2004: 53.
Female: Wing length 2.0-2.2 mm; scutum grayish with reddish brown median
longitudinal stripe diffuse at sides (Fig. 91 A); with some illumination, 1+1 submedian
anterior light subtriangular spots reaching anterior rim; scutum and metanotum dark gray;
pleurae with pruinosity gray; legs yellowish brown with apex of femora and tarsomeres
blackish; abdomen grayish with blackish subquadrate median spots on II-VI and 1+1
sublateral spots wider on III-VI tergites and sometimes on VII (Fig. 91 G); cibarium with
several rows of minute spines except on median depressed area (Fig. 91 J); fronto-ocular
triangle slightly longer than wide; basal sector of R glabrous; sensory vesicle of maxillary
palp about half of palpomere length; claw without subbasal tooth; eighth sternite with about
20 setae on each side; paraproct short, truncate slightly acuminate dorsally (Fig. 92 A).
Male: Wing length 1.9 mm; scutum black with 1+1 subtriangular silvery spots
extending posteriorly as thin stripes surpassing half of scutum length (Fig. 93 A);
abdomen blackish with 1+1 silvery spots on tergites II, VI, VII; gonostylus slightly
acuminate on distal inner portion (Fig. 93 H); ventral plate short with gently curved
border (Fig. 94 A).
Pupa: Cocoon reinforced anterior border, with small anterior projection (Fig. 95 A),
threads evident; length basally 2.9-3.4 mm, at dorsum 2.5-2.8 mm, gills 2.0-2.6 mm;
frontoclypeus and thorax with numerous tubercles (Fig. 95 P); gill with 8 branches
bifurcated not close to base (Fig. 96 A).
Larva: Maximum length 4.6-5.2 mm; cephalic apotome with homogeneous
coloration; ratio of proximal, medial and distal antennal segments = 1:1.3:1.7; cephalic
438 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

A C F D

B G H I E

J L
K

N
M
O

Q
P

S
R
U T V W
Figure 91. Female. A-F: Scutum; A: S. (Psilopelmia) dugesi; B: S. (P.) samboni; C: S. (P.) haematopotum;
D: S. (P.) callidum; E: S. (P.) escomeli; F: S. (P.) mangabeirai. G-I: Abdomen: G: S. (P.) dugesi; H: S. (P.)
iracouboense; I: S. (P.) escomeli. J-Q: Basal portion of cibarium; J: S. (P.) dugesi; K: S. (P.) bivittatum; L: S.
(P.) samboni; M: S. (P.) ircouboense; N: S. (P.) panamense; O: S. (P.) pseudocallidum; P: S. (P.) trivittatum; Q:
S. (P.) dandrettai. R-S: Sensory organ of palp; R: S. (P.) iracouboense; S: S. (P.) haematopotum. T-W:
Claws; T: S. (P.) bivittatum; U: S. (P.) iracouboense; V: S. (P.) panamense; W: S. (P.) callidum.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 439

fan with about 40 rays; mandible with latero-mandibular process; postgenal cleft dome
shaped; postgenal bridge about half of hypostomium length (Fig. 97 J); rectal papillae
with only 3 single lobes; anal sclerite without hair or scales; anal ring with 70 rows of
15-16 hooks each.
Distribution: Mexico: Nueva León, San Luis Potosí, Tamaulipas.
Bionomics: Unknown.
Discussion: Simulium dugesi is very close to S. mediovittatum and S. ochoai; differences
are explained under those species.

Simulium (Psilopelmia) ochoai VARGAS, MARTÍNEZ PALACIOS & DÍAZ NÁJERA


(Figs. 95 B, Q; 96 B; 97 I.)
Simulium (Lanea) ochoai VARGAS et al., 1946: 131-133, 169.
Simulium (Psilopelmia) ochoai: VARGAS & DÍAZ NÁJERA, 1957b: 182; COSCARÓN, 1987: 31; COSCARÓN et
al., 1996: 50-51; CROSSKEY & HOWARD, 1997: 59; 2004: 53.
Female: Wing length 2.1 mm; scutum, pleurae, legs, and metanotum similar to
those of S. dugesi; scutellum light yellowish brown; abdomen as in S. dugesi except
segment II yellowish; cibarium, sensory vesicle of maxillary palp, basal sector of R and
paraproct as in S. dugesi.
Male: Wing length 1.6 mm; coloration of scutum and abdomen as in S. dugesi, but
scutum submedian silvery stripes slightly shorter, not surpassing half of scutum length;
gonostylus elongated on apical third, very similar to S. haematopotum.
Pupa: Cocoon with anterior border reinforced, not projected anteriorly (Fig. 95 B),
weave with evident threads; length along base 2.8-3.2 mm, along dorsum 2.7-2.9 mm, gills
1.5-2.0 mm; frontoclypeus and thorax with scarce tubercles (Fig. 95 Q); trichomes short,
with 1-2 branches; gill with 8 branches arranged very open with primary branches very
short (Fig. 96 B).
Larva: Maximum length 4.5-4.8 mm; cephalic apotome light whitish yellow
scarcely darkened basally with subtriangular or bell shape, but with light median
window; antenna longer than cephalic fan stem; ratio of proximal, medial and distal
segments = 1:1.4:1.8; cephalic fan with about 42 rays; mandible with latero-mandibular
process (Fig. 97 I); postgenal cleft as in S. dugesi; postgenal bridge longer than half of
hypostomium length; lateral sclerite of proleg with 14-15 teeth arranged in 10-11
groups; anal sclerite with scarce hair and scales; anal ring with 64-70 rows of 14-15
hooks each; rectal papillae without diverticula.
Distribution: Mexico: Veracruz.
Bionomics: Unknown.
Discussion: The adults are very similar to those of S. dugesi and S. mediovittatum, but
the pupae are distinguishable. The first species has more numerous tubercles on the
frontoclypeus and thorax, and both species have the pupal gill branching closer to the
base, forming a bunch in S. dugesi.
440 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

B C D

E J F

H
L
N

I Q

K O

Figure 92. Female. A-N: Cercus and paraproct; A: S. (Psilopelmia) dugesi; B: S. (P.) bivittatum; C: S.
(P.) samboni; D: S. (P.) iracouboense; E: S. (P.) haematopotum; F: S. (P.) panamense; G: S. (P.) callidum; H: S.
(P.) escomeli; I: S. (P.) downsi; J: S. (P.) dandrettai; K: S. (P.) bobpetersoni; L: S. (P.) veracruzanum; M: S. (P.)
gonzalezherrejoni; N: S. (P.) trivittatum. O: Eighth sternite and gonapophyses of S. (P.) mediovittatum. P-
Q: Fronto-ocular triangle; P: S. (P.) iracouboense; Q: S. (P.) samboni.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 441

Simulium (Psilopelmia) mediovittatum KNAB (Figs. 92 O; 95 C; 97 K.)


Simulium mediovittatum KNAB, 1915 a: 77; 1915b: 149; VARGAS & DÍAZ NÁJERA, 1954: 69.
Simulium (Lanea) mediovittatum: VARGAS et al., 1946: 107.
Simulium (Psilopelmia) mediovittatum: VARGAS & DÍAZ NÁJERA, 1957b: 154; PETERSON, 1993: 329;
COSCARÓN, 1994: 31; COSCARÓN et al., 1996: 51-54; MOULTON, 1998: 63; CROSSKEY & HOWARD,
1997: 59; 2004: 53; ADLER et al., 2004: 352.
Female: Wing length 2.2 mm; frons and clypeus dark brown with grayish pruinosity;
scutum light grayish brown with variable tones from orange to brown with median
reddish-brown to black stripe, covered by decumbent yellowish pile; scutellum coloration
as for scutum; metanotum brownish black; pleurae yellowish to brownish black; legs
blackish with tibia apex and tarsomeres whitish; abdomen yellow-brown grayish to dark
brown; tergites II-VI with median stripe of subquadrate brown matte spots and 1+1
lateral rectangular-shaped blackish spots with distal rim grayish; VIII-IX yellowish-gray
brownish, pale and shiny; cibarium with 1+1 flat submedian lobes covered by acute teeth,
as in S. dandrettai; sensory vesicle of maxillary palp about 1.5-1.8 of palpomere length;
wing basal sector of R without hair; claw without subbasal tooth; paraproct truncate with
small dorsal projection similar to that of S. dugesi; eighth sternite with 22-28 setae on each
side; gonapophysis divergent with internal rim slightly sclerotized. (Fig. 92 O).
Male: Wing length 1.8 mm; scutum black with 1+1 anterior subtriangular silvery
spots continued posteriorly as thin stripes, surpassing half of scutum length, laterally
grayish (Fig. 93 A); pilosity golden shiny; abdomen black with 1+1 silvery spots on tergites
II, VI, VII; ratio of length/width of hind basitarsus = 6; gonostylus slightly acuminate on
distal inner portion as in S. dugesi; ventral plate wider than long, slightly acuminate distally,
very similar to S. haematopotum and S. trivittatum.
Pupa: Cocoon slipper shaped, slightly elongated basally (Fig. 95 C), threads evident;
length basally 2.5-2.7 mm, dorsally 2.2-2.5 mm, gills 1.5-1.6 mm; frontoclypeus and thorax
with moderate to sparse tubercles, with rounded apical rim; frontal trichomes single and
relatively short; gill with 8 branches, with primary ventral branch relatively elongated.
Larva: Maximum length 5.0-5.5 mm; cephalic apotome as in S. escomeli, darkened
median and basally with median and 1+1 lateral and subbasal light windows; mandible
with 1 latero-mandibular process; antenna surpassing cephalic fan stem length; ratio of
proximal, medial and distal segments = 1:1.3:1.5; cephalic fan with 52-56 rays; postgenal
bridge slightly shorter than hypostomium length (Fig. 97 K); lateral sclerite of thoracic
proleg with about 17 teeth; anal sclerite without hair or scales, similar to S. zempoalense;
anal ring with 68-77 rows of 14-16 hooks each; rectal papillae with 3 single lobes; ninth
segmente with 1 pair of ventral tubercles.
Distribution: Mexico: Chihuahua, Coahuila, San Luis Potosi, Tamaulipas, Veracruz;
USA: Texas.
Bionomics: Simulium mediovittatum larvae live on aquatic vegetation of rivers and
shallow streams. Females bite humans, livestock and wild mammals.
Discussion: The closest species to S. mediovittatum are S. dugesi and S. ochoai. The first
species can be differentiated by the greater number of frontoclypeal and thoracic
tubercles. Simulium ochoai has gill branching nearer the base and more open. These small
442 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

differences, plus the coloration and number of anal ring hook rows of S. mediovittatum,
suggest that it would be useful to compare more material of those species and to also use
additional nonmorphological studies.

Simulium (Psilopelmia) bivittatum MALLOCH (Figs. 91 K, T; 92 B; 93 I; 94 B; 95 D; 96 C.)


Simulium bivittatum MALLOCH, 1914: 31; VARGAS, 1945a: 120.
Simulium (Psilopelmia) bivittatum: VARGAS & DÍAZ NÁJERA, 1958: 13; PETERSON, 1993: 308; COSCARÓN,
1987: 30; PETERSON & KONDRATIEFF, 1995: 35-36; COSCARÓN et al., 1996: 54, 55; 63; CROSSKEY
& HOWARD, 1997: 58; 2004: 53; MOULTON, 1998: 50; ADLER et al., 2004: 349-350.
Simulium idahoense TWINN, 1938: 50-51.
Female: Wing length 2.1-2-3 mm; frons and clypeus with silver pruinosity; scutum
yellowish orange to reddish brown arranged in 1 median and 1+1 sublateral stripes
separated by 1+1 submedian light gray whitish vittae, scutellum brown yellowish and
metanotum black-brown grayish; abdomen grayish with dark brown subquadrate spots on
median line and 1+1 rectangles laterally; legs yellow-grayish with apex of femora, tibiae
and tarsomeres dark brown; cibarium with sclerotized rim with blunt teeth (Fig. 91 K);
basal sector of R glabrous; claw without subbasal tooth (Fig. 91 T); eighth sternite with
20-23 setae; paraproct with short distal projection, with acute apex (Fig. 92 B).
Male: Wing length 2.0-2.1 mm; scutum blackish to reddish brown covered with
recumbent subbasal golden pile; with anterior light showing 1+1 anterior cuneiform
silvery spots, not reaching half of scutum length; scutellum dark yellowish brown, pleurae
dark grayish; metanotum brownish black, grayish pruinose; legs mostly dark brown;
gonostylus subrectangular with small apical concavity (Fig. 93 I); ventral plate short
without apical projection (Fig. 94 B).
Pupa: Cocoon slightly projected anteriorly at base, anterior border reinforced, (Fig.
95 D), threads evident; length along base 2.4-2.7 mm, on dorsum 2.2-2.5 mm, gills 1.9-2.1
mm; frontoclypeus and thorax with tubercles relatively sparse elongated, as long as wide at
base, with rounded apex and some of them acute; trichomes with 1 or 2 branches; gills
with 8 branches, with primary and secondary branches short (Fig. 96 C).
Larva: Maximum length 3.7-4.5 mm; cephalic apotome scarcely pigmented, only
slightly darkened basally; antenna slightly longer than cephalic fan stem; ratio of proximal,
medial and distal segments = 1:1.4:1.6; cephalic fan with 30-38 rays; mandible with latero-
mandibular process; postgenal cleft dome shaped; postgenal bridge slightly shorter than
hypostomium; anal sclerite without scales and very small hairs; anal ring with about 70
rows with 13-16 hooks each; rectal papillae without diverticula on lobes.
Distribution: Canada: Alberta, Saskatchewan; Mexico: Chihuahua, Coahuila, San
Luis Potosi, Tamaulipas, Veracruz, Zacatecas; U.S.A.: Arizona, Colorado, Idaho, Montana,
Nebraska, New Mexico, Oregon, South Dakota, Texas, Utah, Washington, Wyomming.
Bionomics: Simulium bivittatum breeds in rivers with bottoms of rocks and gravel in
lowlands to low foothills. Females bite humans and livestock, and the attacks can make
people stay indoors.
Discussion: Based on genitalia and cibarium characters, S. bivittatum constitutes a
group with S. mediovittatum, S. dugesi and S. ochoai, but the scutum coloration and
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 443

A B E
D
C

G
F
I L

H
K

P N

O
R
S

Figure 93. Male. A-G: Scutum; A: S. (Psilopelmia) dugesi; B: S. (P.) iracouboense; C: S. (P.) escomeli; D: S.
(P.) veracruzanum; E: S. (P.) zempoalense; F: S. (P.) mangabeirai; G: S. (P.) gonzalezherrejoni. H-S: Gonocoxite
and gonostylus; H: S. (P.) dugesi; I: S. (P.) bivittatum; J: S. (P.) samboni; K: S. (P.) iracouboense; L: S. (P.)
haematopotum; M: S. (P.) escomeli; N: S. (P.) downsi; O: S. (P.) dandrettai; P: S. (P.) veracruzanum; Q: S. (P.)
mangabeirai; R: S. (P.) zempoalense; S: S. (P.) callidum.
444 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

ornamentation of the adutls permit it to be differentiated. The presence of yellowish


females and blackish males is similar to the condition for S. escomeli, but the other
morphological characters allow identification.

Simulium (Psilopelmia) samboni JENNINGS (Figs. 91 B, L; 92 C, Q; 93 J; 95 E, R; 97 A.)


Simulium samboni JENNINGS, 1915: 67; FAIRCHILD, 1940: 704-705; VARGAS, 1942a: 240; RAMÍREZ
PÉREZ, 1983: 67; SHELLEY et al., 1984: 155-156.
Simulium (Psilopelmia) samboni: VARGAS & DÍAZ NÁJERA, 1957b, 356, 358:; COSCARÓN, 1987: 37;
COSCARÓN et al., 1995: 292-294; COSCARÓN et al., 1996: 56-59; CROSSKEY & HOWARD, 1997: 59;
2004: 53; SHELLEY et al., 2002: 169.
Simulium (Lanea) samboni VARGAS et al., 1946: 107; VARGAS & DÍAZ NÁJERA, 1948b: 343; DALMAT,
1955: 151-156.
Simulium colvini DALMAT, 1952: 344-347; 131-135; VARGAS & DÍAZ NÁJERA, 1954: 62-63.
Female: Wing length 1.9-2-1 mm; scutum, scutellum and metanotum yellowish, the
first with 1+1 silvery submedian stripes reaching posterior border (Fig. 91 B), and covered
by sparse yellow pile; metanotum with anterolateral areas brownish with gray pruinosity;
abdomen with segments I-II yellowish, III-VI with tergal plate and lateral areas dark
brown but separated by narrow stripes determining 3 rows of subquadrate plates as in S.
escomeli; pleural area light grayish; legs yellowish, darkened on tarsomeres and apex of hind
tibia and femur; cibarium with 1+1 slight submedian elevations, covered with small teeth
(Fig. 91 L); fronto-ocular triangle about as wide as high (Fig. 92 Q); basal sector of R with
sparse hair arranged in 1 row; claw without subbasal tooth; eighth sternite with 14-16
setae; paraproct with short distal projection (Fig. 92 C).
Male: Wing length 1.8-2.0 mm; coloration as in female, but with scutum dark and
submedian silvery stripes not surpassing half of scutum length; abdomen with segments
(III) IV-VI totally dark brown; gonostylus subrectangular, slightly longer than half of
gonocoxite length (Fig. 93 J); ventral plate as in S. escomeli.
Pupa: Cocoon with anterior border reinforced and small projection basal and dorsal
(Fig. 95 E); weave compact with evident threads; length along base 2.2-2.5 mm, on
dorsum 2.0-2.3 mm, gills 2.0-2.1 mm; frontoclypeus and thorax with moderate number of
tubercles (Fig. 95 R), cephalic and thoracic trichomes bifid and elongated; gills with 8
branches, with short dorsal primary and secondary branches.
Larva: Maximum length 5.0-5.2 mm; cephalic apotome with median and 1+1
subbasal windows (Fig. 97 A); antenna surpassing apex of cephalic fan stem; ratio of
proximal, medial and distal segments = 1:0.9-1:1.3-1.5; cephalic fan with 34-36 rays;
postgenal bridge about same length as hypostomium; anal sclerite with single hairs; anal
ring with about 70 rows with 12-13 hooks each; ninth segment with pair of ventral
tubercles; rectal papillae with 1-3 diverticula on each lobe.
Distribution: Belice: Belice, Cayo, Stan Creek, Toledo; Guatemala: Altaverapaz, El
Petén, Escuintla, San Marcos, Santa Rosa, Solalá, Totonicapán; Honduras: Cortés;
Mexico: Chiapas, Tabasco, Oaxaca, Veracruz; Panama: Canal Zone, Chucumaque, Coclé;
Trinidad; Venezuela: Anzoategui, Barinas, Carabobo, Federal District, Guarico, Miran-
da, Monagas; Portuguesa, Tachira, Yaracuy.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 445

Bionomics: Simulium samboni breeds in small to medium-sized creeks, with moderate


to fast-flowing water, attached to aquatic vegetation. Females are zoophilic.
Discussion: The gonostylus shape of S. samboni permits it to be separated from
other S. (Psilopelmia) species. Simulium escomeli is similar but the elongated distal projection
of the paraproct plus the subtrapezoidal gonostylus permit differentiation.

Simulium (Psilopelmia) iracouboense FLOCH & ABONNENC


(Figs. 91 H, M, R, U; 92 D, P; 93 B, K; 94 C; 95 F; 97 B, Q, S.)
Simulium iracouboense FLOCH & ABONNENC, 1946b: 7-12; HAMADA & GRILLET, 2001: 29.
Simulium (Psilopelmia) iracouboense: COSCARÓN, 1987: 31; COSCARÓN et al., 1996: 59-60; SHELLEY et al.,
1997: 30; CROSSKEY & HOWARD, 1997: 59; 2004: 53.
Simulium sucamense NUNES DE MELLO, 1974: 41; PY-DANIEL, 1989a: 254.
Simulium santaelenae RAMÍREZ PÉREZ & PETERSON, 1981: 161; RAMÍREZ PÉREZ, 1983: 20; PY-DANIEL,
1989a: 254.
Female: Wing length 1.5-1.6 mm; scutum, scutellum, metanotum and pleura brown-
yellowish, the first with 1+1 submedian and lateral whitish arrangement as in S. escomeli;
abdomen brown-yellowish with 1 median and 1+1 sublateral blackish subquadrate spots
(Fig. 91 H); legs brown-yellowish with apex of femora. tibiae and tarsomeres dark brown;
fronto-ocular triangle wider than high (Fig. 92 P); sensory vesicle of maxillary palp about
half as long as palpomere (Fig. 91 R); cibarium with 2 protuberances with acute teeth (Fig.
91 M); basal sector of R with hair; claw without subbasal tooth (Fig. 91 U); paraprocts
subquadrate, rounded apically and well-developed basal lobe (Fig. 92 D).
Male: Wing length 1.5 mm; scutum brown yellowish with yellow pile; with certain
lighting, 1+1 submedian cuneiform spots on anterior third visible (Fig. 93 B); legs and
abdomen as in female, but abdominal segments (III) VI-VII darker; gonostylus subtrape-
zoidal, and about 2/3 gonocoxite length (Fig. 93 K). Ventral plate wider than long, slightly
acuminate apically (Fig. 94 C).
Pupa: Cocoon with loose weave, with very obivous threads and dorsally slightly
reinforced (Fig. 95 F); length basally 2.4-3.0 mm, dorsally 2.1-2.4 mm, gills 1.3-2.5 mm;
frontoclypeus and thorax with smooth tubercles in moderate number; cephalic trichomes
single, thoracic single or bifurcated; gills with 8 filiform branches, with primary and
secondary branches arising close to base.
Larva: Maximum length 4.1-4.5 mm; cephalic apotome with darkened median basal
area circumscribing median subbasal windows (Fig. 97 B); antenna surpassing apex of
cephalic fan stem; ratio of proximal, medial and distal segments = 1:1.1-1.3:1.6-1.7;
cephalic fan with 43-48 rays; postgenal bridge slightly shorter than hypostomium length;
anal sclerite without scales and very sparse hair (Fig. 97 Q); anal ring with about 63 rows
with 15-18 hooks each; rectal papillae with 2-3 diverticula on each lobe (Fig. 97 S).
Distribution: Brazil: Amapa, Pará, Roraima; French Guiana; Surinam; Venezuela.
Bionomics: Unknown.
Discussion: The closest species to S. iracouboense is S. samboni, but it can be
differentiated by the absence of hairs on the basal sector of R, presence of a fronto-
ocular triangle, cephalic trichomes bifid, cocoon with compact weave and more
446 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

A B F

D
N
E

H J

K
C I

M G L

Figure 94. Male. Ventral plate. A: S. (Psilopemia) dugesi. B: S. (P.) bivittatum. C: S. (P.) iracouboense. D:
S. (P.) haematopotum. E: S. (P.) panamense. F: S. (P.) callidum. G: S. (P.) escomeli. H: S. (P.) downsi. I: S. (P.)
dandrettai. J: S. (P.) bobpetersoni. K: S. (P.) veracruzanum. L: S. (P.) mangabeirai. M: S. (P.) gonzalezherrejoni.
N: S. (P.) trivittatum.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 447

A
C
E

B
G

D I J

L H

N
O

P
U
Q
R

S
T
W V
Figure 95. Pupa. A-N: General aspect; A: S. (Psilopelmia) dugesi; B: S. (P.) ochoai; C: S. (P.) mediovittatum; D:
S. (P.) bivittatum; E: S. (P.) samboni; F: S. (P.) iracouboense; G: S. (P.) haematopotum; H: S. (P.) downsi; I: S. (P.)
dandrettai; J: nr. S. (P.) dandrettai; K: S. (P.) bobpetersoni; L: S. (P.) veracruzanum; M: S. (P.) mangabeirai; N: S. (P.)
zempoalense. O: Dorsal portion of thoracic integument of S. (P.) zempoalense. P-V: Frontoclypeus; P: S. (P.)
dugesi; Q: S. (P.) ochoai; R: S. (P.) samboni; S: S. (P.) escomeli; T: S. (P.) haematopotum; U: S. (P.) bobpetersoni; V:
S. (P.) panamense. W: Frontoclypeal tubercles, enlarged, of S. (P.) panamense.
448 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

reinforced anterior border and larvae with a small number of primary rays and anal
ring with more rows.

Simulium (Psilopelmia) haematopotum MALLOCH (Figs. 91 C, S; 92 E; 93 L; 94 D; 95 G.)


Simulium haematopotum MALLOCH, 1914: 61; DYAR & SHANNON, 1927: 26; FAIRCHILD, 1940: 704;
IRIARTE, 1943: 191: LANE & VULCANO, 1943: 439: VARGAS, 1945a: 142.
Simulium (Lanae) haematopotum: DALMAT, 1955: 144.
Psilopelmia haematopotum: RUBTZOV & GARCÍA AVILA, 1972: 22.
Simulium (Psilopelmia) haematopotum: COSCARÓN, 1987: 31; COSCARÓN et al., 1996: 60; CROSSKEY &
HOWARD, 1997: 59; 2004: 53.
Simulium pseudohaematopotum HOFFMANN, 1930b: 293.
Simulium boydi DE LEÓN, 1945: 75.
Simulium marathrumi FAIRCHILD, 1940: 715.
Female: Wing length 2.0-2.2 mm; coloration generally brownish; scutum ornament-
ed by median and 1+1 sublateral black stripes, separated by 1+1 thick submedian silvery
pruinose vittae and bordered anteriorly, laterally and posteriorly by thin silvery pruinose
area (Fig. 91 C), hair grayish to yellow in accordance with light position; scutellum and
metanotum brownish; legs grayish, blackish on apex of femur, tibia and tarsomeres of
fore and mid legs, and more expanded darker area on hind femur and tibia; abdomen
grayish, with tergites I-II blackish with posterior border pale gray, III-VI with median row
of subquadrate dark brown plates, and III-IV with 1+1 lateral row of subrectangular
brown-grayish to blackish spots covering anterior 2/3 of tergites; segments V-VI laterally
grayish pollinose; VIII-IX grayish brown shiny; cibarium with 1+1 short sublateral
elevations, covered by teeth as is median depressed area, similar to S. bivittatum but with
more acute teeth; sensory vesicle of maxillary palp shorter than half of palpomere (Fig.
91 S); basal sector of R glabrous; claw without subbasal tooth; eighth sternite with 28-30
setae; paraproct blunt apically and with large basal lobe (Fig. 92 E).
Male: Wing length 1.8-2 mm; scutum black with 1+1 silvery pruinose cuneiform
anterior stripes, extending about 1/4 of scutum length, but with changing light position,
continued as thin stripes to posterior border; abdomen blackish with 1+1 lateral silvery
spots on segments II, V-VII; gonostylus about as long as gonocoxite, becoming thin on
distal 1/3 with enlarged apical spur (Fig. 93 L); ventral plate slightly acuminate distally
(Fig. 94 D).
Pupa: Cocoon compact with anterior border slightly reinforced and with small
dorsal and ventral projections (Fig. 95 G); threads evident; length basally 2.8-3.0 mm,
dorsally 2.4-2.5 mm; gills about 1.5 mm; frontoclypeus and thorax without or with very
sparse tubercles; cephalic and thoracic trichomes single and bifid; gills with 8 branches,
arranged as in S. escomeli.
Larva: Maximum length 4.5-5.0 mm; cephalic apotome ornamentation very similar
to that of S. escomeli; antenna surpassing apex of cephalic fan stem; ratio of proximal,
medial and distal segments = 1:0.9-1:1.3; postgenal cleft slightly short than hypostomium
length; ninth segment with pair of ventral tubercles; anal sclerite without scales; anal ring
with 74-80 rows, with 14-17 hooks each; rectal papillae with 1-3 diverticula on each lobe.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 449

Distribution: Colombia; Costa Rica: Valle Central; Cuba: Las Villas, Oriente, Pinar
del Río; Guatemala: Altaverapaz, Chimaltenango, Chiquimula, El Petén, El Progreso,
Escuintla, Guatemala, Huehuetenango, Izabal, Quezaltwenango, San Marcos, Santa Rosa;
Guyana: Essequibo; Mexico: Chiapas, Colima, Guerrero, Jalisco, Michoacán, Morelos,
Nayarit, Oaxaca, Puebla, Sinaloa and Veracruz; Panama: Canal Zone, Coclé, Colón,
Panama; Puerto Rico: Río Piedras; Venezuela: Bolívar.
Bionomics: Simulium haematopotum breeds in shallow water. Females are anthropo-
philic in Mexico and Guatemala.
Discussion: Simulium haematopotum has female abdominal ornamentation and cibarium
and paraproct shape similar to those of S. bivittatum, S. dugesi, S. ochoai, S. mediovittatum and S.
samboni, but can be differentiated by the scutum and larval cephalic apotome. Simulium
samboni shows similar scutum design but with yellowish coloration. Simulium marathrumi has
similar ornamentation of the scutum and abdomen, and morphology of the wings and legs
Based on the variability of these characters, S. marathrumi can be considered a synonym, but
it needs to be confirmed with more material and the use of cytological or molecular studies.
CROSSKEY & HOWARD (2004: 53) consider it a synonym of S. haematopotum.
Simulium benjamini is very close to S. haematopotum, showing differences such as the
elongated secondary dorsal and median pupal gill branches and a big subbasal tooth on
the female claw, according to the drawings of DALMAT. These characters could represent
variants, because in a pharate female paratype from the USNM, the toothed claw is not
evident. We consider that S. benjamini could be a synonym of S. quadrivittattum, but it
would be convenient to see more material in order to confirm if they are conspecific.

Simulium (Psilopelmia) escomeli ROUBAUD (Figs. 91 E, I; 92 H; 93 C, M; 94 G; 95 S; 96 G; 97 E.)


Simulium escomeli ROUBAUD, 1909: 428-429; SILVA FIGUEROA, 1917: 30; WYGODZINSKY, 1953b: 330-332.
Psilopelmia rufidorsum ENDERLEIN, 1934b: 283.
Simulium (Lanea) escomeli: VARGAS & DÍAZ NÁJERA, 1951: 152.
Simulium (Psilopelmia) escomeli: VARGAS & DÍAZ NÁJERA, 1953a: 17-21; WYGODZINSKY, 1971: 2-12;
COSCARÓN, 1987: 31; 1991: 266-267; SHELLEY et al., 1989: 97; COSCARÓN et al., 1996: 73-76;
COSCARÓN et al., 1995: 289-295; COSCARÓN ARIAS, 2003: 198, 204, 209, 215; CROSSKEY &
HOWARD, 1997: 58; 2004: 53; COSCARÓN ARIAS, 2003: 195.
Female: Wing length 2.3-2.8 mm; general coloration yellowish, some specimens
brownish to blackish; frons and clypeus grayish pollinose; scutum yellow to blackish with
1+1 anterior scutal border and thick submedian silvery stripes reaching posterior area
(Fig. 91 E); prescutellar area brownish; lateral and pleural areas brown with gray
pruinosity; scutellum yellowish brown, metanotum dark brown with silvery pruinosity;
legs yellowish, with apex of femora, tibiae and tarsomeres dark brown; abdomen light
yellow, segment II with brown tergal anterior spot; III-VIII with 1 blackish tergal
subquadrate median, 1+1 sublateral and 1+1 small lateral spots, except VII without
median spot (Fig. 91 I); cibarium with 1+1 small lateral protuberances with thin teeth, and
median portion depressed, smooth, similar to shape in S. trivittatum; sensory vesicle of
maxillary palp about 1/3 of palpomere length; fronto-ocular triangle about as high as
wide; basal sector of R glabrous; claw without subbasal tooth; paraproct strongly
450 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

B
P D

L M

I
J

H
F K

O A

Figure 96. Pupa. Gills and thoracic trichomes. A: S. (Psilopelmia) dugesi. B: S. (P.) ochoai. C: S. (P.)
bivittatum. D: S. (P.) panamense. E: S. (P.) callidum. F: S. (P.) pseudocallidum. G: S. (P.) escomeli. H: S. (P.)
downsi. I: S. (P.) dandrettai. J: nr. S. (P.) dandrettai (also with filament very enlarged). K: S. (P.)
bobpetersoni. L: S. (P.) veracruzanum. M: S. (P.) mangabeirai. N: S. (P.) gonzalezherrejoni. O: S. (P.)
longithallum. P: S. (P.) trivittatum. Q: S. (P.) zempoalense.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 451

projected distally and with well-developed basal posterior protuberance (Fig. 92 H); eighth
sternite with 20 setae on each side.
Male: Wing length 2.2-2.6 mm; scutum velvety black with 1+1 silvery anterior
submedian spots not surpassing anterior half of scutum length (Fig. 93 C) (some
specimens with scutum orange brown to reddish); scutellum light brown and metanotum
black with silvery pruinosity; abdomen black with segment I lighter, II, VI-VIII with 1+1
silvery lateral spots; gonostylus subtrapezoidal and slightly shorter than gonocoxite (Fig.
93 M); ventral plate slightly constricted laterally (Fig. 94 G).
Pupa: Cocoon with reinforced anterior border straight to slightly projected dorsally;
weave thin with evident threads; length basally 3.0-3.5 mm, dorsally 2.2-3.0 mm, gills 2.2-
2.5 mm; frontoclypeus and thorax with numerous smooth tubercles (Fig. 95 S); gills with
8 filamentous branches, relatively open basally (Fig. 96 G).
Larva: Maximum length 5.5-6.0 mm; cephalic apotome with centro-basal and 1+1
basal windows (Fig. 97 E); antenna as long as stem of cephalic fan; ratio of proximal,
medial and distal segments = 1:1.2-1.3:1.1-1.6; cephalic fan with 33-38 rays; postgenal
bridge 3/4 of hypostomium length; sclerite comb of thoracic proleg with about 30 teeth
arranged in 11-12 groups; anal ring with 63-70 rows, with 10-13 hooks each; ninth
segment without ventral tubercles; rectal papillae with 0-3 diverticula on each lobe.
Distribution: Chile: Tarapacá; Colombia; Ecuador: Azuay, Cañar, Carchi, Chimb-
orazo, Cotopaxi, Esmeraldas, Imbabura, La Unión, Los Ríos, Pichincha, Tungurahua;
Peru: Ancash, Arequipa, Lima, Tacna.
Bionomics: Simulium escomeli breeds in clear, fast flowing-streams of regular depth,
attached to aquatic vegetation or stones. Females are anthropophilic and a dangerous pest
in the valley of the Pacific desert.
Discussion: Simulium escomeli is closest to S. downsi, having similar adult coloration
and cibarial shape; some differences are in the paraproct, with a slight posterodorsal
protuberance (Fig. 92 I), and the gonostylus with a more pronounced external median
concavity (Fig. 93 N).

Simulium (Psilopelmia) downsi VARGAS, MARTÍNEZ PALACIOS & DÍAZ NÁJERA


(Figs. 92 I; 93 N; 94 H; 95 H; 96 H; 97 F, P.)
Simulium (Lanea) downsi VARGAS et al., 1946: 126-129; DALMAT, 1955: 136-140.
Simulium downsi: VULCANO, 1967: 10.
Simulium (Psilopelmia) downsi: VARGAS & DÍAZ NÁJERA, 1957b: 154; COSCARÓN, 1995: 294-295,
COSCARÓN, 1987: 31; COSCARÓN et al., 1996: 76-78; CROSSKEY & HOWARD, 1997: 58; 2004: 53.
Female: Wing length 2.3 mm; general coloration yellow; scutum yellow with
brownish pile, with 1+1 silvery submedian vittae reaching posterior border, and lateral
border light gray, ornamented as in S. escomeli; scutellum light yellowish, metanotum
brownish; pleurae light gray with brown episternum; legs yellowish with apex of femora,
tibiae and tarsomeres blackish; abdomen with segments I-II yellowish, III-IX brown-
yellowish; tergites III-VIII with 5 longitudinal rows of blackish subquadrate spots as in S.
escomeli: median row with wider spots, 1+1 sublateral and 1+1 lateral; the last with smaller
spots; median spots absent on tergites VII-VIII; last segments shiny; cibarium with small
452 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

A B C D

E F G H O

J K I P

M N

Q R
S

Figure 97. Larva. A-H: Cephalic apotome; A: S. (Psilopelmia) samboni; B: S. (P.) iracouboense; C: S. (P.)
panamense; D: S. (P.) callidum; E: S. (P.) escomeli; F: S. (P.) downsi; G: S. (P.) mangabeirai; H: S. (P.)
zempoalense. I: Mandible apex of S. (P.) ochoai. J-N: Hypostomium and postgenal cleft; J: S. (P.) dugesi;
K: S. (P.) mediovittatum; L: S. (P.) mangabeirai; M: S. (P.) trivittatum; N: S. (P.) zempoalense. O-P:
Antenna; O: S. (P.) panamense; P: S. (P.) downsi. Q-R: Anal sclerite; Q: S. (P.) iracouboense; R: S. (P.)
pseudocallidum. S: Anal lobes and diverticula of rectal papillae of S. (P.) iracouboense.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 453

teeth on 1+1 submedian protuberances as in S. trivittatum; basal sector of R with 2-3 rows
of hair; claw without subbasal tooth; paraprocts strongly acuminate mediodistally and
with posterobasal protuberance (Fig. 92 I); eighth sternite with 20-28 setae on each side.
Male: Wing length 2.1 mm; scutum coloration as in female, scutum with 1+1
submedian cuneiform stripes only reaching to half of scutum length; gonostylus with
strong distal curvature (Fig. 93 N); ventral plate with body subtriangular (Fig. 94 H).
Pupa: Cocoon with compact structure, slightly projected basally (Fig. 95 H); length
basally 3.0-3.3 mm, dorsally 2.7-3.0 mm, gills 1.8-2.0 mm; frontoclypeus and thorax with
moderate number of tubercles as in S. ochoai; trichomes single or bifid; gills with 8
branches, with branching not close to base (Fig. 96 H).
Larva: Maximum length 5.3-5.5 mm; cephalic apotome darkened mediobasally with
1 light window medially and 1+1 subbasally (Fig. 97 F); antenna surpassing cephalic fan
stem apex; ratio of proximal, medial and distal segments = 1:1-1.1:1.5 (Fig. 97 P); cephalic
fan with 31-36 rays; postgenal bridge about same length as hypostomium; anal ring with
about 70 rows, with 13-15 hooks each; rectal papillae with 4-6 diverticula on each lobe;
anal sclerite without scales; comb of proleg lateral sclerite with 30-36 teeth.
Distribution: Guatemala: Altaverapaz, Bajaverapaz, Chimaltenango, Escuintla, Hue-
huetenango, Jalapa, Jutiapa, El Petén, El Progreso, El Quiché, Retalhuleo, San Marcos; Santa
Rosa, Sololá, Suchitepequez; Mexico: Chiapas, Oaxaca; Venezuela: Miranda.
Bionomics: Unknown.
Discussion: Simulium escomeli is very close to S. downsi, but can be differentiated by
the absence of hair on the basal sector of R, the darker female scutum, especially on the
posterior area, the wider submedian silvery stripes, and the gonostylus without a
distolateral outer expansion. The male of S. callidum is similar externally, but the ventral
plate is subovoidal.

Simulium (Psilopelmia) dandrettai VARGAS, MARTÍNEZ PALACIOS & DÍAZ NÁJERA


(Figs. 91 Q; 92 J; 93 O; 94 I; 95 I, J; 96 I, J.)
Simulium (Lanea) dandrettai VARGAS et al., 1946: 124-126; VARGAS & DÍAZ NÁJERA, 1948b: 323, 342-343.
Simulium (Psilopelmia) dandrettai: VARGAS & DÍAZ NÁJERA, 1957b: 183; COSCARÓN, 1987: 31;
COSCARÓN et al., 1996: 78-81; CROSSKEY & HOWARD, 1997: 58; 2004: 53.
Female: Wing length 2.4-3.0 mm; frons and clypeus with gray pruinosity; scutum
brown with 1+1 submedian and 1+1 lateral whitish stripes bordering median and 1+1
sublateral brown to black vittae; whitish submedian vittae, in some illumination, shows
dark transverse anterior spot, and with posterior light they join posteriorly, changing to
gray-brown; scutum hair golden to silvery, according to light position; scutellum pale
brown; metanotum dark brown grayish; legs brown yellowish with femorae, tibiae and
tarsomeres dark brown to black; abdomen with tergites I-II brown yellowish with dark
spot on median row and 1+1 slightly lighter laterally; VII-VIII shiny light brown grayish
on median area and darker laterally; cibarium with median depression bordered by 1+1
submedian short projections apically covered by spines (Fig. 91 Q); fronto-ocular triangle
higher than wide; basal sector of R without hair; claw without subbasal tooth; eighth
sternite with 18-24 setae; paraproct short (Fig. 92 J).
454 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Male: Scutum black with 1+1 anterior submedian shiny silvery spots not surpassing
1/3 of scutum length, similar to S. veracruzanum in shape; in posterior illumination, with
lighter subquadrate median posterior spot; abdomen black with 1+1 lateral silvery spots
on tergites II-VI; ratio of length/width of hind basitarsus = 4; gonostylus very wide
distally (Fig. 93 O); ventral plate rounded, without lateral constriction (Fig. 94 I).
Pupa: Cocoon with weave consistent, reinforced on anterior margin (Fig. 95 I);
length basally 2.7-3.0 mm, dorsally 3.0-3.1 mm, gills 3.0-3.2 mm; gills dark brown, with 6
branches, dorsal secondary and 2 dorsal tertiary branches longer and thicker than other
branches (Fig. 96 I).
Larva: Cephalic apotome dark on base, with light spots; ratio of proximal, medial and
distal segments = 1:1.3:1.5-1.6; cephalic fan with 26-31 rays; postgenal cleft not deep,
postgenal bridge longer than hypostomium, ratio of postgenal bridge/hypostomium = 1.2;
anal sclerite with hair, but not scales; anal ring with 64-68 rows, with 13-14 hooks each.
Distribution: Mexico: Distritop Federal, Mexico, Oaxaca, Veracruz.
Bionomics: Simulium dandrettai breeds in small clear torrential creeks in mountain areas at
2000-3000 m, attached to aquatic vegetation and rocks. Female feedindg habits are unknown.
Discussion: The pupal gill with 6 branches, thicker dorsal secondary and tertiary
branches, apically enlarged gonostylus and rounded ventral plate permit S. dandrettai to be
differentiated from other Neotropical S. (Psilopelmia) species. Material collected from the
same place as S. dandrettai has cocoons with longitudinal low crest like folds (Fig. 95 J).
The dorsal gill branches are not too much thicker than the other branches (Fig. 96 J) and
the tubercles of the frontoclypeus and exposed portion of the thorax have a wrinkled
apical border. COSCARÓN et al., 1996: 81 suggest that the material could be a different
species, but it was not named as new until more material could be studied in order to
confirm the differences; this point of view is maintained here.

Simulium (Psilopelmia) zempoalense VARGAS, MARTÍNEZ PALACIOS & DÍAZ NÁJERA


(Figs. 93 R; 95 N, O; 96 Q; 97 H, N.)
Simulium (Lanea) zempoaelense VARGAS et al., 1946: 107, 112,136, 138,168, 170-171.
Simulium (Psilopelmia) zempoaelense: VARGAS & DÍAZ NÁJERA, 1957b: 183-184, 187-188; COSCARÓN,
1987: 32; COSCARÓN et al., 1996: 97-98; CROSSKEY & HOWARD, 1997: 59; 2004: 53.
Female: Wing length 2.8-3.0 mm; thorax coloration similar to S. dandrettai; with
lateral light sub-median vittae shown dark gray opaque; pleurae light gray; abdomen with
dorsal spots ornamentation like S. escomeli; basal sector of R, claws and paraprocts as S.
dandrettai but the last with basal lobe more prominent.
Male: Wing length 2.5 mm; scutum black with 1+1 cuneiform vittae reaching thinly
to posterior area; silvery pile; abdomen with 1+1 lateral silvery spots on tergites II, V-VI;
hind basitarsus width and ventral plate shape as in S. dandrettai; gonostylus strongly
acuminate apically (Fig. 93 R).
Pupa: Cocoon with reinforced border (Fig. 95 N); length basally 3.6 mm, dorsally 3.2
mm, gills 2.5-3.6 mm; frontoclypeus and thorax with wrinkled platelets and sparse
tubercles (Fig. 95 O); trichomes single to bifid; gills with 6 branches, with primary and
secondary branches bifurcated near base (Fig. 96 Q) with branches about same width.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 455

Larva: Maximum length 6.0-6.5 mm; cephalic apotome light brown, without visible
ornamentation (Fig. 97 H); ratio of proximal, medial and distal antennal segments =
1:1.3:1.8; cephalic fan with about 38-40 rays; postgenal cleft short, postgenal bridge longer
than hypostomium length (Fig. 97 N); mandible with 2 mandibular processes; anal sclerite
with numerous hairs, but not scales; anal ring with 76-80 rows, with 13-14 hooks each;
rectal papillae with 9-12 diverticula on each lobe.
Distribution: Mexico: Durango, Mexico, Morelos, Oaxaca, Veracruz.
Bionomics: Unknown.
Discussion: Simulium dandrettai is the closest species to S. zempoalense, with similar adults
and pupae with wrinkled integument and 6 gill branches. There are small differences in the
pupa such as the thicker dorsal external pro minence shown in S. dandrettai. It is necessary to
carry out cytological and molecular studies of this species as well as other material close to S.
dandrettai that might be considered possible new species (COSCARÓN et al., 1996).

Simulium (Psilopelmia) trivittatum MALLOCH (Figs. 91 P; 92 N; 94 N; 96 P; 97 M.)


Simulium trivittatum MALLOCH, 1914: 25, 30, 39, 68; DAMPF, 1927: 129; DYAR & SHANNON, 1927: 37;
LANE & VULCANO, 1943: 439.
Simulium (Psilopelmia) trivittatum: VARGAS & DÍAZ NÁJERA, 1957b: 154, 183, 185-186, 264, 360;
PETERSON, 1993: 341; COSCARÓN et al., 1996: 94-97; CROSSKEY & HOWARD, 1997: 59; 2004: 53;
MOULTON, 1998: 66; ADLER et al., 2004: 353.
Simulium distinctum MALLOCH, 1913: 133-134 (preoc.).
Simulium (Psilopelmia) mazzottii DÍAZ NÁJERA, 1979: 553-556.
Female: Wing length 2.0-2.5 mm; scutum ornamentation similar to that of S.
bobpetersoni, with 3 brown-blackish longitudinal vittae delimited by 1+1 submedian and 1+1
lateral light yellowish stripes; on anterior border of sublateral vittae are 1+1 subtrianglular
whitish shiny spots; scutellum and metanotum yellowish brown, the second changing to dark
brownish black; legs yellowish, with apex of femora, tibiae and tarsomeres brown; abdomen
yellowish with 5 rows of blackish subrectangular spots, 1 median and 2+2 at sides as is
common in S. (Psilopelmia); median rows present from I to VI, lateral rows from III to VII;
cibarium with median depression with sclerotized rim, flanked by 2 small protuberances
with short spines (Fig. 91 P); basal sector of R glabrous; claw without subbasal tooth; eighth
sternite with about 40 setae; paraproct with elongated distal projection (Fig. 92 N).
Male: Wing length 1.8-2.3 mm; scutum dark brown to blackish with 1+1 yellowish
submedian vittae reaching to posterior border; anterior border of submedian vittae with
light grayish subcircular spot; abdomen with segments I-II yellow, the following blackish;
sternites yellowish; gonostylus similar to that of S. escomeli but more elongated apically;
ventral plate acuminate distally with rounded apex (Fig. 94 N).
Pupa: Cocoon reinforced on anterior border, weave with threads evident; length
basally about 3.0 mm, dorsally 2.7 mm, gills 2.7-3.0 mm; frontoclypeus and thorax with
sparse tubercles; trichomes single to bifid; gills with 6 branches, with basal trunk and
primary branches relatively short, median secondary branch elongated (Fig. 96 P).
Larva: Maximum length 4.5 mm; cephalic apotome brownish basally, with light
window medially. Antennae dark apically on medial and distal segments; ratio of proximal,
456 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

medial and distal segments = 1:1.3:2; cephalic fan with 36 rays; postgenal bridge about
same length as hypostomium (Fig. 97 M) Anal ring with about 68 rows, with 14 hooks
each; rectal papillae with diverticulum on each lateral lobe.
Distribution: Guatemala; Mexico: Coahuila, Tamaulipas; USA: Oklahoma, Texas.
Bionomics: Simulium trivittatum breeds in shallow rivers and small ditches with
trailing vegetation. Female feeding habits are not well known. There is only one citation of
biting a human.
Discussion: Simulium trivittatum is similar to S. gonzalezherrejoni, S. longithallum, S.
mangabeirai, S. veracruzanum and S. bobpetersoni, but can be differentiated by the pupal gill of
8 branches, except S. mangabeirai, which has 4. As explained for S. bobpetersoni, some
specimens of S. trivittatum can have a gill with 8 branches.

Simulium (Psilopelmia) bobpetersoni COSCARÓN, IBÁÑEZ-BERNAL & COSCARÓN ARIAS


(Figs. 92 K; 94 J; 95 K, U; 96 K.)
Simulium (Psilopelmia) bobpetersoni COSCARÓN et al., 1996: 81-84; CROSSKEY & HOWARD, 1997: 58; 2004:
52; MOULTON, 1998: 66; ADLER et al., 2004: 353.
Simulium (Lanea) trivittatum VARGAS, 1943: 363; VARGAS et al., 1946: 169.
Simulium (Psilopelmia) trivittatum MALLOCH: DAMPF, 1927: 129; VARGAS & DÍAZ NÁJERA, 1957b: 183;
DALMAT, 1955: 156-159.
Female: Wing length 2.2-2.9 mm; scutum with 3 blackish vittae as 1 median and 1+1
sublateral vittae flanked by 1+1 whitish submedian and 1+1 lateral stripes; scutellum and
metanotum dark brown; abdomen with 1 median and 1+1 sublateral and lateral rows of dark
brown subrectangular spots; cibarium with 1+1 small submedian protuberances with short
spines similar to those of S. veracruzanum; basal sector of R glabrous; claw without subbasal
tooth; eighth sternite with 28-30 setae; paraproct with elongated distal projection (Fig. 92 K).
Male: Wing length 1.9-2.1 mm; scutum dark reddish brown, with 1+1 anterior
submedian cuneiform white grayish stripes, generally reaching to half of scutum length
but reaching to posterior border in some specimens; ratio of length/width of hind
basitarsus = 7.5; gonostylus similar to that of S. escomeli but wider distally; ventral plate
slightly projected distally (Fig. 94 J).
Pupa: Cocoon with straight and reinforced anterior border (Fig. 95 K), length basally
2.7 mm, dorsally 2.8 mm, gills 2.2 mm; frontoclypeus and thorax with sparse tubercles
(Fig. 95 U); trichomes short with 1-2 branches on frontoclypeus and 1-3 on thorax; gills
with 8 branches, with primary branches relatively elongated (Fig. 96 K).
Larva: Maximum length 5.5 mm; cephalic apotome darkened on median basal area, bell
shaped, with 1 central and 1+1 basal light spots; antennae as long as or shorter than cephalic
fan stem, ratio of proximal, medial and distal segments = 1:1.2:1.9; cephalic fan with about 28
rays; proleg lateral sclerite with about 22 teeth; anal sclerite without scales; anal ring with about
62 rows, with 11-12 hooks each; rectal papillae with 0-2 diverticula on each lobe.
Distribution: Guatemala; Mexico: Aguascalientes, Chiapas, Chihuahua, Coahuila,
Colima, Durango, Guanajuato, Guerrero, Jalisco, Michoacan, Morelos, Nueva León,
Oaxaca, Puebla, San Luis Potosí, Tamaulipas, Veracruz.
Bionomics: Unknown.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 457

Discussion: The closest species to S. bobpetersoni is S. veracruzanum, but this species


has a shorter bifurcation of the primary and secondary branches and a ventral plate with
the median distal portion more elongated. Males are similar to those of S. zempoalense but
the pupal gill has 6 branches. Simulium trivittatum, of which Mexican material is considered
to be S. bobpetersoni by COSCARÓN (1996), can be differentiated by the 8 pupal gill branches.
ADLER et al., (2004) found S. trivittatum pupae with 8 branches; therefore, it would be useful
to study more material and also to do cytological and molecular studies of this material.

Simulium (Psilopelmia) veracruzanum VARGAS, MARTÍNEZ PALACIOS & DÍAZ NÁJERA


(Figs. 92 L; 93 D, P; 94 K; 95 L; 96 L.)
Simulium (Lanea) veracruzanum VARGAS et al., 1946: 112, 133-136; DALMAT, 1951: 55-57; 1955: 33.
Simulium (Psilopelmia) veracruzanum VARGAS & DÍAZ NÁJERA, 1957b: 184-187, 189; COSCARÓN, 1987:
31; COSCARÓN et al., 1996: 84-87; CROSSKEY & HOWARD, 1997: 59; 2004: 53.
Female: Wing length 2.6-3.0 mm; scutum grayish with 1 median and 1+1 lateral dark
brown vittae; with frontal light, appearing to join posteriorly and showing as silvery the 1+1
submedian light vittae; pile grayish yellow; scutellum light yellowish brown, metanotum dark
grayish brown Pleura brownish yellow; legs yellowish gray with coxae, apex of hind femur,
apex of tibiae and tarsomeres blackish; abdomen brown with tergites I-II yellowish, tergites
II-VII grayish yellow with 5 rows of subquadrate dark brown grayish spots as 1 median, 1+1
sub-lateral and 1+1 lateral rows, as in S. escomeli. Basal sector of R, claw and cibarium as in S.
trivittatum; paraproct very elongated distally (Fig. 92 L).
Male: Wing length 2.7 mm; scutum with 1+1 silvery anterior cuneiform vittae reaching
about half of scutum length (Fig. 93 D); scutellum and metanotum black; pleurae dark gray;
legs brown-blackish with lighter areas on fore tibia, anterior half of middle tibia and tarsus,
anterior third of hind tibia and anterior half of basitarsus; gonostylus with strong distal
curvature (Fig. 93 P); ventral plate with deep lateral constrictions (Fig. 94 K).
Pupa: Cocoon with reinforced anterior border (Fig. 95 L), length basally 3.3 mm,
dorsally 2.8-3.0 mm, gills 3.0-3.4 mm; frontoclypeus and thorax with moderate number of
smooth tubercles; cephalic trichomes with 1-2 branches, thorax with 2-5 branches; gills
with 8 branches, with primary and secondary branches short (Fig. 96 L).
Larva: Maximum length 5.5-6.0 mm; cephalic apotome mildly darkened basally, with
light median and 1+1 subbasal areas; ratio of proximal, medial and distal antennal
segments = 1:1.3-1.4:1.6-1.7; cephalic fan with 32-43 rays; postgenal bridge about same
length as hypostomium; ninth segment with pair of ventral tubercles; anal sclerite with
hair and abundant scales; anal ring with 72-76 rows, with 13-14 hooks each.
Distribution: Belice; Guatemala: Acatenango, Chimaltenango; Mexico: Chiapas,
Chihuahua, Coahuila; Durango, Jalisco, Michoacán, Morelos, Nayarit, Oaxaca, Veracruz.
Bionomics: Unknown.
Discussion: Simulium veracruzanum females are similar to those of S. gonzalezherrejoni,
S. longithallum, S. mangabeirai, S. bobpetersoni and S. trivittatum. The male ventral plate of S.
veracruzanum with the strong lateral constrictions, allow it to be differentiated from those
species. The primary and secondary pupal gill branches are very short and also help
separate these species.
458 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Simulium (Psilopelmia) mangabeirai VARGAS (Figs. 91 F; 93 F, Q; 94 L; 95 M; 96 M; 97 G, L.)


Simulium mangabeirai VARGAS, 1945b: 76.
Simulium (Lanea) mangabeirai: VARGAS et al., 1946: 107; VARGAS, 1948: 309.
Simulium (Psilopelmia) mangabeirai: VARGAS & DÍAZ NÁJERA, 1957b: 184; COSCARÓN, 1987: 31;
COSCARÓN et al., 1996: 87-89; CROSSKEY & HOWARD, 1997: 59; 2004: 53.
Female: Wing length 2.8-2.9 mm; scutum with 3 wide black vittae as 1 median and 2
sublateral vittae separated by 1+1 thinner light grayish submedian stripes; black sublateral
vittae not reaching anterior border (Fig. 91 F); light submedian vittae, with lateral
illumination, appears as shadow stripes, as in S. (Cerqueirellum) species; with posterior
illumination, scutum has light grayish subquadrate area posteriorly; scutellum and metano-
tum brownish gray; pleura light brown-gray; legs brown-yellowish with blackish apex of
femora, tibiae and tarsomeres; abdomen light grayish with 5 rows of blackish subquadrate
spots on tergites from II-VII as in S. escomeli; cibarium, basal sector of R, claw and paraproct
as in S. trivittatum, the latter with distal projection straight; eighth sternite with 24-28 setae.
Male: Wing length 2.5-2.8 mm; scutum black with 1+1 anterior subtriangular spots,
continued as thin stripes near posterior border (Fig. 93 F); hind basitarsus wide, ratio of
length/width = 3.8; gonostylus without lateral projection and slight inner curvature (Fig.
93 Q); ventral plate with slight lateral constrictions (Fig. 94 L).
Pupa: Cocoon slightly projected basally, with reinforced anterior border (Fig. 95 M),
length basally 3.6-4.0 mm, dorsally 3.0-3.5 mm, gills 4.3-4.6 mm; frontoclypeus and thorax
with moderate number of tubercles; trichomes with 1-2 branches; gills with 4 branches,
longer than cocoon length, dorsal thicker than ventral branches (Fig. 96 M).
Larva: Maximum length 6.5-7.0 mm; cephalic apotome slightly darkened on basal half,
with median light window (Fig. 97 G); ratio of proximal, median and distal antennal
segments = 1:1.4:1.5; cephalic fan with about 42 rays; postgenal cleft short, dome shaped
(Fig. 97 L); postgenal bridge 1.7 longer than hypostomium length; lateral sclerite of thoracic
proleg with 15-17 teeth arranged in 9-10 groups; anal sclerite with scales and hair; anal ring
with about 89 rows, with 15-17 hooks each; rectal papillae with 8 diverticula on each lobe.
Distribution: Mexico: Distritop Federal, Mexico, Michoacan, Morelos, Michoacán,
Veracruz, Zempoala.
Bionomics: Unknown.
Discussion: Simulium mangabeirai is very similar, as adults, to S. trivittatum, S.
veracruzanum, S. bobpetersoni and S. gonzalezherrejoni, but can be differentiated by its pupal gill
of 4 branches. Also the ornamentation of the scutum in both sexes shows differences
with the aforementioned species.

Simulium (Psilopelmia) longithallum DÍAZ NÁJERA & VULCANO (Fig. 96 O.)


Simulium (Psilopelmia) longithallum DÍAZ NÁJERA & VULCANO, 1961: 221; COSCARÓN, 1987: 31;
COSCARÓN et al., 1996: 93-94; CROSSKEY & HOWARD, 1997: 59; 2004: 53; MOULTON, 1998: 61;
ADLER et al., 2004: 351-352.
Female: Wing length 2.0-2.8 mm; scutum orange brown to brown-blackish with
yellowish pile, 1+1 silvery-blue pruinose submedian vittae and lateral border; scutellum light
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 459

brown-yellowish; pleural submedian vittae with yellowish brown pruinosity; legs yellowish with
apex of femora, tibiae and tarsomeres dark brown; abdomen yellowish with dorsal dark spots
arranged in median row on segments II-VII and 1+1 lateral rows on V-VIII; cibarium with
median depression well sclerotized and laterally with rim covered by small teeth arranged as in
S. trivittatum; basal sector of R without hair, and claws with subbasal tooth; eighth sternite with
25-28 setae; paraproct elongated and thin distally, similar to that of S. gonzalezherrejoni.
Male: Wing length 2.0-2.6 mm; scutum velvety black with 1+1 silvery blue
submedian vittae reaching posterior border, as in S. gonzalezherrejoni; scutellum and
metanotum black; abdomen velvety black with golden pile; gonostylus half of gonocoxite
length, with large posterolateral prominence, as in S. downsi; ventral plate wider than long,
similar to that of S. downsi and S. gonzalezherrejoni.
Pupa: Cocoon with small anterior dorsal and ventral projection, and reinforced
anterior border; length basally 2.6-3.2 mm, dorsally 2.5 mm, gills 2.0-2.2 mm; fronto-
clypeus and thorax with tubercles of irregular size and in moderate number; trichomes
with 2-3 branches; gill of 8 branches with primary, dorsal secondary and tertiary thicker
than other branches (Fig. 96 O).
Larva: Maximum length 5.0-5.8 mm; cephalic apotome darkened in median and
basal areas, with light median longitudinal and 1+1 subbasal windows, very similar to that
of S. escomeli; mandible with latero-mandibular process; antenna darker than stalk of
cephalic fan; ratio of proximal, medial and distal segments = 1:1.2-1.3:1.8-2; cephalic fan
with about 45 rays; postgenal cleft dome shaped; postgenal bridge slightly longer than
hypostomium; lateral sclerite of thoracic proleg with about 22 teeth arranged in 5-7
groups; anal sclerite without scales; anal ring with about 68 rows, with 11-14 hooks each;
rectal papillae with 1 diverticlum on each lobe.
Distribution: Mexico: Jalisco, Morelos; USA: Arizona.
Bionomics: Simulium longithallum breeds in small rivers with moderate to fast flow
and sandy and rocky bottoms. Larvae are found on trailing vegetation, sticks and stones.
Female feeding habits are unknown.
Discussion: Simulium longithallum is closest to S. gonzalezherrejoni, as explained for that
species. Simulium bobpetersoni is very similar in the thinner trunk and basal branches of the
pupal gill. Simulium downsi is similar in the cibarium and genitalia of both sexes, but the
adults are yellowish.

Simulium (Psilopelmia) gonzalezherrejoni DÍAZ NÁJERA (Figs. 92 M; 93 G; 94 M; 96 N.)


Simulium (Psilopelmia) gonzalezherrejoni DÍAZ NÁJERA, 1969: 22-28; COSCARÓN, 1987: 31; COSCARÓN et
al., 1996: 89-93; CROSSKEY & HOWARD, 1997: 59; 2004: 53.
Simulium (Psilopelmia) jerezense DÍAZ NÁJERA, 1969: 28-32.
Simulium (Psilopelmia) netteli DÍAZ NÁJERA, 1969: 32-35.
Simulium (Psilopelmia) letrasense DÍAZ NÁJERA, 1969: 35-36.
Simulium (Psilopelmia) olympicus DÍAZ NÁJERA, 1969: 36-38.
Simulium (Psilopelmia) vulcanoae DÍAZ NÁJERA, 1969: 38-40.
S. gonzalezherrejoni was redescribed by COSCARÓN et al., 1996 who established the
synonyms. It shows very little difference from S. longithallum; the paraproct (Fig. 92 M),
460 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

scutum ornamentation (Fig. 93 G) and ventral plate shape (Fig. 94 M) are similar. The
biggest differences are in the more elongated primary gill branches (Fig. 96 N). It is
necessary to obtain more material and, if possible, to use cytological and molecular
methods to determine the status of these species.
Distribution: Mexico: Aguascalientes, Zaacatecas.
Bionomics: Simulium gonzalezherrejoni breeds in moderate- to fast-flowing creeks
attached to trailing vegetation. Female feeding habits are undescribed.

Simulium (Psilopelmia) panamense FAIRCHILD (Figs. 91 N, V; 92 F; 94 E; 95 V, W; 96 D; 97 C, O.)


Simulium (Psilopelmia) panamense FAIRCHILD, 1940: 705.
Simulium (Ectemnaspis) panamense: COSCARÓN, 1984: 314; 1987: 29.
Simulium (Psilopelmia) panamense: COSCARÓN et al., 1995: 2297-298; COSCARÓN et al., 1996: 64-66;
CROSSKEY & HOWARD, 1997: 59; 2004: 53.
Female: Wing length 2.4 mm; scutum yellow-orange with 1+1 submedian silvery
pruinose vittae, reaching scutellum base; lateral rim silvery; scutum hair brown-yellowish
to blackish yellow, according to light position; scutellum yellow-lemon and metanotum
yellow-brownish; legs yellowish, darkened on apex of femur, tibia and tarsomeres;
abdomen without longitudinal rows of spots on tergites, as is typical in subgenus; tergites
I-II brown to brown yellowish, the following dark brown, with light spot on tergite III;
cibarium with median depression few evident and abundant teeth at sides until cornuae
border (Fig. 91 N); basal sector of R with 1-2 rows of hair; claw with subbasal tooth (Fig.
91 V); eighth sternite with about 22 setae on each side; gonapophyses divergent,
subovoidal with slightly sclerotized rim; paraprocts becoming slightly thin toward apex,
and with well-developed basal process (Fig. 92 F).
Male: Scutum yellow with 1+1 silvery anterior submedian cuneiform spots,
scutellum, metanotum and abdominal tergites I-II yellowish, following abdominal tergites
grayish brown; ratio of length/width of hind basitarsus = 4.4; gonostylus about as long as
gonocoxite, with strong distal curvature, bigger than that of S. escomeli; ventral plate with
large median process with parallel sides (Fig. 94 E).
Pupa: Cocoon compact with reinforced anterior border, length basally 2.7-3.3 mm,
dorsally 3.0-3.5 mm, gills 1.4-2.1 mm; frontoclypeus and thorax with wrinkled border (Fig.
95 W) numerous tubercles (Fig. 95 V); cephalic trichomes with 2-3 branches, most of
them with curled apex, thoracic with 3-4 branches; gill with 20 branches, with branching
close to base (Fig. 96 D).
Larva: Maximum length 4.5-5.0 mm; cephalic apotome with median basal area
brownish, with light median window (Fig. 97 C); antenna surpassing apex of cephalic fan
stem; ratio of proximal, medial and distal segments = 1:1.4:1.4; shape as in figure 97 O;
cephalic fan with 40-45 rays; mandible with 1 latero-mandibular process; postgenal cleft
dome shaped; postgenal bridge about as long as hypostomium length; lateral sclerite of
thoracic proleg with 26 teeth arranged in 9-10 groups; anal sclerite with scales and hair; anal
ring with about 89 rows, with 14-16 hooks each; rectal papillae with 6 diverticula on each
lobe, with 1 of these diverticula about 2 times thicker and slightly longer than the others.
Distribution: Costa Rica; Panama: Coclé, Panama.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 461

Bionomics: Unknown.
Discussion: Simulium panamense has similarites with other species with a banded
yellow scutum, such as S. samboni and S. callidum, but the unique subtriangular and
elongated paraproct, ventral plate with great median distal process, and pupal gill with 20
branches permit differentiation. Simulium panamense has characters very similar to those of
S. (Ectemnaspis) species, such as in the cibarium, shape of the paraproct and ventral plate,
presence of hair on the base of R and a subbasal tooth on the claw, creating some
confusion at the supraspecific level.

Simulium (Psilopelmia) callidum (DYAR & SHANNON)


(Figs. 91 D, W; 92 G; 93 S; 94 F; 96 E; 97 D.)
Eusimulium callidum DYAR & SHANNON, 1927: 16; LANEA & VULCANO, 1943: 437.
Simulium (Lanea) callidum: DALMAT, 1955: 126-130.
Simulium (Ectemnaspis) callidum: COSCARÓN, 1984: 315-316; COSCARÓN et al., 1995: 295-296.
Simulium (Psilopelmia) callidum: VARGAS & DÍAZ NÁJERA, 1957b: 183; COSCARÓN, 1987: 30; COSCARÓN
et al., 1996: 67-70; CROSSKEY & HOWARD, 1997: 58; 2004: 52.
Eusimulium mooseri DAMPF, 1927: 127.
Female: Wing length 2.7-2.8 mm; scutum and scutellum yellowish, scutum with 1+1
silvery submedian thin vittae extended 2/3 to posterior margin (Fig. 91 D); metanotum
brown-yellowish; pleurae yellow, darkened ventrally; legs yellowish; abdomen without
rows of dark spots typical of subgenus, tergites of segments I-II and III yellowish,
following segments dark brown, IV-V velvety, VI-IX shiny; cibarium with small median
depression and lateral area slightly pronounced with small spicules, without teeth and
spines, as in S. pseudocallidum; basal sector of R with 2 rows of hair; claw with subbasal
tooth (Fig. 91 W); paraprocts subtriangular, becoming thin toward apex (Fig. 92 G).
Male: Wing length 2.4-2.8; scutum yellow with 1+1 silvery anterior submedian
cuneiform spots extending 1/3 of scutum length; metanotum and abdomen yellowish
with abdominal segments III-IX grayish brown; ratio of length/width of hind basitarsus
= 5.5; gonostylus relatively thick, about 3/4 of gonocoxite length (Fig. 93 S); ventral plate
subovoidal, without lateral constrictions (Fig. 94 F).
Pupa: Cocoon compact with threads evident, anterior border slightly reinforced
with median basal projection; length basally 3.0-3.5 mm, dorsally 2.7-3.3 mm, gills 1.4-2.1
mm; frontoclypeus and thorax with abundant tubercles with smooth border; cephalic and
thoracic trichomes with 2-3 long branches; gill with 8 branches, arising close to base and
very open; first dorsal branch slightly thicker than others (Fig. 96 E).
Larva: Maximum length 5.5-6.0 mm; cephalic apotome slightly darkened medioba-
sally (Fig. 97 D); antenna surpassing apex of cephalic fan stem; ratio of proximal, medial
and distal segments = 1:1:1.5-1.6; cephalic fan with 30-32 rays; postgenal bridge as long as
hypostomium; lateral sclerite of thoracic proleg with about 50 teeth, arranged in 13-14
groups; anal sclerite with single hairs and abundant scales with numerous denticles similar
to those of S. pseudocallidum; anal ring with about 105 rows, with 14-17 hooks each.
Distribution: Belice; Colombia: Valle; Guatemala; Mexico: Cordoba, Morelos,
Oaxaca, Veracruz.
462 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Bionomics: Simulium callidum breeds in slow-flowing streams. The larvae are found
on fallen leaves and subaquatic vegetation. Females bite domestic mammals, birds and
humans.
Discussion: Simulium callidum has similarities with the yellowish species. Some
characters distinguish it, such as the shorter and cuneiform submedian silvery stripes,
cibarium with thick rim lacking teeth, paraproct subtriangular and ventral plate subovoi-
dal. The presence of hair on the basal sector of R and a subbasal tooth on the claw, the
presence of scales on the anal sclerite with several spiculae on the border, and the elevated
number of hook rows on the anal ring also differentiate it. As for S. panamense, there are
several characters shared with S. (Ectemnaspis) species, reinforcing the idea that there is a
close relationship between S. (Ectemnaspis) and S. (Psilopelmia).

Simulium (Psilopelmia) pseudocallidum DÍAZ NÁJERA (Figs. 91 O; 96 F; 97 R.)


Simulium (Psilopelmia) pseudocallidum DÍAZ NÁJERA, 1965: 25-45; COSCARÓN, 1984: 316; 1987: 31;
COSCARÓN et al., 1996: 70-73; CROSSKEY & HOWARD, 1997: 59; 2004: 53.
This species is very close to S. callidum and here we mention only the differences.
Female: Wing length 3.1-3.2 mm; metanotum black; abdomen with tergites I-V
yellowish; cibarium generally without spicules on lateral prominences (Fig. 91 O); basal
sector of R without hair.
Male: Wing length 3.2 mm; ratio of length/width of hind basitarsus = 3.7.
Pupa: Thoracic trichomes with 3-5 branches (Fig. 96 F).
Larva: Cephalic fan with 40-42 rays; lateral sclerite of thoracic proleg with 35-38
teeth arranged in 12-14 groups; anal sclerite with scales and abundant hair (Fig. 97 R).
Distribution: Mexico: Mexico, Oaxaca.
Bionomics: Simulium pseudocallidum breeds in cold, fast-flowing creeks and is
sympatric with S. veracruzanum and S. himmani. Female feeding habits are unknown.
Discussion: The small differences could make S. pseudocallidum a variation of S. callidum.
It is necessary to obtain more material in order to see how large the variation is among
populations and also to do cytological and molecular studies to confirm its true identity.

Simulium (Psilopelmia) travisi VARGAS, VARGAS L. & RAMÍREZ PÉREZ


Simulium (Psilopelmia) travisi VARGAS et al., 1993: 175; CROSSKEY & HOWARD, 1997: 59; 2004: 53.
According to its description, S. travisi is very close to S. callidum and S. pseudocallidum;
here we mention only the differences.
Female: Cibarium without spicules on sublateral prominences.
Male: Ventral plate with body laterally and basally slightly concave, contrary to
convex shape in S. callidum and S. pseudocallidum.
Pupa: Dorsal branch thicker and nearly 5 times longer than other filaments, which in
the most similar species is only about 1.5 times longer.
Larva: Ratio of proximal, medial and distal antennal segments= 1: 1.5: 2; cephalic
fan with 40-43 rays.
Distribution: Costa Rica: Alajuela, Heredia.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 463

Bionomics: Simulium travisi breeds in fast, turbulent streams at altitudes of 800-


1325 m, attaching to rocks and aquatic vegetation. No information is available on female
feeding habits.
Discussion: Simulium travisi is closest to S. callidum and S. pseudocallidum. Differenc-
es in the male ventral plate and length of the first dorsal pupal gill filament allow them
to be separated.

BLANCASI species group


The S. (BLANCASI) species group is unique, characterized mainly by the uniformly
gray leaden scutum in adults of both sexes, lack of denticles on the cibarium, simple claw
of the female, elongated subcylindrical gonostylus and subtriangular and elongated
ventral plate of the male, four long branches of the pupal gill and single rectal papillae of
larvae. This combination of characters makes it very difficult to include the species in any
known Simulium subgenus; the closest are S. (Ectemnaspis) and S. (Psilopelmia), two taxa with
a tight relationship (COSCARÓN & MIRANDA ESQUIVELL, unpubl.). However, the similarities
in the male and female grayish coloration and genitalia (gonostylus and ventral plate in
male and paraproct in female) show a close relationship with S. (Psilopelmia), with great
variation in coloration and morphology. Probably it is a desert lineage, with characters that
distinguish it as a separate species group.
Adult: Medium-sized (wing length, 2.2-3.2 mm); color of scutum in both sexes
uniformly gray, except small area near anterior border; abdomen with conspicuous dark
and light pattern; eighth and ninth tergites of female polished; female with fronto-
ocular triangle well developed; cibarium glabrous between cornuae; mandibles and
maxillae serrated on both edges; setae of scutum of uniform type, irregularly arranged,
not forming lines or groups; pleural membrane and katepisternum bare; basal section
of R bare in both sexes; R1 with setae and spines; fore tarsi narrow, approximately 8
times as long as wide; calcipala and pedisulcus present; hair on legs simple, not scale
like; male genitalia with gonostylus about as long as gonocoxite, elongate, subconical
distally, without projection at base, and with single apical spinule; ventral plate
subtriangular, membranous, lacking median keel; median sclerite about as long as wide,
with apical incision; endoparameres with several large hooks; gonapophysis small,
glabrous; paraproct short, simple; spermatheca globular, not reticulate, its surface
minutely punctate; internal spicules present.
Pupa: Cocoon inverted slipper-shaped, lateral margins of anterior opening receding
from dorsal to ventral surface; lateral apertures absent; gill with 4 long, slender filaments
arising from 2 short bases or from common trunk; trichomes of head and thorax simple,
very short; tergites II-V with 4+4 simple hooks, VII and VIII with spine combs; stemites
V-VII each with 2+2 simple, bifid or trifid hooks; apex of abdomen lacking specialized
setae, and with 1+1 apical tubercles.
Larva: Abdomen gradually widening posteriorly, apex not truncate; central tubercles
absent; head and cephalic fans normal; head pattern positive; antennae as long as stem of
cephalic fan; medial antennal segment with 1 or 2 constrictions; apical tooth of mandible
much larger than outer teeth; presence of 1 or 2 well-developed latero-mandibular
464 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

processes; pre-apical teeth decreasing in size from first to third; 2 marginal teeth well
developed; hypostomium with median and lateral teeth about equally prominent,
intermediate teeth much smaller; hypostomial setae arranged in 1+1 single rows, these
rows slightly diverging posteriorly from lateral margins of hypostomium; postgenal cleft
either large or almost absent; cervical sclerite very small, isolated in neck membrane;
lateral sclerite of proleg well developed, elongate subpentagonal; setae arranged in 1 row,
clustered in groups in most cases formed of 3 or 4 long and short setae: overall size of
setae increasing from upper to lower border of sclerite; cuticle of body bare; perianal
setae present; rectal papillae composed of 3 simple lobes.
Discussion: This group is formed of three closely knit species: S. blancasi
WYGODZINSKY & COSCARÓN, S. tenuipes KNAB, and S. penai WYGODZINSKY & COSCARÓN.

Simulium (Psilopelmia) blancasi WYGODZINSKY & COSCARÓN (Fig. 98.)


Simulium blancasi WYGODZINSKY & COSCARÓN, 1970: 6-14; COSCARÓN, 1987: 55; 1991: 143.
Female: Wing length, 2.7-2.8 mm; frons and clypeus dull gray, with few scattered
yellowish hairs; mouthparts piceous; maxillary palps grayish; antennae black, grayish
pruinose; first, second, and base of third segment light orange; scutum dull gray;
anterolateral angles and 1+1 anterior submedian spots silver-white; pleura and sterna with
grayish pruinosity, faintly nacreous; pleural membrane dark brown; hair of scutum short,
scattered, yellowish; scutellum yellowish brown, narrowly gray pruinose along margins;
hair of scutellum from yellowish to black; metanotum glabrous, black, heavily gray
pruinose; general color of legs yellowish white, dark pigment from piceous to black;
pattern as in figures K-M; fore coxae light colored, mid and hind coxae dark; setae of legs
colored with respect to background; anterior wing veins light grayish brown; setae at base
of wing black; stem of halteres yellowish, knob silver-white; color pattern of abdomen as
in figure A; first abdominal segment yellowish white, its fringe silver-white; second tergite
blackish at center, sides with 1+1 very large silvery spots; third to fifth tergites velvety
black, their center at anterior half with transverse dumbbell-shaped light-colored spot,
pinkish or yellowish on third and fourth, silver-white on fifth segment that is larger than 2
before; these spots imperceptible in some specimens; sixth and seventh tergites silver-
white, dark only at extreme sides and in some cases also along posterior margin behind
large silvery area, the latter occasionally indentate posteriorly at middle; eighth and ninth
tergites piceous or black, not conspicuously lighter than anterior tergites; undersurface of
abdomen grayish; fronto-ocular triangle (Fig. B) small, wider than deep; antennae 11
segmented, shape and proportions of segments as in figure C; frons as in figure D;
cibarium as in figure E; maxillary palp as in figure F; apical segment as long as 2 preceding
combined; third segment rather strongly widened; sensory vesicle (Fig. G) large, its
diameter slightly larger than half width of third segment, its mouth wide; tubercles
numerous; lacinia and mandibles serrate on both edges; lacinia (Fig. H) with 20-23 teeth,
mandibles (Fig. I) with 33-35 teeth; scutum lacking scales, setae randomly distributed, not
forming rows or groups; microtrichiae relatively long, of felt-like appearance. Sc and basal
section of R glabrous. R1 with setae and spines arranged in 1 row. Rs with setae arranged
in 1 row beginning slightly beyond base of vein (Fig. J); shape of legs and proportions of
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 465

C D K
B
E
L O
I
N
H
G
Q
M
A F

J
P
W
T
R V
S

U c

Z
a

X b

d e
Y

f
h g

Figure 98. S. (Psilopelmia) blancasi. A-O: Female; A: abdomen, dorsal view; B: fronto-ocular triangle; C:
antenna; D: frons; E: basal portion of cibarium; F: maxillary palp; G: sensory vesicle of maxillary palp;
H: lacinia; I: mandible; J: wing anterior veins; K-M: fore, mid and hind leg, respectively; N: calcipala
and tarsomere I; O: claw. P-W: Male; P: abdomen, lateral view; Q: sensory vesicle of maxillary palp; R-
T: fore, median and hind leg, respectively; U: calcipala and tarsomere I; V: gonocoxite and gonostylus;
W: ventral plate. X-Z, a-c: Pupa; X: dorsal and lateral views; Y: gill; Z: frontoclypeus; a: cephalic
trichomes enlarged; b: portion of thorax showing, at right, trichomes and tubercles with high
magnification; c: abdominal chaetotaxy. d-g: Larva; d: cephalic apotome; e: antenna; f: mandibular
teeth with 2 latero- mandibular processes; g: postgenal cleft and hypostomium.
466 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

segments as in figures K-M; calcipala well developed, but not quite attaining level of
pedisulcus (Fig. N); claws as in figure O; eighth stemite with about 25+25 setae;
gonapophysis glabrous, inner border faintly sclerotized; setae and microtrichiae absent;
cercus much wider than long, apex rounded; apex of paraproct subrectangular.
Male: Wing length, 2.5-2.6 mm; frons and clypeus dull silver-gray; antennae black, faintly
grayish pruinose; first, second, and basal third of third segments orange; eyes purple; scutum
dull gray, faintly tinged with olivaceous; 1+1 submedian spots at anterior margin of
mesonotum faintly silvery white; pleura and sterna gray; scutellum, metanotum and setae of
thorax as in female; general color of legs yellowish white, dark pigment from piceous to
blackish; pattern as in figures R-T; fore coxae light colored, mid and hind coxae dark; white
reflections on leg segments absent; color of wings and halteres as in female; color of abdomen
as in female, but all spots silver, and tergites VI and VII almost completely white (Fig. P); last
segment of maxillary palp slightly longer than 2 preceding combined; sensory vesicle globular,
its diameter equal to half width of third segment; structure of vesicle as in figure Q; chaetotaxy
of thorax as in female; wing venation and chaetotaxy as in female; shape of legs and
proportions of segments as in figures R-T, hind basitarsus 4.6 times as long as wide; calcipala
as in figure U; setae of legs somewhat longer and more numerous than in female; gonocoxite
wider than long; gonostylus approximately as long as gonocoxite, elongate conical, slightly over
twice as long as wide, with 1 apical spine (Fig. V); ventral plate (Fig. W) subtriangular,
membranous, only its short basal arms faintly sclerotized; keel absent; median sclerite broad, as
long as wide, its apical incision occupying approximately 1/3 length of sclerite.
Pupa: Cocoon (Fig. X) subconical, slipper-shaped, dorsal wall salient at middle,
projecting somewhat beyond level of ventral border of anterior opening; ventral wall of
cocoon delicate but complete; cocoon translucent, shining, threads perceptible but not
coarse; color of cocoon light brown; length of cocoon along dorsal surface 3.0-3.5 mm,
along ventral surface, 2.5 mm; length of pupa 2.8-3.0 mm, of gill up to 4.5 mm, viz., up to
1.5 times as long as body of pupa; gills composed of 4 forwardly directed slender filaments
(Fig. Y); 2 primary branches arising from short common trunk; head and thorax of pupa
with numerous small platelets, arranged in irregular groups as shown in figure Z; trichomes
small, hair like, not branched (Fig. a); head with 2+2 frontal and 1+1 facial trichomes,
occasionally only 1 frontal trichome on 1 side; dorsal surface of thorax on each side with 5-
6 simple, very short, hair-like trichomes (Fig. b); abdomen as in Figure c; tergite I with few
small setae; tergite II with 3+3 small, slender, simple spines accompanied laterally by
characteristic group of 2 small spines and 1 long hair; tergites III and IV each with 4+4
simple hooks and few very short hairs; tergite V glabrous; anterior border of tergites VII and
VIII with transverse row of 20-22 spiniform tubercles, these rows connecting laterally with
areas of very small scalelike cuticular processes, the latter areas also present on tergites VI
and IX; apex of abdomen with 1+1 distinct, slightly sclerotized tubercles; sternite IV with
1+1 groups composed of 2-4 simple short hairs; sternites V-VII with 2+2 hooks, closely
approximated on V, more distant on VI and VII; most hooks bifid, occasionally trifid, outer
ones frequently simple, especially on tergites VI and VII; sternites IV-VIII each with 1+1
groups of minute scalelike cuticular processes.
Larva: Length of mature larva 6.0-6.5 mm; width of head capsule 0.7 mm; general
body color whitish, greenish pigment dorsally and ventrally on anterior half of body; head
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 467

yellowish white; pattern of cephalic apotome as in figure d; body integument glabrous,


few short setae in caudal area; ventral tubercles absent; antennae as in figure e, faintly but
distinctly pigmented throughout, medial segment lightest in color; medial segment with
very deep constriction accompanied by unpigmented area; ratio of proximal, medial and
distal segments = 1:1.1:1.25; cephalic fan with 40-45 rays; toothing of mandible as in
figure f; hypostomium with anterior border heavily pigmented; median and lateral teeth
prominent, intermediate teeth much smaller; sublateral teeth and lateral serrations very
well developed; hypostomial setae arranged in single row, their number approximately 5 in
each group; postgenal cleft deep, about as deep as postgenal bridge (Fig. g); anal sclerite as
in S. tenuipes; anal ring with approximately 80 rows each composed of about 17 hooks;
rectal papillae simple, composed of 3 lobes (Fig. h).
Distribution: Chile: Tarapaca; Peru: Lima.
Bionomics: Larval stages of S. blancasi live in small creeks about 1 m wide and 0.30
m deep, in crystalline waters, flowing over aquatic vegetation or stones, with a temperature
of about 15 ºC.
Discussion: The abdominal ornamentation of the female without transverse
anterior light grayish spots on tergites III-V and the pupal gills without primary branches
permit differentiation from S. tenuipes and S. penai, the closest species. Males of S. tenuipes
also have a shorter ventral plate, without basal lateral constrictions.

Simulium (Psilopelmia) tenuipes KNAB (Fig. 99.)


Simulium tenuipes KNAB, 1914a: 21; SILVA FIGUEROA, 1917: 30; WYGODZINSKY & COSCARÓN, 1970: 14-
17; COSCARÓN, 1987: 19; 1991: 144-145.
Female: Wing length, 2.2-2.9 mm; color of head and thorax and their appendages as
in S. blancasi. Abdomen (Fig. A) with tergites I and II as in S. blancasi, III-V dark velvety
brown, VI and VII very light brown with 1+1 silvery white spots on VI, VIII and IX
piceous Undersurface of abdomen grayish; fronto-ocular triangle, antennae, palpi, lacinia,
sensory vesicle, mandible, and cibarium as in S. blancasi; basal portion of cibarium smooth
(Fig. B); claws without subbasal tooth (Fig. C); wings and legs as in S. blancasi; calcipala
about as long as wide (Fig. D); eighth sternite lightly pigmented, and with about 40
trichomes on each side (Fig. E); gonapophysis subtriangular and with abundant microtri-
chiae; genital fork with stem and lateral branches well sclerotized; spermatheca subovoidal
(Fig. F); paraprocts subtriangular, slightly wider than long (Fig. G).
Male: Wing length, 2.4-2.7 mm; color of head and thorax and their appendages
very close to that of S. blancasi, but because of greater number of hairs on scutum, their
golden color is more apparent; legs with tibiae not distinctly darkened at base; abdomen
with color pattern very similar to that of female, but tergites VI and VII with 1+1 small
sublateral silvery spots; shape of antennae, maxillary palp, and sensory vesicle as in S.
blancasi; shape and proportions of segments of legs very similar to those of S. blancasi,
but hind legs somewhat wider, hind basitarsus 3.7 times as long as wide; gonostylus
elongated, subcylinderical, nearly as long as gonocoxite (Figs. H, I), ventral plate
subtriangular (Fig. J); endoparameres with longer spines (Fig. I); median sclerite with
median area translucent (Fig. I).
468 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

B C

A
F
H
D

E
J

N M
O

Figure 99. S. (Psilopelmia) tenuipes. A-G: Female; A: abdomen, dorsal view; B: basal portion of
cibarium; C: claw; D: calcipala and tarsomere I; E: eighth sternite, gonapophysis and genital fork; F:
spermatheca; G: cercus and paraproct: right in posterior view, left mounted on slide. H-J: Male; H:
gonocoxite and gonostylus, dorsal view; I: genitalia dorsal view showing gonocoxite and
gonostylus, ventral plate, endoparameres and median sclerite; J: ventral plate. K-P: Pupa; K: lateral
view; L: portion of thorax and gill; M-P: larva; M: postgenal cleft and hypostomium; N: lateral
sclerite of thoracic proleg; O: anal sclerite; P: scales of anal sclerite area.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 469

Pupa: Color light brown, shape of cocoon as in figure K; length of cocoon along
dorsal surface 2.3-2.9 mm, along ventral surface 1.8-2.0 mm; length of pupa proper 2.5-
2.9 mm, of gills 2.1-2.5 mm, viz., not longer than body of pupa; gill composed of 4
forwardly and downwardly bent filaments (Figs. J, K) arising from short common trunk;
trichomes and platelets of head as in S. blancasi; thorax with single trichomes (Fig. L);
abdominal onchotaxy as in S. blancasi.
Larva: Length of mature larva 6.0-6.8 mm; width of head capsule 0.65-0.70 mm;
general body color whitish, with greenish pigment dorsally and on ventral surface of
anterior half of body; antennae much as in S. blancasi; ratio of proximal, medial and distal
segments = 1:1.1:1.2; cephalic fan with approximately 35 rays; primary teeth slender,
spaced closely, secondary teeth more than half as long as primary ones; mandible much as
in S. blancasi, with 2 or occasionally 3 serrations; hypostomium with 4 or 7 hypostomial
setae in each row (Fig. M); postgenal cleft extremely reduced, almost imperceptible; lateral
sclerites of thoraacic proleg as in figure N; anal sclerite with hair and small spicules (Figs.
O, P); anal ring with about 80 rows each composed of 15-18 hooks.
Distribution: Chile: Tarapacá; Peru: Tacna.
Bionomics: Larvae and pupae were collected in ditches from leaves of Scirpus
(Cyperaceae) and leaves of diverse aquatic plants, on branches of trees and on rocks. The
water was slightly turbid, and the temperature was 16-18 °C. Simulium tenuipes attacks
humans, mules, and horses.
Discussion: This species can be most easily separated from S. blancasi in the larva, by its
obsolete gular cleft, and in the pupa, by the secondary respiratory filaments that seemingly arise
from a common trunk. The females and males of S. tenuipes have different abdominal
ornamentation from S. blancasi and a distinct ventral plate shape, as indicated for that species.

Simulium (Psilopelmia) penai WYGODZINSKY & COSCARÓN (Fig. 100.)


Simulium penai WYGODZINSKY & COSCARÓN, 1970: 18; COSCARÓN, 1987: 19-20; 1991: 145.
Female: Wing length 2.8-3.2 mm; color of head and thorax and their appendages as
in S. blancasi, except scutum and metanotum which are darker, contrasting more with pale
yellow scutellum; pattern of abdomen similar to that of S. tenuipes but with posterior
border of tergites III-V light colored, light-colored portion on V occupying entire
posterior half; general structure of fronto-ocular triangle, antennae, maxillary palp with
sensory vesicle, and cibarium as in S. blancasi; lacinia with approximately 20 teeth,
mandible with 35 teeth; shape and pigmentation of segments of legs much as in S.
blancasi, but calcipala much reduced, subtriangular; wings as in S. blancasi.
Male: Wing length, 2.8 mm; color of head and thorax with their appendages as in S.
blancasi, but scutum and metanotum darker; color pattern of abdomen unknown;
structure of head and appendages and of wings as in S. blancasi; hind basitarsus 4.0-4.5
times as long as wide; calcipala (Fig. A) similar to that of female; genitalia similar to those
of S. blancasi, but gonostylus (Fig. B) slightly wider.
Pupa: Color and shape of cocoon much as in S. blancasi, but more closely woven
and, when observed in liquid, somewhat parchment-like, viz., with lead like sheen; length
of cocoon along dorsal surface 2.8-3.5 mm., along ventral surface 2.6-3.0 mm; length of
470 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

A B

E
D
G F

Figure 100. S. (Psilopelmia) penai. A-B: Male; A: obsolescent calcipala and pedisulcus; B: gonostylus.
C-E: Pupa; C: dorsal portion of thorax and gill; D: frontoclypeus; E: trichomes and tubercles of
thorax with high magnification; G-F: larva; F: mandibular marginal teeth; G: cephalic capsule,
ventral view.

body of pupa 3.4-3.7 mm., of gills 3.8-4.0 mm; gill either straight or downwardly curved
(Fig. C); head (Fig. D) and thorax without or with only few scattered platelets (Fig. E);
abdominal onchotaxy as in S. blancasi.
Larva: Length of mature larva, 6.5-7.0 mm.; width of head capsule, 0.65-0.70 mm;
overall color of body whitish; pattern of cephalic apotome as in S. blancasi; antennae
similar to those of S. blancasi; ratios of proximal, medial and distal segments = 1:1.1:1.2;
cephalic fan with 38-42 rays Primary teeth slender, not spaced closely, secondary teeth
somewhat over half as long as primary ones; mandible and hypostomium much as in S.
blancasi; marginal teeth as in figure F; postgenal cleft extremely reduced (Fig. G); anal
sclerite as in S. tenuipes; anal ring with 76-80 rows, each composed of 13-17 hooks.
Distribution: Chile: Antofagasta.
Bionomics: The larvae and pupae were collected on aquatic plants in a small, slightly
brackish stream.
Discussion: Simulium penai seems to be closer to S. tenuipes than to S. blancasi. It
agrees with the former in the structure of the gills and virtual disappearance of the
postgenal cleft, a specialized character in this group. The pupae of S. tenuipes can be
identified by the more abundant platelets and the adults by the bigger calcipala.

Species inquirenda

Simulium jacobsi DALMAT


Simulium (Lanea) jacobsi DALMAT, 1953: 40; 1955: 149.
Simulium (Psilopelmia) jacobsi: CROSSKEY & HOWARD, 1997: 59; 2004: 53.
This species was described based only on the larva that, according to the author, is
similar to S. haematopotum and S. samboni. The characters described do not provide security
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 471

for differentiating it from other species. It will be necessary to obtain more material of other
stages to determine its identity; until then, we maintain S. jacobsi as species inquirenda.

Simulium (Aspathia) ENDERLEIN (Figs. 101-112; 150.)


Aspathia ENDERLEIN, 1935: 359 (as genus).
Type-species: Simulium hunteri MALLOCH, 1914: 59.
Jalacingomyia PY-DANIEL, in PY-DANIEL & MOREIRA SAMPAIO, 1994: 127, 129-137. Type species:
Simulium anduzei VARGAS & DÍAZ NÁJERA, 1948b: 328.
Adults: Female and male blackish. Frontal pruinosity dull blackish to shining black;
scutum blackish with color pattern variable, but generally with 2+2 shining silvery nacreous
longitudinal stripes to 1+1 subquadrate diffuse silvery pruinose anterior spots in females and
usually 1+1 silvery diagonal anterior stripes in males, with some variation, sometimes with
greenish blue iridescence or silvery pruinose areas; female pleural membrane and katepiste-
mum bare, sometimes shining; abdomen blackish without special ornamentation, tergites V-IX
shining; male with typical 1+1 silvery spots on both sides. Wing with basal section of R bare.
Base of cibarium smooth but sometimes with small nodular granulations or with median
projection that can be covered with small denticles at apex. Frons slightly convergent below.
Fronto-ocular triangle deep. Arms of furcastemum with conspicuous projections. Sensory
vesicle of palpus well developed, about 1/3 to 1/2 length of palpomere. Legs generally with
petaloid hair; tarsal claws with strong basal tooth. Female gonapophysis distally truncate to
bluntly rounded; paraprocts about as long as wide at base, slightly acuminate ventrally or
rounded distally as are cerci. Genital fork with lateral branches relatively short but apodemes
elongate and acute distally, spermatheca well developed, with inner spiculae. Male gonocoxite
subquadrate; gonostylus about 1.5 times as long as gonocoxite, subparallel-sided, apex
rounded, with subterminal spine, and internal basal process frequently strong and as long as
transverse diameter of gonostylus basally; ventral plate generally with narrow curved body that
is interspecifically variably shaped, sometimes hairy, and narrower than basal arms; parameres
large, with numerous robust hooks.
Pupa: Cocoon generally slipper shaped, with distinct threads, sometimes with
reinforced edge and projecting dorsally, covering only gill bases. Gill with 5-130
filamentous branches to membranous with 3-24 terminal branches. Frontoclypeus with
2+2 frontal and 1+1 facial trichomes, absent on cephalic, epicranial, genal and ocular
areas. Thorax with 5 multibranched to 25-140 simple or bifurcated trichomes on disc.
Tubercles of frontoclypeus and thorax variable, smooth to rugose, abundant to sparse or
absent. Abdominal onchotaxy with following cuticular processes dorsally on each side of
tergites: I with 1 short median and 1 long sublateral seta; II with 3 median, 3 submedian
(that are more robust, like small hooks in some species), and 3 lateral setae; III and IV
with 4 large hooks near posterior margin and 1 median, 1 submedian and 2 or 3 lateral
setae on each tergite; V with 3 short submedian setae and small scales on anterior edge
present also on following posterior tergites; VI with 2 submedian and 2 lateral setae and 2-
4 small spines on anterior edge as on tergites VII-VIII that have 8-10 and 14-16 spines,
respectively; IX with short and robust distal hook. Ventrally, sternites I-II without setae;
III with 4 submedian and 1 lateral seta; IV with submedian simple or bifurcated hook plus
472 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

2 submedian and 3 lateral setae; V with 2 submedian trifurcated hooks and 2 setae plus 2
lateral setae; VI-VII with submedian bifurcated or trifurcated and 1 sublateral simple to
bi- or trifurcate hooks plus 1 submedian and 1 or 2 sublateral setae. III-VII with small
rugosities like scales on submedian area; VIII with 1 or 2 sublateral setae.
Larva: Cephalic apotome variable, without unusual ornamentation, frequently with
positive spots on median longitudinal area and 1+1 anterior groups with subbasal
position, or with negative spots and gradually darkened basally. Postgenal cleft shallow,
internal edge rounded or subacute, postgenal bridge generally longer than height of
hypostomium. Hypostomial teeth well differentiated, median tooth at same level as lateral
teeth, 3-9 lateral serrations on each side with 4-9 lateral setae. Mandible normal; inner
teeth 8-12, generally in 2 rows; 2 marginal teeth present, the posterior one very small;
lateral mandibular process generally present. Antenna generally longer than stem of
cephalic fan. Antennal segments medial and distal with transverse striations evenly
sclerotized, without light areas and medial segment generally longer than proximal and
distal. Cephalic fan with 29-45 primary rays. Cervical sclerites small and free. Body shape
normal. Thoracic cuticle bare. Sclerite of proleg with 30-36 teeth, generally arranged in
groups. Anal sclerite X-shaped, without struts, generally with simple and sparse hair, rarely
with scales. Anal ring generally with 66-90 rows but may be up to 109, with 12-15 hooks
on each. Rectal papillae with 3 lobes and 0 to 13 branches on each lobule. Body without
hair or scales. Body generally with ventral tubercle on ninth segment.
This subgenus in the Neotropical realm is composed of 25 species, which occur at
elevations of 500-3700 m. Several species are anthropophilic and vectors of the agent of
human onchocerciasis.
The Neotropical species are in the METALLICUM and HUNTERI species groups
(CROSSKEY & HOWARD, 1997, 2004).

KEYS TO NEOTROPICAL SIMULIUM (ASPATHIA) SPECIES

Females
Simulium covagarciai, S. kompi, S. ruizi and S. sandyi are excluded due to incomplete information.

1 Scutum with 1+1 anterior grayish spots not continued as light grayish vittae ..... 2
– Scutum with 3 light nacreous or silvery stripes (Figs. 101 A, F) between 5 blackish
vittae (Figs. 101 B, G) ....................................................................................................... 5
2(1) Scutum with 1+1 whitish subtrapezoidal anterior spots (Fig. 101 H); cibarium
with median basal area smooth, without median process or reinforcement on edge
(Fig. 102 C); paraprocts scarcely produced ventrally, with rounded edge (Fig. 103
D) .....................................................................................................................Simulium putre
– Scutum with 1+1 whitish anterior spots not subtrapezoidal; cibarium with median
process or reinforced on edge; paraprocts produced ventrally and gradually
acuminate at apex (Figs. 103 J, K) .................................................................................. 3
3(2) Scutum with 1+1 anterior subtriangular silvery spots; cibarium without median
process or only slightly elevated medially (Fig. 102 G) ...............................................
............................................................................................. Simulium hechti, Simulium wygoi
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 473

– Scutum with 1+1 anterior subcircular silvery spots or arranged as continuous


anterior band; cibarium with median process (Fig. 102 O) ...................................... 4
4(3) Small species, wing length about 3.0 mm; cibarium with sparse denticles (Fig. 102
O) .................................................................................................................. Simulium bustosi
– Larger species, wing length about 3.8 mm; cibarium without denticles ..................
.................................................................................................................. Simulium costalimai
5(1) Scutum, when viewed with lateral illumination, with pollinose iridescent area;
paraprocts produced ventrally and acuminate at apex (Fig. 103 B) ..........................
.................................................. Simulium metallicum, Simulium horacioi, Simulium jobbinsi,
Simulium morae, Simulium puigi, Simulium racenisi
– Scutum, when viewed with lateral illumination, not iridescent; paraprocts
produced or not produced ventrally .............................................................................. 6
6(5) Relatively large species; wing length 3.3-4.0 mm; cibarium smooth, without
denticles, papillae or median process; sensory vesicle of palp less than 1/2 length
of palpomere ...................................................................................................................... 7
– Medium to small species; wing length less than 3.2 mm; cibarium smooth or not;
sensory vesicle of palp generally 1/2 or more length of palpomere ..................... 9
7(6) Wings with hair on basal section of R ............................................... Simulium anduzei
– Wings without hair on basal section of R .................................................................... 8
8(7) Wing length 4.0 mm; sensory vesicle of palp slightly less than 1/2 length of
palpomere ............................................................................................... Simulium marquezi
– Wing length 3.4 mm; sensory vesicle of palp 1/2 length of palpomere .................
...................................................................................................................... Simulium tricorne
9(6) Cibarium smooth; wing length 2.0-2.5 mm; subbasal tooth of tarsal claw absent or
very small ................................................................................................. Simulium spilmani
– Cibarium with denticles or spherules; wing length 2.7-3.0 mm; subbasal tooth of
tarsal claw well developed (Fig. 102 M) ...................................................................... 10
10(9) Cibarium without median process, with nearly straight edge; paraproct shorter
than wide at base (Fig. 103 G) ...................................................................................... 11
– Cibarium with median process (Figs. 102 J-L); paraprocts about as long as wide
.............................................................................................................................................. 12
11(10) Cibarium with papillae (Fig. 102 N); paraproct not acuminate distally; sensory
vesicle of palp about 1/2 length of palpomere .................................. Simulium parrai
– Cibarium with small acuminate denticles (Fig. 102 F); paraproct gradually
acuminate distally (Fig. 103 G); sensory vesicle of palp less than 1/2 length of
palpomere (Fig. 101 L) ............................................................ Simulium matteabranchium
12(10) Paraproct rounded distally; sternite VIII with 29-35 setae ..................................... 13
– Paraproct acuminate distally; sternite VIII with 20-25 setae .................................. 14
13(12) Inner edges of gonapophyses divergent below .................................... Simulium piperi
– Inner edges of gonapophyses parallel .............................................. Simulium jacumbae
14(12) Inner edges of gonapophyses divergent below .................................. Simulium iriartei
– Inner edges of gonapophyses parallel to gradually divergent below ........................
.......................................................................................................................Simulium hunteri
474 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Males
The following species are not considered due to lack of information: Simulium covagarciai,
S. sandyi, S. racenisi, S. kompi.

1 Ventral plate subquadrate, not laterally flattened (Figs. 106 A-L) ........................... 2
– Ventral plate subconical, subcircular to subcylindrical, gradually flattened from
base (Figs. 106 M-Q) ......................................................................................................... 8
2(1) Gonostylus with inner basal process long and slender; scutum with 1+1 anterior
short subtriangular silvery spots (Fig. 104 I) .......... Simulium jobbinsi, Simulium piperi
– Gonostylus with inner basal process relatively short and stout, frequently wrinkled
and dentate apically; scutum variable; 1+1 short anterior to long convergent
stripes diagonally directed to midline (Figs. 104 A, B) .............................................. 3
3(2) Scutum with 1+1 submedian and 1+1 sublateral silvery spots or bands (Figs. 104
L-N) ...................................................................................................................................... 4
– Scutum with 1+1 silvery spots of variable shape, subtriangular to subquadrate or
subrhomboidal, or stripe-like ........................................................................................... 5
4(3) Median process of ventral plate wide basally (Fig. 106 G) ................. Simulium puigi
– Median process of ventral plate narrow basally (Fig. 106 L) ............ Simulium hechti
5(3) Scutum with 1+1 silvery diagonal spots projecting posteriorly (Figs. 104 A, B, L)
................................................................................................................................................ 6
– Scutum with 1+1 silvery spots not diagonal but nearly transverse (Fig. 104 D)
................................................................................................................................................ 7
6(5) Ventral plate with narrow, short median process (Fig. 106 I); scutum with 1+1
short and basally stout silver spots (Fig. 104 J) ................................... Simulium parrai
– Ventral plate with medium to robust process (Figs. 106 A, B); scutum with 1+1
elongate and distally slender silver spots (Figs. 104 A, B) ..........................................
................................................. Simulium metallicum, Simulium horacioi, Simulium spilmani
7(5) Ventral plate with robust median process edged by cleft, and lateral margins
curved (Figs. 106 D, E); gonostylus inner basal process short and smooth (Fig.
105 F) ........................................................................ Simulium marquezi, Simulium tricorne
– Ventral plate with medium-sized median process not edged by cleft, and lateral
margins straight (Figs. 106 C, H, K); gonostylus inner basal process medially long
and dentate distally (Figs. 105 G, K) ................................................................................
........ Simulium bustosi, Simulium iriartei, Simulium jacumbae, Simulium morae, Simulium wygoi
8(1) Scutum without isolated silver spots or stripes, entire disc silvery margined (Fig.
104 K) ...................................................................... Simulium costalimai, Simulium anduzei
– Scutum with silver spots or stripes ................................................................................ 9
9(8) Scutum with 1+1 submedian anterior subquadrate to subrhomboidal silvery spots
joining silver lateral area; body of ventral plate as wide basally as median
transverse diameter (Fig. 106 M) ...........................................................Simulium hunteri
– Scutum with 1+1 submedian anterior silvery subtriangular to subtrapezoidal spots or
diagonally curved bands (Figs. 104 E, G), not joining silver lateral area; body of ventral
plate narrower basally than median transverse diameter (Figs. 106 O-Q) ................. 10
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 475

10(9) Gonostylus with distinct curvatures in lateral view (Fig. 105 E); body of ventral
plate strongly flattened laterally from base, subcylindrical (Fig. 106 Q) ..................
..........................................................................................................................Simulium putre
– Gonostylus with subparallel straight sides in lateral view; body of ventral plate not
strongly flattened laterally, subconical (Figs. 106 J, P) ............................................. 11
11(10) Gonostylus inner basal process shorter than basal transverse diameter ..................
.......................................................................................................................... Simulium ruizi
– Gonostylus inner basal process longer than basal transverse diameter (Fig. 105 H)
...................................................................................................... Simulium matteabranchium

Pupae
1 Gill stout and membranous (Figs. 107 H, J, 109 A-C, G, I) ..................................... 2
– Gill filamentous (Figs. 107 A, C, F) ............................................................................... 9
2(1) Gill without filaments; frontoclypeus and thorax with tubercles ............................ 3
– Gill with flexible filaments (Figs. 109 A, E, I); frontoclypeus and thorax without
tubercles ............................................................................................................................... 6
3(2) Gill branches digitiform, without small tubuliform secondary branches; cocoon
without anterior projection (Fig. 107 H) ....................................................................... 4
– Gill branches tubuliform with small tubuliform secondary branches (Figs. 109 B,
C); cocoon with (Fig. 107 I) or without anterior projection ..................................... 5
4(3) Gill with 3 branches directed anteriorly, with annulations; cocoon without dorsal
carina ............................................................................................................. Simulium kompi
– Gill with 2 branches directed anteriorly and 1 directed posteriorly, without
annulations (Fig. 107 H); cocoon with dorsal keel ........................... Simulium tricorne
5(3) Gill with 2 globose, basally subcylindrical branches that are progressively
narrower distally and main branch subdivided distally into 7 small and narrower
branches (Fig. 109 B); abdominal tergites I and II with tubercles ............................
................................................................................................................... Simulium marquezi
– Gill arborescent, with 25 relatively narrow terminal branches emerging from 3
short, stout primary branches (Fig. 109 C); abdominal tergites I-II without
tubercles ....................................................................................................... Simulium sandyi
6(2) Gill with main branch subdivided distally (Fig. 107 J) ............................................... 7
– Gill subdivided basally into 4 large branches ............................................................... 8
7(6) Gill distally with main branch and basally with only 1 short branch (Fig. 109 E)
.......................................................................................................................... Simulium ruizi
– Gill with robust stalk expanded distally, with several small and short branches and
basally with 3 short branches (Fig. 109 A) ......................... Simulium matteabranchium
8(6) Gill branches with pseudosegmentation, 2 branches directed anteriorly and 2
ventrally (Fig. 109 I) .................................................................................. Simulium parrai
– Gill branches without pseudosegmentation, the largest directed anteriorly, with
small subtriangular protuberances and 2 ventral subglobose protuberances ..........
................................................................................................................. Simulium covagarciai
9(1) Gill terminal branches greater than 8.5 mm; frontoclypeus and thorax with simple
trichomes (Figs. 108 G, H) ............................................................................................. 10
476 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

– Gill terminal branches less than 3.5 mm; frontoclypeus and thorax with simple or
multibranched trichomes ................................................................................................ 12
10(9) Gill with 3 stout primary branches, subdivided into 7-10 main branches and 85-
100 small terminal branches (Fig. 108 H); thorax with about 50 trichomes on each
side ............................................................................................................... Simulium iriartei
– Gill with about 110-135 branches (Fig. 108 G); thorax with 70-100 trichomes on
each side ............................................................................................................................. 11
11(10) Frontoclypeus and thorax without tubercles; cocoon with dorsal exterior
projection ............................................................................................... Simulium costalimai
– Frontoclypeus and thorax with moderate number of tubercles; cocoon with
straight anterior edge ................................................................................Simulium hunteri
12(9) Gill with 5 branches (Fig. 108 E) ....................................................... Simulium spilmani
– Gill with 6-30 branches ................................................................................................... 13
13(12) Gill with 6 branches about same length (Fig. 108 A); cocoon with straight edge
(Figs. 107 A, C) ................................................................................................................. 14
– Gill with 10-30 branches; cocoon generally with anterodorsal projection or
aperture circle-shaped anteriorly................................................................................... 16
14(13) Frontoclypeal and thoracic trichomes with 2-6 branches. (Figs. 107 B, 108 A),
thorax with 5 trichomes on each side .......................................................................... 15
– Frontoclypeus and thorax with simple trichomes, those of thorax number 60-85
on each side (Figs. 107 C-E) ......................................................................Simulium putre
15(14) Cocoon flattened, subcircular in dorsal view .................................... Simulium horacioi
– Cocoon normal, not flattened (Fig. 107 A) ....................................................................
.... Simulium metallicum, Simulium jobbinsi, Simulium morae, Simulium puigi, Simulium racenisi
16(13) Cocoon flattened, subcircular, with anterior edge fused ventrally (Fig. 107 F); gill
with 10-16 terminal branches ........................................................................................ 17
– Cocoon with normal shape, not flattened, with or without anterodorsal
projection; gill with 10-30 branches ............................................................................. 19
17(16) Gill with 10 terminal branches, with short stem at base and 4 primary branches;
thorax with 25 trichomes on each side, their diameter diminishing gradually
toward apex ................................................................................................ Simulium bustosi
– Gill with 2 or 3 main branches, with 9-11 narrower flexible branches; thorax with
5-120 trichomes that are very stout in their basal 1/3 (Fig. 107 K) ..................... 18
18(17) Gill with 3 main branches; thorax with 100-200 trichomes; thorax smooth (Fig.
108 J) .............................................................................................................. Simulium hechti
– Gill with 2 main branches; thorax with 5 trichomes on each side; thorax wrinkled
(Fig. 108 I) .................................................................................................... Simulium wygoi
19(16) Cocoon with anterodorsal projection; thorax with 25-65 trichomes on each side
.............................................................................................................................................. 20
– Cocoon without anterodorsal projection; thorax with 5+5 trichomes on each side
..................................................................................................................... Simulium anduzei
20(19) Gill with 10 terminal branches; thorax with 50-65 trichomes .......... Simulium piperi
– Gill with 25 terminal branches (Fig. 108 F); thorax with 25-35 trichomes .............
................................................................................................................... Simulium jacumbae
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 477

Larvae
Not included: Simulium costalimai, S. covagarciai, S. kompi, S. matteabranchium, S. morae, S.
racenisi and S. sandyi.

1 Postgenal cleft only indented or scarcely developed; ratio of hypostomium


length/postgenal bridge = 0.4-0.7 (Figs. 112 A-D) .................................................... 2
– Postgenal cleft scarcely to well developed; ratio of hypostomium/postgenal
bridge = 0.8-1.4 .................................................................................................................. 5
2(1) Medial antennal segment shorter than distal, and proximal segment generally
longer or equal to medial (Fig. 111 B) ........................................................................... 3
– Medial antennal segment longer than distal, and proximal segment shorter than
medial (Fig. 111 L) ............................................................................................................. 4
3(2) Rectal papillae lobes without diverticula (Fig. 111 G) .......................Simulium putre
– Rectal papillae lobes with several diverticula .............................................................
.................................................................................. Simulium anduzei, Simulium jacumbae
4(2) Anal disc with about 106 rows; inner teeth of mandible 13-15 ...........................
................................................................................................................... Simulium marquezi
– Anal disc with 80 or less rows; inner teeth of mandible 10-12 .................................
.......................................................................................... Simulium piperi, Simulium tricorne
5(1) Hypostomium larger than postgenal bridge ................................................................. 6
– Hypostomium as short as or shorter than postgenal bridge (Figs. 112 B, C, G)
................................................................................................................................................ 7
6(5) Ratio of hypostomium/postgenal bridge = 1; anal ring with about 90 rows of
hooks ............................................................................................................... Simulium ruizi
– Ratio of hypostomium/postgenal bridge = 1.2-1.4; anal ring with about 78-80
rows of hooks ............................................................ Simulium parrai, Simulium spilmani
7(5) Scales on anal sclerite present ...........................................................................................
........................ Simulium metallicum, Simulium jobbinsi, Simulium horacioi, Simulium puigi
– Scales on anal sclerite absent ........................................................................................... 8
8(7) Medial antennal segment shorter than proximal and distal segments ......................
....................................................................................................................... Simulium bustosi
– Medial antennal segment longer than proximal and sometimes distal segment as
well (Figs. 111 H, N) ......................................................................................................... 9
9(8) Cephalic apotome without spots ............................................................. Simulium hechti
– Cephalic apotome with median and submedian positive or negative spots ........ 10
10(9) Rectal papillae with 13-14 diverticula on each lobe ............................ Simulium wygoi
– Rectal papillae with 5-8 diverticula on each lobe ...................................................... 11
11(10) Ratio of proximal, medial and distal antennal segments = 1:1.1:1.2 ........................
....................................................................................................................... Simulium iriartei
– Ratio of proximal, medial and distal antennal segments = 1:2.1-2.2:1.3-1.5 ..........
.......................................................................................................................Simulium hunteri
478 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

CLAVES DE ESPECIES DE SIMULIUM (ASPATHIA)


Hembras
Simulium covagarciai, S. kompi, S. ruizi y S. sandyi excluidos por escasa información.

1 Escudo con 1+1 manchas grisáceas no continuadas como bandas claras ............ 2
– Escudo con 3 bandas nacaradas claras o plateadas (Figs. 101 A, F) entre 5 bandas
negras (Figs. 101 B, G) ...................................................................................................... 5
2(1) Escudo con 1+1 manchas blanquecinas subtrapezoidales anteriores (Fig. 101 H);
cibario con área media basal lisa, sin prolongación mediana o reforzada el borde
(Fig. 102 C); paraproctos escasamente desarrollados ventralmente, con borde
redondeado (Fig. 103 D) ............................................................................Simulium putre
– Escudo con 1+1 manchas blanquecinas no subtrapezoidales; cibario con un
proceso mediano o reforzado sobre el borde; paraproctos con desarrollo ventral y
aguzado en el ápice (Figs. 103 J, K) ............................................................................... 3
3(2) Escudo con 1+1 mancha plateada subtriangular; cibario sin una prolongación
mediana o solo ligeramente elevado medianamente (Fig. 102 G) .............................
............................................................................................. Simulium hetchi, Simulium wygoi
– Escudo con 1+1 manchas plateadas subcirculares o dispuestas como una banda
continua anterior; cibario con una pequeña prolongación (Fig. 102 O) ................ 4
4(3) Especies pequeñas, largo ala 3,0 mm; cibario con pequeños dientes esparcidos
(Fig. 102 O) ................................................................................................ Simulium bustosi
– Especies grandes, largo ala aproximadamente 3,8 mm; cibario sin dientes ............
.................................................................................................................. Simulium costalimai
5(1) Escudo visto con iluminación lateral con área polinosa iridiscente; paraproctos
desarrollados ventralmente y aguzados en el ápice (Fig. 103 B) ................................
.... Simulium metallicum, Simulium horacioi, Simulium morae, Simulium puigi, Simulium racenisi
– Escudo visto con iluminación lateral no iridiscente; paraproctos desarrollados o
no ventralmente .................................................................................................................. 6
6(5) Especies relativamente grandes; largo ala 3,3-4,0 mm; cibario liso, sin dentículos,
papilas o prolongación mediana; vesícula sensorial del palpo menos de 1/2 del
largo del palpómero ........................................................................................................... 7
– Especies medianas a pequeñas; largo ala menos de 3,2 mm; cibario liso o no;
vesícula sensorial del palpo generalmente la mitad o más del largo del palpómero
................................................................................................................................................ 9
7(6) Alas con pelos en sector basal de R .................................................... Simulium anduzei
– Alas sin pelos en sector basal de R ................................................................................ 8
8(7) Largo ala 4,0 mm; vesícula sensorial del palpo ligeramente menor de 1/2 del largo
del palpómero ........................................................................................ Simulium marquezi
– Largo ala 3,4 mm; vesícula sensorial del palpo 1/ 2 del largo del palpómero .......
...................................................................................................................... Simulium tricorne
9(6) Cibario liso; largo del ala 2,0-2,5 mm; diente subbasal de la uña ausente o muy
pequeño .................................................................................................... Simulium spilmani
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 479

– Cibario con dentículos o esférulas; largo ala 2,7-3,0 mm; diente subbasal de la uña
bien desarrollado (Fig. 102 M) ...................................................................................... 10
10(9) Cibario sin prolongación mediana con bordes casi rectos; paraprocto más corto
que ancho en la base (Fig. 103 G) ................................................................................ 11
– Cibario con una prolongación mediana (Figs. 102 J-L); paraprocto
aproximadamente tan largo como ancho .................................................................... 12
11(10) Cibario con papilas (Fig. 102 N); paraprocto no aguzado distalmente; vesícula
sensorial del palpo aproximadamente la mitad del largo del palpómero .................
........................................................................................................................ Simulium parrai
– Cibario con pequeños dientes aguzados (Fig. 102 F); paraprocto afinándose
escasamente distalmente (Fig. 103 G); vesícula sensorial del palpo menos de la
mitad del largo del palpómero (Fig. 101 L) .......................... Simulium mattebranchium
12(10) Paraprocto redondeado distalmente; VIII esternito con 29-35 cerdas ................ 13
– Paraprocto aguzado distalmente; VIII esternito con 20-25 cerdas ....................... 14
13(12) Borde interno de las gonapófisis divergente hacia abajo ................... Simulium piperi
– Borde interno de las gonapófisis paralelo ....................................... Simulium jacumbae
14(10) Borde interno de las gonapófisis divergente hacia abajo ................. Simulium iriartei
– Borde interno de las gonapófisis paralelo a gradualmente divergente hacia abajo
.......................................................................................................................Simulium hunteri

Machos
Simulium covagarciai, S. sandyi, S. racenisi y S. kompi no se incluyen por falta de información.

1 Placa ventral subcuadrada, no achatada lateralmente (Figs. 106 A-L) ................... 2


– Placa basal subcónica, subcircular a subcilíndrica, gradualmente achatada desde la
base (Figs. 106 M-Q) ......................................................................................................... 8
2(1) Gonostilo con una prolongación basal larga y delgada; escudo con 1+1 mancha
anterior plateada subtriangular (Fig. 104 I) ............ Simulium jobbinsi, Simulium piperi
– Gonostilo con una prolongación basal relativamente corta y robusta,
frecuentemente rugosa y dentada apicalmente; escudo variable; generalmente con
1+1 banda plateada anterior, convergente, diagonalmente dirigidas hacia la línea
media (Figs. 104 A, B) ....................................................................................................... 3
3(2) Escudo con 1+1 submediana y 1+1 sublateral mancha o banda plateada (Figs. 104
L-N) ...................................................................................................................................... 4
– Escudo con 1+1 mancha plateada de forma variable, subtriangular a subcuadrada
o subromboidal, o como una banda .............................................................................. 5
4(3) Prolongación mediana de la placa ventral ancha basalmente (Fig. 106 G) .............
.......................................................................................................................... Simulium puigi
– Prolongación mediana de la placa ventral angosta basalmente (Fig. 106 L) ...........
.........................................................................................................................Simulium hechti
5(3) Escudo con 1+1 manchas plateadas diagonales prolongándose posteriormente
(Figs. 104 A, B, L) .............................................................................................................. 6
– Escudo con 1+1 mancha plateada no diagonal prolongándose posteriormente
(Fig. 104 D) ......................................................................................................................... 7
480 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

6(5) Placa ventral con una prolongación, mediana, corta y angosta (Fig. 106 I); escudo
con 1+1 manchas plateadas cortas y robustas (Fig. 104 J) ............... Simulium parrai
– Placa ventral con una prolongación mediana a robusta (Figs. 106 A, B)); escudo
con 1+1 manchas plateadas, alargadas y afinadas distalmente (Figs. 104 A, B) ....
................................................. Simulium metallicum, Simulium horacioi, Simulium spilmani
7(5) Placa ventral con una prolongación mediana robusta bordeada por una hendidura,
y los lados curvos (Figs. 106 D-E); prolongación interna basal del gonostilo corta
y lisa (Fig. 105 F) .................................................... Simulium marquezi, Simulium tricorne
– Placa ventral con una prolongación mediana no bordeada por una hendidura y
lados rectos (Figs. 106 C, H, K); prolongación interna basal del gonostilo larga y
dentada distalmente (Figs. 105 G, K) ..............................................................................
........ Simulium bustosi, Simulium iriartei, Simulium jacumbae, Simulium morae, Simulium wygoi
8(1) Escudo sin manchas o bandas aisladas, todo el disco marginalmente plateado (Fig.
104 K) ...................................................................... Simulium costalimai, Simulium anduzei
– Escudo con manchas o bandas plateadas ..................................................................... 9
9(8) Escudo con 1+1 mancha plateada anterior subcuadrada o subromboidal
uniéndose al área lateral plateada; cuerpo de la placa ventral tan ancho como el
diámetro mediano transversal (Fig. 106 M) .........................................Simulium hunteri
– Escudo con 1+1 mancha plateada anterior subtriangular o subtrapezoidal o
bandas diagonales curvas (Figs. 104 E, G) no unidas al área lateral plateada;
cuerpo de la placa ventral más angosto basalmente que el diámetro mediano
transversal (Figs. 106 O-Q) ............................................................................................ 10
10(9) Gonostilo con distintas curvaturas en vista lateral (Fig. 105 E); cuerpo de la placa
ventral fuertemente achatado lateralmente desde la base, subcilíndrico (Fig. 106
Q) .....................................................................................................................Simulium putre
– Gonostilo con lados rectos subparalelos en vista lateral; cuerpo de la placa ventral
no fuertemente achatado lateralmente, subcónico (Figs. 106 J, P) ........................ 11
11(10) Gonostilo con prolongación interna basal más corta que el diámetro basal
transvesal ........................................................................................................ Simulium ruizi
– Gonostilo con prolongación interna basal más larga que el diámetro basal
transversal (Fig. 105 H) ........................................................... Simulium matteabranchium

Pupas
1 Branquias robustas y membranosas (Figs. 107 H, J, 109 A-C. G, I) ....................... 2
– Branquias filamentosas (Figs. 107 A, C, F) ................................................................... 9
2(1) Branquias sin filamentos; frontoclípeo y tórax con tubérculos ............................... 3
– Branquias con filamentos flexibles (Figs. 109 A, E, I); frontoclípeo y tórax sin
tubérculos ............................................................................................................................ 6
3(2) Branquias con ramas digitiformes sin pequeñas ramas secundarias tubuliformes;
capullo sin prolongación anterior (Fig. 107 H) ........................................................... 4
– Branquias con ramas digitiformes con pequeñas ramas secundarias tubuliformes
(Figs. 109 B, C); capullo con (Fig. 107 I) o sin anterior prolongación ................... 5
4(3) Branquia con 3 ramas dirigidas anteriormente, con anillaciones; capullo sin carena
dorsal ............................................................................................................ Simulium kompi
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 481

– Branquia con 2 ramas dirigidas posteriormente y 1 anteriormente, sin anillaciones


(Fig. 107 H); capullo con carena dorsal .............................................. Simulium tricorne
5(3) Branquia con ramas basalmente globosas, subcilíndricas que se angostan
progresivamente distalmente, y la rama principal subdividida distalmente en 7
ramas pequeñas y angostas (Fig. 109 B); tergitos abdominales I y II con tubérculos
................................................................................................................... Simulium marquezi
– Branquia arborescente con 25 ramas terminales relativamente angostas
emergiendo de 3 ramas primarias cortas y robustas (Fig. 109 C); tergitos
abdominales I-II sin tubérculos .............................................................. Simulium sandyi
6(2) Branquia con rama principal subdividida distalmente (Fig. 107 J) .......................... 7
– Branquia subdividida basalmente en 4 grandes ramas ............................................... 8
7(6) Branquia con la rama principal distalmente y basalmente con 1 corta rama
únicamente (Fig. 109 E) .............................................................................. Simulium ruizi
– Branquia con un robusto tallo expandido distalmente, con varias ramas cortas
y pequeñas y basalmente con 3 cortas ramas (Fig. 109 A) ....................................
...................................................................................................... Simulium matteabranchium
8(6) Ramas de las branquias con pseudosegmentaciones, 2 dirigidas anteriormente y 2
ventralmente (Fig. 109 I) .......................................................................... Simulium parrai
– Ramas de las branquias sin pseudosegmentaciones, la más larga dirigida
anteriormente con pequeñas protuberancias subtriasngulares y 2 protuberancias
ventrales subglobosas ......................................................................... Simulium covagarciai
9(1) Ramas terminales de la branquia superando los 8,5 mm; frontoclípeo y tórax con
tricomas simples (Figs. 108 G, H) ................................................................................ 10
– Ramas terminales de la branquia menores de 3,5 mm; frontoclípeo y tórax con
tricomas simples o multirramosas ................................................................................ 12
10(9) Branquia con 3 ramas primarias robustas, subdividida en 7-10 ramas principales
con 85-100 ramas terminales pequeñas (Fig. 108 H); tórax con alrededor de 50
tricomas de cada lado ............................................................................... Simulium iriartei
– Branquias con alrededor de 110-135 ramas (Fig. 108 G); tórax con 70-100
tricomas de cada lado ...................................................................................................... 11
11(10) Frontoclípeo y tórax sin tubérculos, capullo con prolongación dorsal anterior .....
.................................................................................................................. Simulium costalimai
– Frontoclípeo y tórax con moderado número de tubérculos; capullo con borde
anterior recto ..............................................................................................Simulium hunteri
12(9) Branquias con 5 ramas (Fig. 108 E) ................................................... Simulium spilmani
– Branquias con 6-30 ramas .............................................................................................. 13
13(12) Branquias con 6 ramas de aproximadamente el mismo largo (Fig. 108 A); capullo
con borde anterior recto (Figs. 107 A, C) ................................................................... 14
– Branquias con 10-30 ramas; capullo generalmente con prolongación dorsal
anterior o abertura en forma de círculo ...................................................................... 16
14(13) Tricomas del frontoclípeo y tórax con 2-6 ramas (Figs. 107 B, 108 A), tórax con 5
tricomas de cada lado ...................................................................................................... 15
– Tricomas de frontoclípeo y tórax simples; aquellos del tórax en número de 60-85
de cada lado (Figs. 107 C-E) .....................................................................Simulium putre
482 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

15(14) Capullo achatado, subcircular en vista dorsal ................................... Simulium horacioi


– Capullo normal no achatado (Fig. 107 A ) .....................................................................
........ Simulium metallicum, Simulium jabbinsi, Simulium morae, Simulium puigi, Simulium racenisi
16(13) Capullo achatado, subcircular con borde anterior unido ventralmente (Fig. 107 F);
branquia con 10-16 ramas terminales .......................................................................... 17
– Capullo con forma normal, achatado con o sin prolongación dorsal anterior;
branquia con 10-30 ramas .............................................................................................. 19
17(16) Branquia con 10 ramas terminales, con un corto tallo en la base y 4 ramas
primarias; tórax con 25 tricomas de cada lado, con el diámetro disminuyendo
gradualmente hacia el ápice .................................................................... Simulium bustosi
– Branquia con 2-3 ramas principales con 9-11 ramas angostas y flexibles, con 5-120
tricomas que son muy robustos en su 1/3 basal (Fig. 107 K) ............................... 18
18(17) Branquias con 3 ramas principales; tórax con 100-200 tricomas; tórax liso (Fig.
108 J) .............................................................................................................. Simulium hechti
– Branquias con 2 ramas principales; tórax con 5 tricomas de cada lado; tórax
rugoso (Fig. 108 I) ...................................................................................... Simulium wygoi
19(16) Capullo con prolongación dorsal anterior; tórax con 25-65 tricomas de cada lado
.............................................................................................................................................. 20
– Capullos sin proyección anterior dorsal; tórax con 5+ 5 tricomas de cada lado
..................................................................................................................... Simulium anduzei
20(19) Branquia con 10 ramas terminales; tórax con 50-65 tricomas .......... Simulium piperi
– Branquia con 25 (Fig. 108 F) ramas terminales; tórax con 25-35 tricomas ............
................................................................................................................... Simulium jacumbae

Larvas
No incluidas Simulium costalimai, S. covagarciai, S. kompi, S. matteabranchium, S. morae, S.
racenisi y S. sandyi.

1 Hendidura postgenal no evidente o escasamente desarrollada; relación largo


hipostomio/puente postgenal = 0,4-0,7 (Figs. 112 A-D) .......................................... 2
– Hendidura postgenal escasa a bien desarrollada; relación largo hipostomio/
puente postgenal = 0,8-1,4 .............................................................................................. 5
2(1) Artejo antenal mediano más corto que el distal, y el artejo proximal generalmente
más largo o igual al mediano (Fig. 111 B) .................................................................... 3
– Artejo antenal mediano más largo que el distal, y el artejo proximal generalmente
más corto que el mediano (Fig. 111 L) ......................................................................... 4
3(2) Lóbulos de las papilas rectales sin divertículos (Fig. 111 G) ..............Simulium putre
– Lóbulos de las papilas rectales con varios divertículos ................................................
.................................................................................... Simulium anduzei, Simulium jacumbae
4(2) Disco anal con alrededor de 106 hileras de ganchos; mandíbula con 13-15 dientes
internos .................................................................................................... Simulium marquezi
– Disco anal con 80 o menos hileras de ganchos; mandíbula con 10-12 dientes
internos ........................................................................... Simulium piperi, Simulium tricorne
5(1) Hipostomio más largo que el puente postgenal .......................................................... 6
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 483

– Hipostomio tan corto o más corto que el puente postgenal (Figs. 112 B, C, G) ....... 7
6(5) Relación largo hipostomio/puente postgenal = 1; anillo anal con alrededor de 90
hileras de ganchos ........................................................................................ Simulium ruizi
– Relación largo hipostomio/puente postgenal = 1.2-1.4; anillo anal con alrededor
de 78-80 hileras de ganchos .................................... Simulium parrai, Simulium spilmani
7(5) Escamas en el esclerito anal presente ..............................................................................
........................ Simulium metallicum, Simulium jobbinsi, Simulium horacioi, Simulium puigi
– Escamas en el esclerito anal ausente .............................................................................. 8
8(7) Artejo antenal mediano más corto que el proximal y distal ............ Simulium bustosi
– Artejo antenal mediano más largo que el proximal y a veces también del distal
(Figs. 111 H, N) .................................................................................................................. 9
9(8) Apotoma cefálico sin manchas ................................................................. Simulium hechti
– Apotoma cefálico con manchas medianas y submedianas positivas o negativas
.............................................................................................................................................. 10
10(9) Papilas rectales con 13-14 divertículos sobre cada lóbulo ................. Simulium wygoi
– Papilas rectales con 5-8 divertículos sobre cada lóbulo ........................................... 11
11(10) Relación de los artejos antenales proximal, mediano y distal = 1:1.1:1.2 ................
....................................................................................................................... Simulium iriartei
– Relación de los artejos antenales proximal, mediano y distal = 1:2.1-2.2:1.3-1.5
.......................................................................................................................Simulium hunteri

Simulium (Aspathia) anduzei VARGAS & DÍAZ NÁJERA


Simulium (Simulium) anduzei VARGAS & DÍAZ NÁJERA, 1948b: 328-330; 1949: 296-298; 1957b: 264;
COSCARÓN, 1987: 32; CROSSKEY & HOWARD, 1997: 65; 2004: 59; COSCARÓN et al., 1999: 554.
Simulium (Aspathia) anduzei: ADLER et al., 2004: 365-366.
Jalacingomyia anduzei: PY-DANIEL & MOREIRA SAMPAIO, 1994: 101.
Simulium (Simulium) patziciaense TAKAOKA & TAKAHASI, 1982: 63.
Female: Wing length 3.3 mm. Head blackish, frons and clypeus black; clypeus
pollinosity shining blue. Thorax black with median and 1+1 whitish submedian vittae,
subtriangular anteriorly, and 1+1 lateral silvery stripes. Abdominal tergites black with pale
median spots on T II-V. Cibarium smooth. Basal section of R haired. Tarsal claw with
subbasal tooth. Gonapophysis distally truncated and with parallel internal sides, paraproct
short, gradually acuminate ventrally.
Male: Wing length 3.3 mm. Head blackish, clypeus gray pollinose. Thorax blackish,
only ornamented dorsally by lateral silver stripe. Abdominal tergites black with long black
hair; T II-V with 1+1 ovoid sublateral shining gray spots. Gonostylus with prominent
internal basal process smooth distally; ventral plate subconical, becoming gradually
narrower toward apex, similar to S. costalimai and S. hunteri.
Pupa: Cocoon slipper-shaped, without anterior projection but with anterior collar.
Gill length 4.5-4.7 mm, with 17-19 branches originating from long stem. Frontoclypeus
and thorax with spiculae on surface. Frontoclypeus with 1+1 frontal and 1+1 facial simple
trichomes. Thorax with 5+5 simple trichomes. Abdominal sternites VI-VII with only 1
simple or bifurcated posterior hook on each side (from PY-DANIEL, 1994).
484 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Larva: Length of body 7.8-8.0 mm. Body with 1+1 ventral tubercles posteriorly.
Cephalic apotome with positive spots. Cephalic fan with 27-31 primary rays. Antennae longer
than stem of cephalic fan. Ratio of segments proximal, medial and distal = 1:0.9-1.0:1.0-1.2.
Mandible with 6-10 inner teeth and 3 lateral mandibular processes. Hypostomium with 13-15
teeth and 6-7 lateral serrations, lateral setae 5-8; postgenal cleft rounded, shallow. Ratio of
hypostomium/postgenal bridge = 0.5. Anal sclerite with hair, no scales. Anal ring with 65-74
rows of 11-12 hooks each. Rectal papillae with 9-13 diverticula on each lobe.
Distribution: Mexico: Distrito Federal, Oaxaca, Veracruz; USA: Arizona, New Mexico.
Bionomics: This species breeds in spring-fed streams at an altitude of about 2000
m, attached to fallen leaves and stones. Female feeding habits are unknown.
Discussion: The female of S. anduzei is similar to that of S. spilmani and S. tescorum,
but those species have a bare basal section of R. The male is similar to S. costalimai, but
this species has about 102 gill branches. We accept the synonymy of S. patziciaense
proposed by ADLER et al. (2004: 365).

Simulium (Aspathia) bustosi VARGAS, MARTÍNEZ PALACIOS & DÍAZ NÁJERA (Fig. 102 O.)
Simulium (Simulium) bustosi VARGAS et al., 1946: 138-140; VARGAS & DÍAZ NÁJERA, 1957b: 264;
COSCARÓN, 1987: 32; CROSSKEY & HOWARD, 1997: 65; 2004: 59; COSCARÓN et al., 1999: 555.
Female: Wing length 2.8-3.0 mm. General coloration blackish, clypeus and pleurae
gray pollinose. Scutum black with golden hair and 1+1 small anterior silvery subcircular
spots. Abdomen black. Frons parallel sided; fronto-ocular triangle about as long as wide.
Palpus sensory vesicle a little shorter than 1/2 length of palpomere. Basal portion of
cibarium with small median process and sparse denticles (Fig. 102 O). Ratio of length/
width of posterior basitarsus = 7.3. Paraproct shorter than wide at base, distal edge
gradually acuminate, directed ventrally; sternite VIII with about 18-20 hairs on each side;
gonapophysis weakly produced and curved distally; spermatheca well developed.
Male: Wing length 3.1 mm. Scutum velvety black with 1+1 short anterior silver
spots, connected with silver lateral band that extends to posterior margin. Gonostylus
narrow, subcylindrical, with internal basal process robust and dentate on apex. Ventral
plate wider than high, with short median process, shape similar to that of S. metallicum.
Pupa: Cocoon slipper-shaped, with collar present; length basally 3.6-4.0 mm,
dorsally 3.0 mm. Frontoclypeus and exposed portion of thorax with small and sparse
tubercles; thorax with 25-30 simple trichomes. Gill filaments 3.7 mm long, with 10
branches, short stem at base, and 4 primary branches whose first and second dorsal
branches are narrower than others.
Larva: Length 5.7-6.3 mm. Cephalic apotome with 1+1 median longitudinal and 2+2
submedian positive spots. Ratio of hypostomium/postgenal bridge = 0.8-0.9; hypostomium
with 5+5 lateral setae and 3-4 lateral serrations. Cephalic fans with 37-43 primary rays. Ratio
of segments proximal, medial and distal = 1:0.9:0.9-1.0, palpomere robust. Mandible with 8
inner teeth, second marginal tooth well separated from first; 2 anterolateral setae present.
Comb of proleg with about 36 teeth. Anal sclerite with hair, no scales. Anal ring with 66-72
rows of 12-13 hooks each. Rectal papillae lobes without diverticula.
Distribution: Mexico: Oaxaca, Veracruz.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 485

Bionomics: Unknown.
Discussion: The number of gill branches (10) is similar to that for S. piperi, but the
dorsal projection of the cocoon, 50-54 thoracic trichomes, vittae on the female scutum,
and smooth apex of the inner basal process of the male gonostylus allow the separation
of these 2 species.

Simulium (Aspathia) costalimai VARGAS, MARTÍNEZ PALACIOS & DÍAZ NÁJERA


(Figs. 103 K; 104 K; 106 N.)
Simulium (Simulium) costalimai VARGAS et al., 1946: 108, 140-142; VARGAS & DÍAZ NÁJERA, 1957b: 264, 372-
373; COSCARÓN, 1987: 32; CROSSKEY & HOWARD, 1997: 65; 2004: 60; COSCARÓN et al., 1999: 556.
Simulium (Simulium) jacumbae: VARGAS et al., 1943: 289; (not DYAR & SHANNON).
Female: Wing length 3.8 mm. Blackish, frons bright pollinose. Scutum black with
golden pilosity and 1+1 anterior whitish spots connected laterally with silvery marginal
area. Tarsal claw with subbasal tooth well developed. Cibarium with median process not
covered by denticles. Paraproct shorter than high, produced ventrally and gradually
acuminate distally (Fig. 103 K); gonapophysis longer than wide at base, truncated apically;
sternite VIII with 28-30 hairs on each side.
Male: Wing length 3.0 mm. Scutum velvety black without spots or vittae, with
golden pilosity and edged by grayish iridescent pollinosity on lateral, posterior and
anterior margins (Fig. 104 K). Ratio of length/width of hind basitarsus = 3.4. Abdominal
tergites II, IV-VIII with 1+1 silver pollinose spots. Gonostylus subcylindrical, with
internal-basal process well developed, not denticulate apically. Ventral plate subconical,
1.5 times longer than wide, gradually projected distally (Fig. 106 N).
Pupa: Cocoon with thick silk reinforced on edge, with very long dorsal projection
curved ventrally; basal length 4.5-5.0 mm, dorsal length 5.5-6.0 mm. Gill 1.2-1.3 mm long.
Frontoclypeus and thorax without tubercles. Thoracic trichomes simple, elongate, about
80-100 on each side. Gill filaments with about 110-135 branches, with irregular branching.
Abdominal sternite IV with bifurcated hooks.
Larva: Unknown.
Distribution: Mexico: Morelos, Oaxaca.
Bionomics: Unknown.
Discussion: The high number of gill terminal branches (over 100) is similar to that
of S. hunteri, but the cocoon’s straight anterior edge and only about 70 thoracic trichomes
in S. hunteri differentiates these 2 species.

Simulium (Aspathia) covagarciai RAMÍREZ PÉREZ, YARZABAL, TAKAOKA, TADA & RAMÍREZ
Simulium covagarciai RAMÍREZ PÉREZ et al., 1982: 41-44.
S. (Simulium) covagarciai COSCARÓN, 1987: 32; CROSSKEY & HOWARD, 1997: 65; 2004: 60; COSCARÓN
et al., 1999: 556.
(Based on original description).
Female (pharate): Frons and vertex blackish with nacreous pruinosity and silver hair.
Thorax black with golden hair, color pattern unknown. Abdomen blackish, tergites VI-
486 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

VIII shining. Fronto-ocular triangle as long as wide. Sensory vesicle of palp 1/3 length of
palpomere. Cibarium base smooth but with small, sclerotized, rounded papillae. Paraproct
1/2 as long as wide at base.
Male (pharate): Thorax and abdomen velvety black, thorax with golden hair;
abdominal tergites II, VI-VIII with 1+1 silver lateral spots. Gonostylus with median
constriction, with internal basal process. Ventral plate about as long as wide, with short
median process.
Pupa: Cocoon threads thick and conspicuous. Basal length 4.75 mm, dorsal length
4.50 mm. Frontoclypeus and thorax without tubercles, frontoclypeus with 1+1 facial and
2+2 frontal, simple, filiform trichomes. Thorax with about 40 simple, filiform trichomes.
Gill membranous, with 4 subcylindrical, smooth dorsal lobes, 2 ventral lobes subglobose,
with some hair, the frontal largest, with small subtriangular protuberances and some long
hair basally.
Larva: Unknown.
Distribution: Venezuela: Amazonas, Sierra de Parima (950 m).
Bionomics: Unknown.
Discussion: The closest species is S. matteabranchium, but S. covagarciai can be differentiat-
ed by the gills of the pupa.

Simulium (Aspathia) hechti VARGAS, MARTINEZ-PALACIOS & DÍAZ NÁJERA


(Figs. 104 M, N; 105 J; 106 L, 107 K; 108 J.)
Simulium (Simulium) hechti VARGAS et al., 1946: 142; VARGAS & DÍAZ NÁJERA, 1957b: 264, 372-373;
COSCARÓN, 1987: 32; CROSSKEY & HOWARD, 1997: 65; 2004: 60; COSCARÓN et al., 1999: 557.
Simulium (Aspathia) hechti: ADLER et al., 2004: 366.
Simulium (Simulium) sayi: VARGAS et al., 1943: 288; (not DYAR & SHANNON).
Female: Wing length 3.3 mm. Blackish, clypeus silvery pollinose, scutum black with 1+1
anterior whitish triangle-shaped spots and silvery edging posteriolaterally. Abdomen dark
brownish with 1+1 spot on tergite II. Basal portion of cibarium smooth, weakly reinforced on
edge. Frons slowly convergent, fronto-ocular triangle about as long as wide at base. Sensory
vesicle of palp near half of palpomere length. Ratio of length/width of hind basitarsus = 6.0.
Paraprocts about as long as wide at base, slightly acuminate distally. VIII stemite with 25-26
setae; gonapophysis with convex margin, inner margins parallel and reinforced.
Male: Wing length 3.1 mm. Scutum reddish brown with 1+1 median ovoid, and 1+1
diagonal cuneiform spots (Figs. 104 M, N), anterior stripes silvery pollinose. Hind
basitarsus wide, ratio of length/width = 3.0. Gonostylus subcylindrical with strongly
wrinkled internal basal process denticulate apically (Fig. 105 J). Ventral plate wider than
high, slightly flattened, with short and wide subconical median process (Fig. 106 L).
Pupa: Cocoon with marginal collar and projected antero ventrally, threads well
defined. Basal length 3.4-4.2 mm, dorsal length 2.0-2.5 mm. Gill length 25-30 mm.
Frontoclypeus and exposed portion of thorax with abundant tubercles. Trichomes of
head and thorax simple, with peculiar thickening on basal 1/3 and filiform on distal 3/4
(Fig. 107 K), 100-120 on each side of thorax. Gill with 3 main filiform branches bearing 9-
11 narrower flexible branches (Fig. 108 J).
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 487

Larva: Cephalic apotome without ornamentation or spots. Postgenal cleft short,


ratio of hypostomium/postgenal bridge = 0.8; hypostomium with 7-8 lateral setae and 5-
7 lateral serrations. Ratio of segments proximal, medial and distal = 1:1.5:1.3. Cephalic fan
with 32-34 primary rays. Mandible with 8 inner teeth and 1 lateral mandibular process.
Comb of proleg with about 28-32 teeth. Anal sclerite with sparse hair, no scales. Anal ring
with about 75-80 rows of 11 -13 hooks each. Rectal papillae with 6 diverticula on each
lobe, with distal lobule larger than others.
Distribution: Mexico: Durango, Distrito Federal, Morelos; USA: Arizona, New
Mexico.
Bionomics: Simulium hechti breeds in mountainous areas. Female feeding habits are
unknown.
Discussion: The most similar species is S. wygoi, especially in the female, but the
pupal gill with two main branches and wrinkled thorax with five trichomes on each side in
S. wygoi will differentiate them.

Simulium (Aspathia) horacioi OKAZAWA & ONISHI


Simulium (Simulium) horacioi OKAZAWA & ONISHI, 1980: 167-173; COSCARÓN, 1987: 32; CROSSKEY &
HOWARD, 1997: 69; 2004: 63; GRILLET & BARRERA, 1997: 197, 200, 206; COSCARÓN et al., 1999: 558.
Simulium horacioi: TAKAOKA, 1982: 49.
?Simulium metallicum (cytoform “H”) CONN et al., 1989: 1217.
This species is very close to S. metallicum. In S. horacioi, small differences occur: the
paraprocts are shortest distally, the median process of the ventral plate is more projected
ventrally, the pupal cocoon is flattened and rounded, and the cephalic apotome of the
larva has positive spots and segments VII-IX are blackish brown dorsally. Simulium horacioi
may correspond to any cytotype of the S. metallicum species complex. It would be useful to
do complementary studies.
Distribution: Guatemala; Mexico: Chiapas, Oaxaca; Venezuela: Carabobo.
Bionomics: The species breeds in small rivers, tributaries and channels in mountain-
ous areas at about 1300 m. It is an anthropophilic and zoophilic species.

Simulium (Aspathia) hunteri MALLOCH (Figs. 102 K; 106 M; 108 G.)


Simulium (Simulium) hunteri MALLOCH, 1914: 59; PETERSON & KONDRATIEFF, 1995: 39; CROSSKEY &
HOWARD, 1997: 65; 2004: 60; COSCARÓN et al., 1999: 559-560.
Simulium (Aspathia) hunteri: ADLER et al., 2004: 366.
Simulium (Simulium) lassmanni VARGAS, MARTÍNEZ PALACIOS & DÍAZ NÁJERA, 1946: 149, 174; VARGAS
& DÍAZ NÁJERA, 1957b: 264; COSCARÓN, 1987: 32; CROSSKEY & HOWARD, 1997: 65.
Female: Wing length 3.1 mm. General coloration blackish. Scutum ornamented with
silvery lyre-shaped stripes; 1+1 anterior, subtriangular silvery spots when viewed with
other illumination angles. Abdomen blackish, last segment shining. Frons convergent
below. Sensory vesicle of palp less than 1/2 length of palpomere. Fronto-ocular triangle
about as long as wide. Basal portion of cibarium with denticulate median process (Fig. 102
K). Ratio of length/width of hind basitarsus = 6.0. Paraprocts about as long as wide with
488 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

curved posterior edge. Stemite VIII with 20-25 hairs on each side; gonapophysis
truncated, with parallel inner margins.
Male: Wing length 3.0 mm. Scutum blackish, with yellow pilosity and 1+1 silvery
anterior subrhomboidal spots meeting laterally, lateral and posterior margins silvery. Legs
blackish with fore coxa and hind basitarsus whitish on basal half. Ratio of length/width
of hind basitarsus = 4.7. Abdomen black, with 1+1 silvery spots on tergites II and VII.
Gonostylus curved, subcylindrical, with internal-basal process as long as transverse
diameter of gonostylus at base and not dentate apically. Ventral plate subconical-elongate,
slightly longer than wide at base (Fig. 106 M).
Pupa: Cocoon with straight, reinforced anterior edge, loosely woven with well–
defined threads. Basal length 4.3 mm, dorsal length 3.6 mm. Frontoclypeus and thorax
with moderate number of tubercles arranged in groups; trichomes simple and elongate,
those of thorax about 70 on each side. Gill arborescent, with about 130 branches,
branching from 7 primary branches (Fig. 108 G), length about 1.2-1.5 mm.
Larva: Length 5.8 mm. Cephalic apotome with median longitudinal and 1+1
discontinuous sublateral positive spots. Ratio of hypostomium/postgenal bridge = 0.8.
Postgenal bridge 1.2 times height of hypostomium; hypostomium with 7-9 lateral setae
and 3-4 lateral serrations. Ratio of segments proximal, medial and distal = 1:2.1-2.2:1.3-
1.5. Cephalic fan with about 35-38 primary rays, combs of rays with 7 intermediate minor
teeth alternating with 1 bigger tooth. Inner mandibular teeth 10-11. Proleg comb with 28
teeth. Anal sclerite with hair, crochet circlet with 72-76 rows of 13-14 hooks. Rectal
papillae with 5 diverticula on each lobe.
Distribution: Canada: Alberta, British Columbia, Saskatchewan, Yukon; Mexico:
Distrito Federal, Michoacan; USA: Alaska, Arizona, California, Colorado, Idaho, New
Mexico, Nevada, Oregon, South Dakota, Texas, Utah, Washington, Wyoming.
Bionomics: The species breeds in small creeks, frequently with fast current. Females
bite livestock and humans.
Discussion: The closest species is S. costalimai; differences are discussed under that
species.

Simulium (Aspathia) iriartei VARGAS, MARTÍNEZ PALACIOS & DÍAZ NÁJERA


(Figs. 105 G; 106 H; 108 H; 111 N.)
Simulium iriartei: VARGAS et al., 1946: 144-146, 173-174; VARGAS & DÍAZ NÁJERA, 1957b: 264;
COSCARÓN, 1987: 32; CROSSKEY & HOWARD, 1997: 65; 2004: 60; COSCARÓN et al., 1999: 560.
Simulium (Aspathia) iriartei: ADLER et al., 2004: 366-367.
Simulium hirtipes: VARGAS, 1943a: 143; (not FRIES).
Female: Wing length 3.0 mm. Body blackish. Scutum with 1+1 anterior light grayish,
subtriangular spots continued as longitudinal, very narrow median stripes. Frons conver-
gent below, fronto-ocular triangle about as long as wide. Sensory vesicle of palp more
than 1/2 length of palpomere. Tarsal claw with subbasal tooth well developed. Ratio of
length/width of hind basitarsus = 6.5. Basal portion of cibarium with median process
covered by denticles. Paraprocts longer than wide at base; gonapophyses reinforced on
inner margins and gradually divergent below; stemite VIII with 21-25 hair.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 489

Male: Wing length 2.8 mm. Head blackish, clypeus shining whitish. Scutum black
with 1+1 subrhomboidal anterior silvery spots with green hair, edged laterally by green
metallic pollinose stripe. Gonostylus with internal-basal process well developed, wrinkled
and dentate distally (Fig. 105 G). Ventral plate subquadrate basally, horseshoe-like, with
small median process, subacute distally (Fig. 106 H).
Pupa: Cocoon with reinforced anterior margin and strongly projected dorsally, with
conspicuous threads. Basal length 4.4-5.0 mm, dorsal length 4.2-5.0 mm. Gill 2.0 mm
long, arborescent, with about 85-100 terminal filaments branching basally from 3 stout
primary branches, subdivided 3-6 times to end (Fig. 108 H). Frontoclypeus and thorax
with sparse tubercles, trichomes simple and elongate; thorax with about 50 trichomes on
each side. Scales on abdominal stemites III-VIII conspicuous as small spines.
Larva: Length 6.3 mm. Cephalic apotome gradually darkened basally, with 1 median
anterior, 1 subbasal and 1+1 submedian positive spots. Postgenal bridge 1.2 times longer
than height of hypostomium, ratio of hypostomium/postgenal bridge = 0.8, hypostomi-
um with 7 lateral setae and 4-6 lateral serrations. Antennae longer than stem of cephalic
fan. Ratio of segments proximal, medial and distal =1:1.1: 1.1-1.2 (Fig. 111 N). Cephalic
fan with 32-34 primary rays. Mandible with 10-11 internal teeth. Thoracic proleg comb
with 22-24 teeth arranged in 6-8 groups. Anal sclerite with hair, no scales; anal ring with
72-76 rows of 13-14 hooks each; rectal papillae with 5-8 diverticula on each lobe.
Distribution: Mexico: Distrito Federal, Durango, Guerrero, Oaxaca, Veracruz;
USA: Arizona, New Mexico.
Bionomics: Simulium iriartei breeds in small creeks in mountainous areas. Female
feeding habits are unknown.
Discussion: Simulium hunteri and S. costalimai are the closest species. The first can be
separated from S. iriartei by the elongate and apically smooth inner basal process of the
gonostylus; the second species can be separated by the female scutum with 1+1 anterior
grayish spots. Both of these similar species can be distinguished from S. iriartei in the
pupal stage by having a gill with more than 100 terminal branches.

Simulium (Aspathia) jacumbae DYAR & SHANNON (Figs. 102 J; 106 K; 108 F.)
Simulium jacumbae DYAR & SHANNON, 1927: 44-45, VARGAS & DÍAZ NÁJERA, 1948b: 333-334.
Simulium (Aspathia) jacumbae: ADLER et al., 2004: 367.
Simulium (Simulium) jacumbae: VARGAS et al., 1946:146; DALMAT, 1951: 54; 1955: 208-213; VARGAS &
DÍAZ NÁJERA, 1957b: 264, 376-377; STONE & BOREHAM, 1965: 170; COSCARÓN, 1987: 32;
CROSSKEY & HOWARD, 1997: 65; 2004: 60; COSCARÓN et al., 1999: 561-562.
Simulium guatemalense DE LEÓN, 1945: 75.
Female: Wing length 2.5-2.8 mm. General coloration dark grayish brown. Scutum
ornamentation with lyre-shaped pattern as in S. metallicum, with gray pollinosity and
golden to silver hair. Legs blackish, with basal half of median and hind tibiae whitish.
Abdomen black, tergites V-VIII shining. Frons convergent below, fronto-ocular triangle as
long as wide. Basal portion of cibarium with median process, wrinkled on anterior edge
(Fig. 102 J). Palpal sensory vesicle 1/2 length of palpomere. Sc and basal section of R not
haired. Tarsal claw with subbasal tooth well developed. Paraprocts as long as wide,
490 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

rounded distally. Stemite VIII with 29-35 setae. Gonapophyses with straight distal edge,
internal margins sclerotized and parallel-sided.
Male: Wing length 2.6 mm. General coloration as in female. Scutum similar to that
of S. metallicum but 1+1 anterior triangles smaller, hair golden to tin color. Hair on legs
and first abdominal segment yellowish brown. Ratio of length/width of hind basitarsus =
4.3. Abdomen black with 1+1 silvery spots on tergites II, V-VII. Gonostylus about twice
length of gonocoxite, subcylindrical, acuminate distally, with internal basal process
narrow, almost as long as transverse diameter of gonostylus at base and dentate on edge;
ventral plate subquadrate basally, gradually elongate distally and about as long as wide,
with sparse hair (Fig. 106 K).
Pupa: Cocoon with anterodorsal process, loosely woven and with threads visible.
Basal length 3.0-3.3 mm, dorsal length 3.6-4.2 mm. Gill length 2.0 mm, with 24-25
filaments, stem short, with 3 primary branches: ventral primary branch short and stouter,
with 2 large secondary branches as stout as other primary branches (Fig. 108 F).
Frontoclypeus and thorax with sparse tubercles, cephalic and thoracic trichomes simple,
those of thorax about 23-35 on each side. Hooks of stemites VI-VIII bi- or trifurcated.
Larva: Length 6.7 mm. Cephalic apotome with median and submedian positive spots.
Postgenal cleft shallow. Ratio of hypostomium/postgenal bridge = 0.4-0.6. Hypostomium
with 4 lateral setae, and 5-7 lateral serrations. Antennae longer than base of cephalic fans.
Ratio of segments proximal, medial and distal = 1:0.9:1.2, without evident transverse
striations. Cephalic fan with 55-60 primary rays. Inner mandibular teeth in number 8-10 in 2
series, with 2 lateral mandibular processes. Anal sclerite without hair or scales; anal ring with
about 70 rows of 12 hooks each; rectal papillae with 1 -3 diverticula on each lobe.
Distribution: Guatemala; Mexico: Chiapas, Hidalgo, Jalisco, Michoacan, Oaxaca,
Veracruz; USA: Arizona, California, Colorado, Kansas, Nebraska, Nevada, Texas, Utah,
Wyoming.
Bionomics: The species breeds in lowlands, in small streams. Females bite livestock
and humans.
Discussion: Simulium piperi is closest to S. jacumbae but the latter can be differentiat-
ed by the female inner margins of the gonapophyses being parallel and the pupal gill
having 25 terminal branches.

Simulium (Aspathia) jobbinsi VARGAS, MARTÍNEZ PALACIOS & DÍAZ NÁJERA


Simulium (Simulium) jobbinsi VARGAS et al., 1946: 147-149; VARGAS & DÍAZ NÁJERA, 1957b: 264, 378-
379; DALMAT, 1951: 54; 1955: 213; COSCARÓN, 1987: 32; CROSSKEY & HOWARD, 1997: 69; 2004:
63; COSCARÓN et al., 1999: 562-563.
Female: Wing length 2.6 mm. General color blackish, with ornamentation on thorax and
abdomen similar to that of S. metallicum (with iridescent pruinosity on thorax and 1+1 silver
spots on tergite II). Frons with subparallel sides; fronto-ocular triangle as long as wide at base.
Cibarium smooth. Sensory vesicle of palp about 1/2 length of segment. Tarsal claw with
subbasal tooth well developed; ratio of length/width of hind basitarsus = 5.8. Basal portion of
R without hair. Paraproct longer than wide, gradually acuminate. Stemite VIII with 37-40 hairs
on each side. Gonapophyses truncate, inner margins reinforced and divergent posteriorly.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 491

Male: Wing length 2.4 mm. Color black, very similar to S. metallicum but 1+1 latero-
frontal cuneiform silvery spots shorter (not prolonged posteriorly). Legs slightly darker
than in S. metallicum. Gonostylus subcylindrical, curved, with well-developed internal basal
process almost as long as transverse diameter of gonostylus at base; ventral plate wide,
with short median process very similar to that of S. metallicum.
Pupa: Cocoon with straight edge, loosely woven. Basal length 3.3-3.5 mm, dorsal
length 3.0-3.2 mm; gills 4.0-4.3 mm. Frontoclypeus and exposed portion of thorax with
abundant rounded tubercles. Gill with 6 filaments, stem short, and with well-defined
primary branches. Cephalic trichomes with 3-4 branches.
Larva: Length 6.2-6.4 mm. Cephalic apotome with small gradually darkened median
longitudinal spot. Ratio of hypostomium/postgenal bridge = 0.9; hypostomium with 4 to
5 lateral setae on each side and 6-8 lateral serrations. Antenna typical of the subgenus;
ratio of segments proximal, medial and distal = 1:1.1:1. Cephalic fan with 40-45 primary
rays. Mandible with 9-10 inner teeth. Anal sclerite with hair and scales basally; anal ring
with 76-80 rows of 13-14 hooks each. Ninth sternite with 2 tubercles. Rectal papillae with
3 lobes, each with 6 diverticula.
Distribution: Guatemala: Altaverapaz, Bajaverapaz, Chimaltenango, Guatemala,
Huehuetenango, Quezaltenango, San Marcos, Sacatepequez, Solola, Suchitepequez, Toto-
micapan; Mexico: Chiapas, Oaxaca, Veracruz.
Bionomics: Unknown.
Discussion: Simulium jobbinsi is very close to S. metallicum, separated only by small
differences in the male: the ventral plate and silvery diagonal stripes on the thorax are shorter.
Females cannot be separated from those of S. metallicum and S. horacioi (TAKAOKA, 1982).
Molecular and cytological studies would be useful to assure the validity of this species.

Simulium (Aspathia) kompi DALMAT


Simulium (Simulium) kompi DALMAT, 1951: 38; 1955: 218; COSCARÓN, 1987: 32; CROSSKEY & HOWARD,
1997: 66; 2004: 60; COSCARÓN et al., 1999: 563.
(Diagnosis based on DALMAT, 1955) (female and male pharate).
Female: Paraproct higher than long, gradually acuminate distally; cercus with convex
edge; gonapophysis with internal edge parallel, distal edge straight.
Male: Gonostylus parallel-sided, with small bifid terminal spur; subbasal process
small; ventral plate longer than wide, gradually acuminate and haired distally, without
strong lateral constriction.
Pupa: Cocoon slipper-shaped, without anterior collar, aperture reinforced. Gill thick
and membranous, composed of 3 inflated, tube-like branches, with transverse striations,
the dorsal one measuring 2.1 mm, the middle one 1.8 mm and the ventral one 1.3 mm.
Larva: Length 6.2 mm. Cephalic apotome darkened, with 1 basal median and 1+1
sublateral elongate spots. Cephalic fan with 28 primary rays. Antennae as long as stalk of
cephalic fan, with transverse striations on proximal and medial segments. Medial segment
longer than palpomere, and medial and distal subequal. Hypostomium with 6+6 lateral
serrations. Postgenal cleft shallow, not acuminate apically. Thoracic proleg comb sclerite
with 38-40 teeth. Anal sclerite with hair, no scales; anal ring with 78-82 rows of hooks.
492 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Distribution: Guatemala: Chimaltenango; Mexico: Chiapas.


Bionomics: Unknown.
Discussion: The three-lobed pupal gill branches are similar to those of S. kompi and
S. tricorne, but the latter have no annulations and the branches have a different orientation.
There are slight differences in the female, such as the distally truncate cerci and
gonapophyses divergent below.

Simulium (Aspathia) marquezi VARGAS & DÍAZ NÁJERA


(Figs. 104 D; 105 F; 106 D; 107 I; 109 B; 111 L; 112 A.)
Simulium (Simulium) marquezi VARGAS & DÍAZ NÁJERA, 1957a: 37; COSCARÓN, 1987: 33; CROSSKEY &
HOWARD, 1997: 66; 2004: 60; COSCARÓN et al., 1999: 564-565.
Simulium vandalicum: VARGAS et al., 1946: 109 (larva); (not DYAR & SHANNON).
Female: Wing length 4.0 mm. General coloration blackish. Scutum ornamented with
lyre-shaped markings and silvery posterolateral margin, pleurae brownish, with silver
pollinosity. Legs blackish, with basal 1/3 of middle basitarsus and basal 2/3 of hind
basitarsus whitish. Abdomen blackish, with 1+1 silvery spots on tergites II-IV, tergites V-
VIII shining. Frons relatively wide; fronto-ocular triangle as long as wide at base. Sensory
vesicle of palp a little shorter than 1/2 length of palpomere. Basal portion of cibarium
smooth. Sc with 1-3 hairs, basal section of R bare. Ratio of length/width of hind
basitarsus = 8; tarsal claw with subbasal tooth well developed. Paraprocts rounded distally,
shorter than wide at base, cerci truncate. Stemite VIII with 15-20 setae on each side;
gonapophysis truncate distally, with internal edge divergent and more sclerotized.
Male: Wing length 3.8 mm. Scutum dark brown, with pilosity and silver pruinosity
on lateral and posterior margins, prolonged into subapical diagonal bands separated from
each other by black band at midline (Fig. 104 D). Pleura as in female. Abdomen dark
brown or black, concolorous with thorax, with median silvery pollinose spots on tergites
II-VII when viewed from behind. Gonostylus subcylindrical, with strong internal basal
process, without denticles on edge (Fig. 105 F), ventral plate wider than long, with
anterolateral angular projections and small rounded median projections (Fig. 106 D).
Pupa: Cocoon with dorsal projection keel-shaped, loosely woven, anterior edge
reinforced (Fig. 107 I). Basal length 5.0-5.2 mm, dorsal length 5.5-6.3 mm. Frontoclypeus
and exposed portion of thorax with abundant tubercles irregularly arranged and with
small spiculae on surface, frontoclypeal trichomes simple and small, difficult to see.
Thorax with about 30-33 short, curved trichomes. Gill membranous, with 2 globose
subcylindrical basal branches bifurcated at median distal portion and becoming narrower
at 1/4-1/3 of length distally, main dorsal branch subdivided distally into 7 small, narrower
branches (Fig. 109 B). Abdominal onchotaxy evident, tergites I and II with tubercles.
Larva: Length 6.8-7.3 mm. Cephalic apotome with 1+1 submedian spots not well
defined, diffuse brownish area present mediobasally. Hypostomium with 9+9 lateral setae
and 6-7 lateral serrations; Postgenal cleft reduced (Fig. 112 A); ratio of hypostomium/
postgenal bridge = 0.6. Antennae with second segment wrinkled (Fig. 111 L); ratio of
segments proximal, medial and distal = 1:1.3:1. Cephalic fan with 40-42 primary rays,
several small teeth, with 1 longest, alternating on combs. Inner mandibular teeth 13-15,
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 493

with secondary tooth row, marginal tooth with 1 large cusp and another smaller one at
base. Thoracic proleg comb with about 22 teeth, not arranged in groups. Anal sclerite with
abundant setae, no scales; crochet circlet with 106 rows of 15 hooks each.
Distribution: Mexico: Chihuahua, Distrito Federal, Morelos.
Bionomics: Unknown.
Discussion: Simulium tricorne is very similar morphologically, but the small body size,
proportionally shorter sensory vesicle of the palp in the female, absence in the pupa of
small tubuliform gill branches and cocoon with a straight anterior edge permit separation.

Simulium (Aspathia) matteabranchium ANDUZE


(Figs. 101 L; 102 F; 103 F-H; 105 H, I; 106 P; 107 J; 109 A.)
Simulium matteabranchia ANDUZE, 1947: 374; VARGAS & DÍAZ NÁJERA, 1948b: 333-334; PERRUOLO
LANETI, 1971: 397; RAMÍREZ PÉREZ, 1971: 290; 1983: 53.
Simulium (Simulium) matteabranchia: COSCARÓN, 1987: 33.
Simulium (Simulium) matteabranchium: CROSSKEY, 1987: 471; CROSSKEY & HOWARD, 1997: 66; 2004: 60;
COSCARÓN et al., 1999: 565-566.
Female: Wing length 2.7-2.8 mm. Body blackish. Frontoclypeus silvery pollinose.
Thorax with lyre-shaped pattern; scutum with decumbent hair, some darker and stronger
than others. Abdomen black with 1+1 silvery spots on tergite II. Legs dark brownish, hind
tibia whitish on basal 3/4; ratio of length/width of hind basitarsus = 7.5; tarsal claw with
subbasal tooth well developed. Frons parallel-sided, moderately wide; fronto-ocular
triangle about as long as wide. Cibarium with anterior edge not elevated and with
abundant small sclerotized papillae at base (Fig. 102 F). Palpal sensory vesicle slightly
shorter than 1/2 length of palpomere (Fig. 101 L). Sc and base of R bare. Stemite VIII
with 17-21 setae; gonapophysis curved distally, with internal edge sclerotized and 5+5
setae (Fig. 103 F); paraprocts shorter than wide at base, gradually acuminate distally (Fig.
103 G); genital fork with long, acuminate apodemes (Fig. 103 H).
Male: Wing length 2.3 mm. Body blackish; scutum black with 1+1 anterior
subtriangular silvery spots. Legs brownish black, lighter only on anterior half of second
and third basitarsi. Sensory vesicle of palp more than 1/2 length of palpomere. Ratio of
length/width of hind basitarsus = 3.6-3.7. Gonostylus relatively narrow with large
internal basal process slightly longer than height of gonocoxite (Fig. 105 H). En-
doparameres with strong hooks and poorly defined median sclerite (Fig. 105 I). Ventral
plate subconical, laterally flattened, acuminate distally (Fig. 106 P).
Pupa: Cocoon with reinforced edge, tightly woven, without visible threads (Fig. 107
J). Gill membranous, with 3 branches over thorax and head produced anteriorly as thick
stalk distally expanded into several lobules and with thin, filiform branches (Fig. 109 A).
Frons with 2+2 frontal and 1+1 simple facial trichomes; frontoclypeus and exposed
portion of thorax without tubercles. Thorax with about 52-104 trichomes on each side,
very narrow beyond middle, curved and simple, and relatively wide. Abdominal tergite I
with sparse acuminate tubercles medially.
Larva: Ratio of hypostomium/postgenal bridge = 1.1. Antenna as long as stalk of
cephalic fans. Mandible with 11 inner teeth.
494 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

A B

G
F

C
E

Figure 101. Female. A-E: S. (Aspathia) metallicum; A-B: scutum with different illumination; C-E:
anterior, middle and posterior legs, respectively. F-G: S. (A.) piperi, scutum with different
illumination. H-J: S. (A.) putre; H: scutum; I: maxillary palp; J: sensory vesicle of palp. K: S. (A.)
wygoi, sensory vesicle of palp. L: S. (A.) matteabranchium, sensory vesicle of maxillary palp.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 495

Distribution: Colombia: Cundinamarca; Mexico(?); Venezuela: Aragua, Carabo-


bo, Distrito Federal, Merida, Monagas, Tachira, Trujillo.
Bionomics: Simulium matteabranchium is an anthropophilic species and occurs
between 700 and 1800 m, according to RAMÍREZ PÉREZ (1971).
Discussion: Simulium parrai is the most closely related species and is differentiated
by the female scutum not ornamented with vittae, cibarium base with papillae, paraproct
rounded distally, and pupa with the gill having 4 large lobes: 2 basal and 2 anterior.

Simulium (Aspathia) metallicum BELLARDI


(Figs. 101 A-E; 102 S; 103 A, B; 104 A-C; 105 A-D; 106 A, B; 107 A, B; 108 A; 110; 111 M; 112 C.)
Simulium metallicum BELLARDI, 1859: 13; DYAR & SHANNON, 1927: 41; FAIRCHILD, 1940: 712; VARGAS,
1942a: 234-236; VARGAS et al., 1946: 175; DALMAT, 1955: 220-223; VARGAS & DÍAZ NÁJERA,
1957b: 264; LEWIS & LEE POTTER, 1964: 99; WYGODZINSKY, 1971: 12-14; RAMÍREZ PÉREZ, 1971:
275; 1983: 54; 1984: 79; SHELLEY & ARZUBE, 1985: 328.
Simulium (Simulium) metallicum OKAZAWA & ONISHI, 1980: 167-179; COSCARÓN, 1987: 33; CROSSKEY &
HOWARD, 1997: 69; 2004: 63; COSCARÓN et al., 1999: 566-568; SHELLEY et al., 2002: 171-176;
COSCARÓN ARIAS, 2003: 196, 204, 208, 215.
Simulium (Aspathia) metallicum: ADLER et al., 2004: 365 (as species group).
Simulium riveti ROUBAUD, 1906b: 108; WYGODZINSKY, 1953b: 332; SHELLEY & ARZUBE, 1985: 328.
Simulium nitidum MALLOCH, 1912: 652.
Simulium versicolor LUTZ & NUÑEZ TOVAR, 1928: 47.
Simulium avidum HOFFMANN, 1930a: 51; 1930b: 293.
Simulium violacescens ENDERLEIN, 1934b: 285.
“A”-”K” CONN et al., 1989 (cytoform).
“L” ARTEAGA & MUÑOZ DE HOYOS, 1999 (cytoform).
“X” MILLEST, 1989 (thesis) (cytoform).
Female: Wing length 1.9-2.3 mm. Head blackish, frons and clypeus grayish
pollinose; frons parallel-sided. Scutum black, with 3 longitudinal black stripes separated by
2 narrower light ones plus 1+1 submedial grayish pollinose stripes in frontal view, that
change to black when viewed from behind as do lateral margins (Figs. 101 A, B), in lateral
view scutum has shining metallic greenish iridescence, hair of scutum black and
recumbent; scutellum blackish brown; pleurae blackish gray. Legs brownish black,
ornamentation as in figures 101 C-E. Abdomen blackish brown, tergites II, IV-VIII with
1+1 silvery pollinose spots that become iridescent when viewed with lateral light, tergites
VI-VIII shining. Sensory vesicle of palp about 1/3 length of palpomere. Cibarium
smooth. Basal section of R bare. Calcipala and pedisulcus as in figure 102 S; tarsal claw
with subbasal tooth well developed. Ratio of length/width of hind basitarsus about 6.
Stemite VIII with 27-30 hairs on each side; gonapophyses with subparallel sides and
posterior edge truncate; spermatheca well developed (Fig. 103 A); paraproct subtriangular,
gradually acuminate from base to apex, longer than wide at base; cercus with rounded
edge (Fig. 103 B).
Male: Wing length 2.0-2.3 mm. General color blackish. Scutum velvety black with
silvery anterior diagonal bands from anterolateral band to midline, but not quite reaching
496 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

0.05 mm

O
C F

G N J

K L

H P

B
0.1 mm 0.1 mm

E
S Q
A
0.03 m

R
m
0.02 m
m

m
3m
0.1 mm 0.0

I M

Figure 102. Female. A-E: S. (Aspathia.) putre. A: front; B: fronto-ocular triangle; C: basal portion of
cibarium; D: furcasternum; E: subbasal claw tooth. F: S. (A.) matteabranchium, cibarium. G-I: S. (A.)
wygoi; G: cibarium; H: frons; I: subbasal claw tooth. J: S. (A.) jacumbae, cibarium. K: S. (A.) hunteri,
cibarium. L-M: S. (A.) piperi; L: cibarium, M: claw. N: S. (A..) parrai, cibarium. O: S. (A.) bustosi,
cibarium. P-R: S. (A.) sandyi; P: frons; Q-R: subbasal claw tooth in different orientations. S: S. (A.)
metallicum, calcipala and pedisulcus.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 497

E
H
m
5m
0.0

J B

mm
0.05 mm

0.05
G

F
0.32 mm

K
C

Figure 103. Female. A-B: S. (Aspathia) metallicum. A: eighth sternite and gonapophysis, spermatheca,
genital fork; B: paraproct and cercus in different position. C-E: S. (A.) putre; C: eighth sternite and
gonapophysis; D: paraproct and cercus; E: genital fork. F-H: S. (A.) matteabranchium; F: eighth sternite
and gonapophysis; G: paraproct and cercus; H: genital fork. I-J: S. (A.) wygoi; I: eighth sternite and
gonapophysis; J: paraproct and cercus. K: S. (A.) costalimai; paraproct and cercus.
498 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

midline and not joined (Figs. 104 A, B) in lateral light, silver areas become iridescent. Legs
similar to those of female; ratio of length/width of hind basitarsus about 4.8-5.4.
Abdomen black, with 1+1 silvery spots on tergites II, IV-IX (Fig. 104 C). Gonocoxite and
gonostylus as in figures 105 A-C; gonostylus with well-developed internal basal process
and rounded edge; ventral plate subquadrate, with short median process (Figs. 106 A, B);
endoparameral organ with robust spines (Fig. 105 D).
Pupa: Cocoon basal length 3.6-4.0 mm, dorsal length 2.4-3.2 mm; body length 3.2
mm; gills 3.6-4.0 mm long; cocoon slipper-shaped, loosely woven, slightly reinforced
anteriorly (Fig. 107 A). Frontoclypeus and thorax with abundant tubercles. Frontoclypeus
with 1+1 facial and 2+2 bifid trichomes (Fig. 107 B); thorax with 5+5 bifid or trifid
trichomes on disc (Fig. 108 A). Gill with 6 elongate filaments, with 3 short primary
branches emerging from short stem. Abdominal chaetotaxy as in figure 110.
Larva: Length 4.8-5.6 mm. Head light yellowish brown; cephalic apotome slightly
brownish pigmented midbasally, with negative spots. Antenna longer than stem of
cephalic fan; ratio of proximal, medial and distal segments = 1:1.5-1.6:0.9-1.2 (Fig. 111
M). Cephalic fan with 35-40 primary rays. Mandible with 9-10 inner teeth and 2 lateral
mandibular processes. Postgenal cleft shallow (Fig. 112 C); ratio of hypostomium/
postgenal bridge = 0.9:1.0-1.1; hypostomium with 4+4 lateral setae and 4-5 lateral
serrations. Sclerite of thoracic proleg with 18-22 teeth arranged in 5-6 groups. Anal
sclerite with sparse hair and abundant spiculae; anal ring with 80 rows of 12-14 hooks
each; rectal papillae with 3-4 diverticula on each lobe.
Distribution: Colombia: Cundinamarca, Magdalena, Valle; Costa Rica; Belize; Cayo,
Toledo; Ecuador: Azuay, Carchi, Chimborazo, Esmeraldas, Imbibura, Napo, Pichincha,
Tungurahua, Zamora; Guatemala: Altaverapaz, Bajaverapaz, Chimaltenango, Chiquimula,
El Peten, El Progreso, El Quiche, Escuintla, Guatemala, Huehuetenango, Jutiapa, Quetzalt-
enango, Retalhuleu, Sacatepequez, San Marcos, Santa Rosa, Solola, Suchitepequez; Jamaica;
Mexico: Chiapas, Chihuahua, Colima, Durango, Guerrero, Hidalgo, Jalisco, Mexico,
Michoacan, Morelos, Nayarit, Nuevo León, Oaxaca, Puebla, San Luis Potosi, Sinaloa,
Sonora, Tabasco, Veracruz; Panama: Cocle, Colon; Peru: Huanuco; Trinidad; Venezuela:
Anzoategui, Aragua, Carabobo, Cojedes, Distrito Federal, Falcon, Guarico, Lara, Merida,
Miranda, Monagas, Nueva Esparta, Portuguesa, Sucre, Tachira, Trujillo, Yaracuy, Zulia.
Bionomics: Simulium metallicum breeds in small creeks frequently in shaded areas and
also in rivers, attached to submerged vegetation and fallen leaves, in slow to fast currents.
Females attack humans and livestock and are considered vectors of Onchocerca volvulus.
Discussion: Simulium metallicum is a widely distributed species complex and shows
some morphological variation. Cytological studies indicate that it is a species complex of
at least 7 cytospecies. CONN (1988) established cytotypes “A” and “B” from Guatemala
and Mexico, “C” from Colombia, “D” and “E” from Venezuela, “F”, “J” and “K” from
Panama, “G” from Costa Rica, “H” from Guatemala, Mexico, Panama, and “I” from
Guatemala and Mexico. ARTEAGA & MUNOZ DE HOYOS (1999) found 2 cytotypes in
Colombia, and MILLEST (1990) established differences between cytotypes and larval
morphology. The closest species are S. horacioi, S. jobbinsi and S. racenisi; differences from
each species are explained under those species. A detailed explanation of similarities with
those species was given by SHELLEY et al. (2002: 174-175).
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 499

A
B

I
E G

M F

J
H
C

L K N

Figure 104. Male. A-C: S. (Aspathia) metallicum; A-B: scutum dorsal and lateral view, respectively. C:
abdomen, lateral view. D: S. (A.) marquezi, scutum, dorsal view. E-G: S. (A.) putre, scutum, different
position and illumination. H: S. (A.) putre, hind leg. I: S. (A.) piperi, scutum. J: S. (A.) parrai, scutum.
K: S. (A.) costalimai, scutum. L: S. (A.) puigi, scutum. M-N: S. (A.) hechti, scutum, dorsal view from
different position.
500 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

F
E
m
m
05
0.

H
m
5m

0.08 mm
0.0

I G
0.08 mm

m
5m
0.0

J K L

Figure 105. Male. Gonocoxite and gonostylus. A-D: S. (Aspathia) metallicum; A: with ventral plate,
endoparameral organ and median sclerite. D: endoparameres, median sclerite and aedeagal
membrane in different position. E: S. (A.) putre. F: S. (A.) marquezi. G: S. (A.) iriartei. H-I: S. (A.)
matteabranchium; H: gonocoxite, gonostylus; I: endoparameral organ and median sclerite. J: S. (A.)
hechti. K: S. (A.) wygoi. L: S. (A.) spilmani.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 501

Simulium (Aspathia) morae RAMÍREZ PÉREZ, RASSI & RAMÍREZ


Simulium (Simulium) morae RAMÍREZ PÉREZ et al., 1977: 167; COSCARÓN, 1987: 33; CROSSKEY &
HOWARD, 1997: 69; 2004: 63; COSCARÓN et al., 1999: 569.
Simulium morae: RAMÍREZ PÉREZ et al., 1982: 48-50; 1983: 59; RAMÍREZ PÉREZ, 1983: 98.
(Based on original description).
Female: Vertex and frontoclypeus blackish, bright nacreous. Scutum dark, with
golden hair, ornamentation similar to that of S. metallicum but without shining iridescence.
Pleurae blackish with nacreous pollinosity. Abdomen with tergite II nacreous, the
following tergites shining black with golden pilosity. Fronto-ocular triangle narrower than
high at base. Sensory vesicle of palp about 1/2 length of segment. Basal portion of
cibarium elevated medially, with abundant teeth. Tarsal claw with subbasal tooth well
developed. Paraproct subtriangular, shorter than wide at base.
Male: General coloration blackish as in female. Thorax black, with golden pilosity
and with 1+1 anterior submedian dark triangles edged by nacreous areas. Gonostylus
subcylindrical, acuminate distally, with robust internal basal process; ventral plate shorter
than wide, subquadrate, with median haired process.
Pupa: Cocoon length 4.0 mm, with loose, irregular weave. Frontoclypeus and thorax
with tubercles abundant medially. Cephalic trichomes with 2-3 branches, thoracic
trichomes with 2-5 branches. Gill composed of 6 branches with short stem and primary
branches.
Larva: Unknown.
Distribution: Venezuela: Amazonas, Dpto. of Atabapo, Sierra Parima.
Bionomics: Unknown.
Discussion: This species is close to S. metallicum and S. racenisi but is differentiated in
the female by a medially elevated cibarium with small denticles, which is unique among
Neotropical Simulium (Aspathia) species, and in the male by a narrower gonostylus.

Simulium (Aspathia) parrai VARGAS, MARTÍNEZ PALACIOS & DÍAZ NÁJERA


(Figs. 102 N; 104 J; 106 I; 109 H, I.)
Simulium (Simulium) parrai VARGAS et al., 1946: 151-175; VARGAS & DÍAZ NÁJERA, 1948b: 336; 1957b:
264; DALMAT, 1951: 55; 1955: 230; COSCARÓN, 1987: 33; CROSSKEY & HOWARD, 1997: 65; 2004:
60; COSCARÓN et al., 1999: 569-570.
Female: Wing length 3.0 mm. Scutum blackish not ornamented, covered with dense
silvery pilosity. Sensory vesicle of palp about 1/2 length of palpomere. Frons gradually
convergent below, fronto-ocular triangle about as long as wide. Basal portion of cibarium
with abundant papillae in middle (Fig. 102 N). Scutum without special ornamentation,
with dark, strong hair intermixed with paler, weaker hair. Sc and basal section of R
without hair. Ratio of length/width of hind basitarsus = 6.0. Tarsal claw with subbasal
tooth well developed. Paraproct slightly shorter than wide, not acuminate distally. Stemite
VIII with 25-29 hairs on each side; gonapophysis truncate, with slightly diverging,
reinforced inner edge; spermatheca oval-shaped; genital fork with small submedian
process on lateral branches.
502 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

B C
A
0.03 mm

0.05 mm

0.04 mm

D
E F

G H I

J
0.03 mm

L
K 0.04 mm
M

0.05 mm

N O Q

Figure 106. Male. Ventral plate. A-B: S. (Aspathia) metallicum, in different position. C: S. (A.) wygoi.
D: S. (A.) marquezi. E: S. (A.) tricorne. F: S. (A.) piperi. G: S. (A.) puigi. H: S. (A.) iriartei. I: S. (A.)
parrai. J: S. (A.) spilmani. K: S. (A.) jacumbae. L: S. (A.) hechti. M: S. (A.) hunteri. N: S. (A.) costalimai.
O: S. (A.) ruizi. P: S. (A.) matteabranchium. Q: S. (A.) putre.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 503

Male: Wing length 2.7 mm. Scutum reddish brown, with pale golden iridescent
pilosity and 1+1 anterior silvery diagonal cuneiform spots; lateral and posterior margins
with silver pollinosity (Fig. 104 J). Ratio of length/width of hind basitarsus = 4.1.
Gonostylus subcylindrical, with strong internal basal process; ventral plate body subquad-
rate, with short median process (Fig. 106 I).
Pupa: Basal length 4.1 mm, dorsal length 3.0 mm; gill about 1.5 mm long. Cocoon
flattened, anterior rim reinforced with conspicuous threads. Frontoclypeus and thorax without
tubercles; thorax with 7-12 simple trichomes (Fig. 109 H). Gill membranous and pseudo-
annulated, with 1 basal and 4 large distal lobes with abundant and long filaments on median
basal area (Fig. 109 I). Abdominal tergite II with 3 submedian setae developed as small hooks.
Larva: Length 5.2-5.6 mm. Cephalic apotome with diffuse ornamentation on basal half,
1+1 spots and median basal darkened area present. Postgenal cleft miter-shaped. Postgenal
bridge slightly shorter than height of hypostomium; ratio of hypostomium/postgenal bridge
= 1.1; hypostomium with 6-7 lateral setae and 3-4 lateral serrations. Cephalic fan with 40-44
primary rays. Ratio of proximal, medial and distal segments = 1:1.1:0.8. Mandible with 9-11
inner teeth arranged in 2 rows, with 1 lateral mandibular process. Anal sclerite with hair, no
scales; anal ring with 78 rows of 14-15 hooks each; rectal papillae not seen.
Distribution: Guatemala: Chimaltenango, Solola; Mexico: Oaxaca, Veracruz.
Bionomics: Unknown.
Discussion: This species is similar to S. matteabranchium. Simulium covagarciai is similar
in the globose membranous respiratory organ, but the unique morphology and disposi-
tion of the lobules in S. covagarciai allows differentiation.

Simulium (Aspathia) piperi DYAR & SHANNON (Figs. 101 F, G; 102 L, M; 104 I; 106 F.)
Simulium piperi DYAR & SHANNON, 1927: 38.
Simulium (Aspathia) piperi: ADLER et al., 2004: 367.
Simulium (Simulium) piperi PETERSON & KONDRATIEFF, 1995: 39; CROSSKEY & HOWARD, 1997: 66;
2004: 60; COSCARÓN et al., 1999: 572-573.
Simulium knowltoni TWINN, 1938: 53.
Simulium stonei STAINS & KNOWLTON, 1943: 277.
Simulium sayi DYAR & SHANNON, 1927: 40.
Female: Wing length 3.0 mm. General coloration grayish brown. Head brownish,
frons and clypeus grayish pollinose; frons gradually converging, fronto-ocular triangle
about as high as wide. Sensory vesicle of palp about 1/2 length of palpomere. Basal
portion of cibarium smooth, with small median prominence (Fig. 102 L) Thorax blackish
brown with light gray pilosity; scutum when viewed from behind with silvery lyre-shaped
pattern delimited by median and 1+1 sublateral black stripes (Fig. 101 F), and with
anterior illumination, 1+1 subtriangular light grayish spots, followed by 1+1 black vittae
that reach posterior edge (Fig. 101 G); metanotum velvety grayish brown. Abdomen
brown with 1+1 silvery spots on tergite II, tergites VI-VII shining. Wing veins yellowish
brown, with brown hair, Sc with 3-4 hairs. Legs brown, darker on apices of tibiae and tarsi,
ratio of length/width of hind basitarsus = 6.1, tarsal claw with subbasal tooth well
developed (Fig. 102 M). Stemite VIII strongly sclerotized, about 27-29 setae on each side;
504 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

gonapophyses with distal edge truncate, inner edges more sclerotized, divergent below;
paraprocts rounded distally, shorter than wide at base.
Male: Wing length 2.8 mm. General coloration brownish, darker than female.
Scutum blackish brown with grayish yellow decumbent hair, when viewed with lateral light
from front, with 1+1 silvery triangular spots about 1/6 length of scutum (Fig. 104 I); in
other lights, scutum silvery laterally and posteriorly. Ratio of length/width of hind
basitarsus = 3.5. Gonostylus subcylindrical, twice as long as gonocoxite; internal-basal
process long and smooth; ventral plate wider than high, with broad median process
gradually acuminate distally (Fig. 106 F).
Pupa: Cocoon projected dorsally; basal length 4.5-4.7 mm, dorsal length 4.0 mm, threads
conspicuous. Gill with 10 very open branches and with short basal stem, 4 primary branches of
about same length present. Thorax with 50-65 simple hairs. Frontoclypeus and dorsal portion
of thorax with moderate number of small and smooth tubercles. Frontoclypeus with 1+1
facial and 2+2 frontal simple trichomes; thorax with 50-54 simple trichomes.
Larva: Cephalic apotome with positive spots edged by diffuse pigmentation;
cephalic fan with 35-38 primary rays; ratio of proximal, medial and distal segments =
1:1.3-1.5:1.0-1.1. Ratio of hypostomium/postgenal bridge = 0.5; hypostomium with 4-5
lateral setae and 8-9 lateral serrations. Inner mandibular teeth 10-12. Anal sclerite with
hair, no scales. Comb of thoracic proleg with 12-15 teeth, not grouped. Anal ring with 72-
76 rows of 14-15 hooks each; rectal papillae lobes with 5-7 diverticula each.
Distribution: Canada: Alberta, British Columbia, Saskatchewan; Mexico: Distrito
Federal; USA: Arizona, California, Colorado, Idaho, Montana, Nebraska, New Mexico,
Oregon, South Dakota, Texas, Utah, Washington, Wyoming.
Bionomics: Simulium piperi breeds in small turbulent streams attached to aquatic
vegetation. Females attack livestock and also bite humans.
Discussion: Simulium jacumbae is the most closely related species; differences
between them were discussed under S. jacumbae. Variation in morphology (e.g., gill
branches numbering 9-13) suggests that S. piperi is a species complex (ADLER et al., 2004).

Simulium (Aspathia) puigi VARGAS, MARTÍNEZ PALACIOS & DÍAZ NÁJERA


(Figs. 104 L; 106 G; 108 B, C.)
Simulium (Simulium) puigi VARGAS et al., 1946: 153-155, 176; VARGAS & DÍAZ NÁJERA, 1957b: 191;
TAKAOKA & TAKAHASI, 1982: 63; COSCARÓN, 1987: 33; CROSSKEY & HOWARD, 1997: 69; 2004:
63; MOULTON, 1996: 741; COSCARÓN et al., 1999: 573.
Simulium (Aspathia) puigi: ADLER et al., 2004: 368.
Female: Wing length 3.1 mm. Body color blackish. Scutal ornamentation lyre-
shaped, very similar to S. metallicum, appearing metallic green iridescent when viewed with
lateral illumination. Legs predominantly yellowish. Abdomen dark dorsally, tergite II with
1+1 silvery pollinose spots, tergites VII-IX shining cinereous. Basal portion of cibarium
smooth. Paraprocts 1/2 as long as wide, rounded distally; gonapophysis subcircular, with
sclerotized inner margin. Stemite VIII with 26-30 hairs on each side.
Male: Wing length 3.0 mm. Scutum velvety blackish with 1+1 suboval median and
1+1 diagonal anterior silver spots, joining basally with lateral and posterior marginal silver
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 505

band (Fig. 104 L). Legs blackish, whitish basally on fore tibia, on basal half of middle
tarsus, and hind basitarsus; ratio of length/width of hind basitarsus = 3.2. Abdomen
blackish with 1+1 spots on tergite II, tergites VI-VII with 1+1 silvery greenish pollinose
spots. Gonostylus subcylindrical, with internal basal process slightly developed; ventral
plate subquadrate, as long as wide at base with wide median process basally (Fig. 106 G).
Pupa: Cocoon slipper-shaped, heavy, with texture well defined and strongly reinforced
on anterior edge, slightly projected ventrally; basal length 4.2 mm, dorsal length 3.4 mm; gill
length 4.0-5.5 mm. Frontoclypeus and exposed portion of thorax with abundant large,
elevated tubercles, surface wrinkled, and bearing spinules. Facial trichomes with 3 branches.
Thoracic trichomes with 3-5 long branches on disc and 2-4 simple ones on lateral edge. Gill
with 6 branches, with short stem and relatively short primary branches (Figs. 108 B, C).
Larva: Length 6.0-6.5 mm. Cephalic apotome without spots or special ornamenta-
tion. Ratio of hypostomium/postgenal bridge = 0.8; hypostomium with 4-5 lateral setae
and 2-4 lateral serrations on each side. Cephalic fan with 28-36 primary rays. Postgenal
cleft medially developed, subtriangular; ratio of hypostomium/postgenal bridge = 0.8-1.0.
Ratio of proximal, medial and distal segments = 1:1.2-1.3:1.1. Mandible with 8-10 inner
teeth. Anal sclerite with scales and hair; anal ring with 64 rows of 11-12 hooks each; rectal
papillae lobes with 5-8 diverticula each.
Distribution: Costa Rica; Guatemala; Mexico: Aguascalientes, Chiapas, Oaxaca,
Veracruz; USA: New Mexico.
Bionomics: This species breeds in creeks with moderate flow and aquatic vegeta-
tion, from lowlands to 2700 m of altitude. Female feeding habits are unknown.
Discussion: Simulium puigi is very difficult to separate from other species of
Simulium (Aspathia). The wider spindle-shaped posterior basitarsus, the silver thoracic
spots meeting with the lateral and posterior silvery area, the shorter basal process on the
gonostylus in the male and absence of ornamentation on the cephalic apotome of the
larva help identify S. puigi.

Simulium (Aspathia) putre COSCARÓN & MATTA


(Figs. 101 H, J; 102 A-E; 103 C-E; 104 E-H; 105 E; 106 Q; 107 C-E; 108 D; 111 A-G; 112 D-F.)
Simulium putre COSCARÓN & MATTA, 1982: 52-54; 1987; CROSSKEY & HOWARD, 1997: 81; 2004: 60.
Simulium (Simulium) putre COSCARÓN, 1987: 33; 1991: 268-269; COSCARÓN et al., 1999: 574-575;
CROSSKEY & HOWARD, 2004: 60.
Female: General coloration blackish. Frons and clypeus blackish, frons shining
nacreous, clypeus with silver pollinosity. Scutum blackish, when viewed with lateral
illumination with 1+1 anterior subtrapezoidal silvery spots and marginal silvery pollinosity
laterally and posteriorly (Fig. 101 H), with shifting light, becoming black; hair of scutum tin
colored. Frons convergent below (Fig. 102 A); fronto-ocular triangle about as long as tall
(Fig. 102 B). Maxillary palp as in figure 101 I. Cibarium smooth basally (Fig. 102 C). Sensory
vesicle of palp about 1/2 length of palpomere (Fig. 101 J). Leg hair filiform; tarsal claw with
well-developed subbasal tooth (Fig. 102 E). Gonapophyses parallel-sided along inner
margins and with sclerotized margin truncate distally (Fig. 103 C); paraproct gradually
produced ventrally, cerci with rounded edge (Fig. 103 D); genital fork as in figure 103 E.
506 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

B
C

1 mm

F 1 mm

J
1 mm 0.2 mm

G
E
0.05 m
m
0.02 mm 0.2 mm

Figure 107. Pupa. A-B: S. (Aspathia) metallicum; A: lateral view; B: frontoclypeus. C-E: S. (A.) putre;
C: lateral view; D: frontoclypeus; E: thoracic trichomes. F-G: S. (A.) wygoi; F: dorsal view; G:
frontoclypeus. H: S. (A.) tricorne. I: S. (A.) marquezi. J: S. (A.) matteabranchium. K: S. (A.) hechti,
thoracic trichomes.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 507

D
m
0.4 m

0.5 mm
E

0.2 mm
I 0.4
mm
B

0.2
mm

H C

mm
0.2

F
0.4 m
m

J 0.4
mm

Figure 108. Pupa. Gill and thoracic trichomes. A: S. (Aspathia) metallicum. B-C: S. (A.) puigi. D: S.
(A.) putre. E: S. (A.) spilmani. F: S. (A.) jacumbae. G: S. (A.) hunteri. H: S. (A.) iriartei. I: S. (A.) wygoi.
J: S. (A.) hechti.
508 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Male: Wing length 2.8 mm. Scutum black, when viewed with lateral illumination with
1+1 silvery anterior spots and marginal silver pollinosity laterally and posteriorly (Figs. 104 E,
F), with the same but more inclined illumination, color changes to black, with light from front,
sublateral elongate silvery spots appear (Fig. 104 G). Abdomen black with 1+1 silvery spots on
tergites II, V-VII. Sc and basal section of R bare. Pigmentation of hind leg as in figure 104 H;
ratio of length/width of posterior basitarsus = 3.5. Gonostylus without internal-basal process
but with high lateral keel from base to middle length and lateral submedian sinuosity, apically
flattened with elongate depression on inner side where small subapical spur is inserted (Fig.
105 E); body of ventral plate flattened and curved, with ventral setae (Fig. 106 Q); parameres
with well-developed teeth, median sclerite subrhomboidal.
Pupa: Cocoon basal length 3.5-3.7 mm, dorsal length 3.0 mm, loosely woven; like chenille
(Fig. 107 C). Frontoclypeus and thorax with moderate to abundant tubercles (Fig. 107 D), and
simple slender hair (Fig. 107 E); frontoclypeus with 2+2 frontal and 1+1 facial trichomes;
thorax with about 60-85 trichomes on each side. Gill grayish, with 6 filaments, stem and
primary branches short (Fig. 108 D). Abdomen with terminal hooks relatively little developed.
Larva: Maximum length 6.5 mm. General aspect as in figure 112 F. Head yellowish
brown, body greenish gray. Cephalic apotome with positive spots (Fig. 111 A). Antenna
longer than stem of cephalic fans; ratio of proximal, medial and distal segments = 1:0.8:1;
proximal and medial segments with conspicuous transverse striations (Figs. 111 B-C).
Mandible with 10 inner and 12 acuminate marginal teeth (Fig. 111 D). Cephalic fan with 30-
34 primary rays. Postgenal cleft very short, subtriangular (Fig. 112 D); ratio of hypostomi-
um/postgenal bridge = 0.5; hypostomium teeth as in figure 112 E, lateral setae 3-4 and 5-7
lateral serrations on each side. Proleg sclerite with about 20 comb teeth arranged in groups
and in 2 rows (Fig. 111 E). Anal sclerite with hair, no scales (Fig. 111 F); anal ring with 72
rows of 12-14 hooks each; rectal papillae with 3 simple lobes (Fig. 111 G).
Distribution: Chile: Arica, Parinocota, Tarapaca.
Bionomics: Simulium putre inhabits creeks at 3500 m elevation, where larvae and
pupae are attached to aquatic vegetation.
Discussion: Simulium putre is an isolated species and has the southernmost
distribution of this subgenus. It has a unique female and male scutal ornamentation and
male gonostylus morphology. The pupa, with 6 filamentous gill branches, is similar to that
of S. metallicum, but the presence of trichomes with 2-6 branches and the 5+5 thoracic
trichomes permit separation.

Simulium (Aspathia) racenisi RAMÍREZ PÉREZ


Simulium racenisi RAMÍREZ PÉREZ, 1971: 332; 1983: 66.
Simulium (Simulium) racenisi COSCARÓN, 1987: 33; CROSSKEY & HOWARD, 1997: 69; 2004: 63;
COSCARÓN et al., 1999: 575.
Simulium sp. G. 1962 LEWIS & IBAÑEZ ALDECOA, 1962: 449; LEWIS, 1963: 62.
(Based primarily on original description).
Female: Body color blackish. Frontoclypeus nacreous pollinose; fronto-ocular triangle
slightly wider than high. Sensory vesicle of palp about 1/3 length of segment. Basal portion of
cibarium smooth. Scutum ornamentation very similar to that of S. metallicum. Scutellum dark
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 509

brownish. Pleurae blackish, with silver pollinosity. Legs blackish, with femur and fore tibia, base
of hind femur and most of middle and hind basitarsi whitish. Abdomen blackish, tergites V-
VIII shining; tergite II with 1+1 silvery spots. Tarsal claw with subbasal tooth well developed.
Paraproct about as long as high, with truncate edge; gonapophysis wider than tall.
Male: Gonostylus subcylindrical, with rounded apex and short intemal-basal
process; ventral plate wider than tall.
Pupa: Gill with 6 branches. Frontoclypeus with 3+3 bifid trichomes. Thorax with 5-
6 bifid or trifid trichomes on each side.
Larva: Cephalic apotome with small 1+1 median longitudinal and 1+1 sublateral
spots; postgenal bridge as long as hypostomium.
Distribution: Venezuela: Miranda, Monagas.
Bionomics: Unknown.
Discussion: Simulium racenisi is very close to S. metallicum, but according to RAMÍREZ-
PEREZ, S. racenisi has longer primary and secondary gill branches, a shorter median process
on the ventral plate, a stouter gonostylus distally in the male, and shorter paraprocts in the
female. Because we were unable to examine specimens, we are uncertain about the validity
of this species, particularly in view of the differences in genitalic characters, which depends
on how they are viewed.

Simulium (Aspathia) ruizi VARGAS & DÍAZ NÁJERA (Figs. 106 O; 109 E-G.)
Simulium (Simulium) ruizi VARGAS & DÍAZ NÁJERA, 1948b: 330; 1957b: 265, 392; COSCARÓN, 1987: 33;
CROSSKEY & HOWARD, 1997: 66; 2004: 60; COSCARÓN et al., 1999: 576-577.
Female: Wing length 2.6 mm. Body color blackish. Frons and clypeus blackish,
clypeus with dirty white pruinosity. Cibarium with small rounded denticles. Sensory
vesicle of palp nearly 1/2 as long as segment. Scutum black, when seen from behind with
frontal light, there is 1 slender median blackish stripe and 1+1 wider submedian silvery
pollinose vittae anteriorly, lateral and posterior margins silvery pollinose, when seen in
frontal view, the color is inverted, with 1+1 submedian vittae separated by 0.25 of width
of one, and 1+1 sublateral vittae nearly twice as wide as submedian bands of silver
pollinosity. Scutellum yellowish brown. Abdomen concolorous with thorax, tergite II
silvery pollinose. Legs predominantly yellowish brown. Paraprocts half-moon-shaped,
nearly as long as basal width; stemite VIII with 16-18 hairs on each side.
Male: Wing length 3.0 mm. Body blackish. Scutum black with 1+1 diagonal curved
bands not joining silvery marginal band. Legs blackish, with basal 1/3 of hind basitarsus
yellowish; posterior basitarsus wide, ratio of length/width = 2.7-3.2. Abdomen black,
with 1+1 silvery spots on tergites I-VII. Gonostylus subcylindrical, with internal basal
process smooth and shorter than basal transverse diameter; ventral plate subconical, 1.3
times longer than wide, with strong basal branches (Fig. 106 O).
Pupa: Cocoon projected dorsally with anterior edge reinforced, tissue compact with
conspicuous threads. Frontoclypeus and thorax without tubercles. Trichomes simple,
curved, about 45-50 on each side. Gill membranous, haired and subcylindrical, with short
basal and 3 apical branches (Fig. 109 E), and filaments simple (Fig. 109 F) or with several
branches (Fig. 109 G).
510 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

B
A

C
0.4 mm

F
D

0.03 mm
mm
0.05

G I
0.1 mm
0.
4m

H
m

Figure 109. Pupa. Gill and thoracic trichomes. A: S. (Aspathia) matteabranchium. B: S. (A.) marquezi.
C-D: S. (A.) sandyi; C: gill; D: thoracic trichome and tubercles. E-G: S. (A.) ruizi; E: gill; F-G:
appendages of gill. H-I: S. (A.) parrai; H: thoracic sclerite with gill insertion and trichomes; I:
frontal view.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 511

I
II
II

III III

IV IV

V V

VI
VI
VII
VII

VIII
VIII
IX

Figure 110. Pupa. Abdominal chaetotaxy of S. (Aspathia) metallicum.

Larva: Length 6.5 mm. Cephalic apotome scarcely darkened subbasally with 1 median
and 1+1 submedian elongated spots. Postgenal cleft subtriangular, acuminate cephalad.
Ratio of hypostomium/postgenal bridge = 1.1; hypostomium with subdental platform, 8-9
lateral setae and 2-3 lateral serrations on each side. Ratio of antennal segments proximal,
medial and distal = 1:1.3:0.8. Cephalic fan with about 35-40 primary rays. Mandible with 9-
11 inner teeth arranged in 2 rows, second marginal tooth very close to first. Thoracic proleg
comb with 30 teeth arranged in about 7 groups with 1+1 well defined apodemes. Anal
sclerite with simple hair, no scales; anal ring with 90 rows of 15 hooks each.
Distribution: Mexico: Oaxaca, Veracruz.
Bionomics: Unknown.
Discussion: This species is very close to S. matteabranchium. The peculiar membra-
nous and flexible filaments of the gill are similar to those of S. matteabranchium, S. parrai
and S. covagarciai, but the subdivision of branches allows differentiation from other
Neotropical species of the subgenus Aspathia.
512 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Simulium (Aspathia) sandyi COSCARÓN, IBÁÑEZ BERNAL & COSCARÓN ARIAS


(Figs. 102 P-R; 109 C, D.)
Simulium (Simulium) sandyi COSCARÓN et al., 1999: 577-578; CROSSKEY & HOWARD, 2004: 60.
Female (Description based on pharate female): Frons wide (Fig. 102 P), fronto-
ocular triangle about as long as high. Basal portion of cibarium smooth, without denticles
or spherules. Sensory vesicle of palp 1/2 length of palpomere. Basal section of R bare.
Tarsal claw with subbasal tooth well developed (Figs. 102 Q, R).
Male: Unknown.
Pupa: Cocoon loosely woven and covered by detritus. Frontoclypeus and thorax
with abundant tubercles, with spiculate surface (Fig. 109 D). Thorax with 16-19 simple,
robust trichomes on each side. Gill membranous, without hair, with 25 terminal branches
emerging from 3 short, stout primary branches, secondary branches digitiform, and
terminal portion subdivided (Fig. 109 C).
Larva: Unknown.
Distribution: Mexico: Distrito Federal, Lagunas de Zempoala.
Bionomics: Unknown.
Discussion: The pupa of S. marquezi is closest to S. sandyi, but the gill has 2 robust
basal branches and only 7 terminal branches, and thus is easily separated.

Simulium (Aspathia) spilmani STONE (Figs. 105 L, 106 J; 108 E .)


Simulium (Simulium) spilmani STONE, 1969: 313; CROSSKEY, 1987: 474; CROSSKEY & HOWARD, 1997: 69;
2004: 60; COSCARÓN et al., 1999: 578.
(Diagnosis based on original description).
Female: Wing length 2.0 mm. Frons shining black, convergent below. Sensory
vesicle of palp about 0.4 times length of palpomere. Basal portion of cibarium flat,
unarmed. Scutum black with 1+1 submedian and 1+1 sublateral silvery pollinose stripes,
anterior portion of sublateral stripes iridescent. Abdominal tergite II with 1+1 iridescent
spots, tergites II-V velvety black, tergites VI-VIII shining black. Wing with Sc and basal
section of R bare. Stemite VIII, gonapophysis, paraprocts and cerci unknown.
Male: Scutum black with 1+1 oblique silvery stripes on frontal declivity, broad blue
to gold velvety lateral stripe, iridescent blue on anterior edge. Leg hair filiform and
petaloid; ratio of length/width of posterior basitarsus = 6. Gonostylus spatuliform, with
thin subapical spur; subbasal process short, with abundant denticles (Fig. 105 L); ventral
plate wide basally, with narrow median process edged by deep emargination (Fig. 106 J).
Pupa: Body length 2.2 mm. Head and thorax with small tubercles, sparser on thorax;
head with 2+2 bifid trichomes, thorax with 5-6 simple to bifid and 1+1 simple trichomes
on each side. Gill 1.5-1.8 mm long, with 5 terminal branches, stalk short, with primary
dorsal and ventral branches short and bifurcated, lateral branch simple, length similar to
secondary branches (Fig. 108 E).
Larva: Length 5.0-5.5 mm. Cephalic apotome darkened mediobasally, subtriangular,
without positive head spots. Postgenal cleft deep, with anterior edge subacute; ratio of
hypostomium/postgenal bridge = 1.1; hypostomium with 4+4 lateral serrations and 5
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 513

setae on each side. Antennae transversally striated on proximal and medial segments; ratio
of antennal segments proximal, medial and distal = 1:1.2-1.3:0.8-0.9. Mandible with 10-11
inner teeth and 1 lateral mandibular process, secondary marginal teeth very small.
Cephalic fan with 38-40 primary rays. Thoracic proleg comb with 24-26 teeth arranged in
8-9 main groups. Anal sclerite with scarce hair, no scales; anal ring with 75-80 rows with
14 hooks each; rectal papillae with 3 simple lobes.
Distribution: Dominica.
Bionomics: Unknown.
Discussion: The unique 5 branches of the gill of the pupa is a useful character to
differentiate S. spilmani from other species of the subgenus Aspathia. The scutal
ornamentation of the female and shape of the male genitalia place S. spilmani close to S.
metallicum.

Simulium (Aspathia) tricorne DE LEÓN (Figs. 106 E; 107 H.)


Simulium tricornis DE LEÓN, 1945: 68.
Simulium (Simulium) tricornis: VARGAS et al., 1946: 155-176; VARGAS & DÍAZ NÁJERA, 1957b: 265, 398;
DALMAT, 1955: 234; COSCARÓN, 1987: 33.
Simulium (Simulium) tricorne CROSSKEY & HOWARD, 1997: 66; 2004: 60; COSCARÓN et al., 1999: 580.
Female: Wing length 3.4 mm. Body blackish. Scutum black with dark golden and
blackish pilosity lyre-shaped, scutum hair of different sizes. Abdomen blackish. Legs
black, with coxa, anterior side of fore tibia, basal 1/2 of tibia and middle basitarsus, basal
1/2 of tibia, hind basitarsus and second tarsomere yellowish; posterior basitarsus wide,
ratio of length/width = 6.5. Sensory vesicle of palp 1/4 length of palpomere. Frons
convergent. Basal portion of cibarium smooth. Cerci wide, with truncate distal edge;
paraprocts 1/2 length of basal width; sternite VIII with about 17-22 setae; gonapophyses
divergent below, with internal margin more sclerotized.
Male: Wing length 3.1 mm. Body blackish. Scutum with 1+1 cuneiform spots
indented toward middle, lateral margin edged by silvery band; abdomen black, with 1+1
silver spots on tergite II, IV-VI. Ratio of length/width of posterior basitarsus = 3.7.
Gonostylus subcylindrical, with short internal-basal process apically smooth and shorter
than transverse diameter of gonostylus; ventral plate about as long as wide with short
median process edged by submedian depressions; basal branches very robust (Fig. 106 E).
Pupa: Cocoon basal length 3.0-3.5 mm, dorsal length 2.8-3.0 mm, with anterior edge
reinforced, projected anteroventrally, tissue dense and threads conspicuous, general aspect
as in figure 107 H. Frontoclypeus and exposed portion of thorax with sparse, small,
irregularly arranged tubercles, superficially with microscopic spiculae as seen in S. sandyi.
Thorax with about 25-40 simple trichomes, 2-3 of median anterior area about twice as
large as others. Gills membranous, with 3 strong subcylindrical branches. Abdominal
tergites I and II with flat, multicuspidate tubercles.
Larva: Length 6.0-6.2 mm. Cephalic apotome with positive spots arranged in 2 groups:
basal-median longitudinal spot and 1+1 submedian spots. Postgenal cleft subquadrate to
rounded. Ratio of hypostomium/postgenal bridge = 0.7; hypostomium with 7-8 lateral
setae and 5-6 lateral serrations on each side. Cephalic fan with 26-32 primary rays. Ratio of
514 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

antennal segments proximal, medial and distal = 1:1.3:1.2. Mandible with 10-12 inner teeth
in 2 series, small platform present below median tooth. Thoracic proleg sclerite with 27
teeth. Anal sclerite without hair or scales; anal ring with 76-78 rows of 12-14 hooks each;
rectal papillae with 9-12 lobules each (DALMAT, 1955: 239).
Distribution: Guatemala: Chimaltenango, Guatemala, Huehuetenango, San Mar-
cos, Solola, Totonicapán; Mexico: Chiapas.
Bionomics: Unknown.
Discussion: The pupal gill with three branches is similar to that of S. kompi, but the
branches are all orientated in same direction in S. kompi, permitting differentiation.

Simulium (Aspathia) wygoi COSCARÓN, IBAÑEZ BERNAL & COSCARÓN ARIAS


(Figs. 101 K; 102 G-I; 103 I, J; 105 K; 106 C; 107 F, G; 108 I; 111 H-K; 112 G, H.)
Simulium (Simulium) wygoi COSCARÓN et al., 1999: 581; 582; CROSSKEY & HOWARD, 2004: 60.
Female (pharate): Scutum dark brown, with 1+1, anterior light grayish spots. Frons
convergent below (Fig. 102 H); fronto-ocular triangle deep. Sensory vesicle of palp large,
about 1/2 length of palpomere (Fig. 101 K). Cibarium scarcely elevated medially, bearing
spherules (Fig. 102 G). Basal section of R bare. Tarsal claw with small subbasal tooth (Fig.
102 I); ratio of length/width of hind basitarsus = 6.5; hair of legs filiform. Thorax with
decumbent hair. Stemite VIII with about 25 hairs on each side; gonapophyses with parallel
sides on inner margin and rounded distally (Fig. 103 I); paraproct about as long as wide, not
acuminate, but with small process on apex (Fig. 103 J); spermatheca with rows of spiculae.
Male (pharate): Sensory vesicle about 1/3 length of palpomere. Hind basitarsus
wide, ratio of length/width = 3.6. Gonocoxite as in figure 105 K; gonostylus subcylindri-
cal, with internal-basal process relatively short, covered with acuminate denticles; ventral
plate wide, subquadrate, with small, curved median process (Fig. 106 C); paramere with
robust hooks.
Pupa: Cocoon basal length 5.0 mm, dorsal length 3.3 mm, body length 4.4 mm, gill
length 3.0 mm. Cocoon flattened, translucent, subcircular in dorsal view, thorax free
dorsally, fused anteriorly into ring, anterior edge reinforced along thoracic edge and
enlarged anteriorly at gill insertion (Fig. 107 F). Frontoclypeus and thorax with abundant
tubercles and conspicuously rugose (Fig. 107 G), similar to S. parnassum MALLOCH. Frontal
(2+2) and facial (1+1) trichomes simple; thorax with 5+5 simple trichomes. Gill with
short stem and 2 main branches with 9-10 smaller, and flexible branches: 3-4 branches on
stem, and 3 on each large branch, some can be subdivided into 2 or 3 branches, gill base
with sclerotized apodeme (Fig. 108 I). Abdominal tergite II with stout setae resembling
small hooks. Tergite V with 4 setae, the largest on margin, without anterior scales; hooks
on sternites V-VII large, bifid.
Larva: Cephalic apotome with negative spots and slightly darkened mediobasally
(Fig. 112 H). Postgenal cleft scarcely developed (Fig. 112 G). Ratio of hypostomium/
postgenal bridge = 0.7-0.9; hypostomium with 5-8 lateral setae and 4+4 lateral serrations
on each side. Antenna as in figure 111 H; ratio of proximal, medial and distal segments =
1:1.3-1.4:1.1-1.2. Mandible with 8-10 inner teeth and 2 lateral mandibular processes (Fig.
111 I), second marginal tooth small and variably shaped (Fig. 111 J). Cephalic fan with 38-
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 515

A M

mm
B

m
0.05 m
0.05

0.05 mm
0.05 m
m
I
G

J
C

0.02 mm

N
F

E
m
2m
0.0

K
0.1 mm
D

Figure 111. Larva. A-G: S. (Aspathia) putre; A: dorsal view of head; B: antenna; C: Portion of
antenna under high magnification; D: dental portion of mandible; E: comb of thoracic proleg
sclerite; F: anal sclerite; G: rectal papillae. H-K: S. (A.) wygoi; H: antenna; I: dental portion of
mandible; J: mandibular marginal teeth; K: rectal papillae. L: S. (A.) marquezi, antenna. M: S. (A.)
metallicum, antenna. N: S. (A.) iriartei, antenna.
516 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

0.1 mm

A
D

G
B
0.1 mm

F
mm
0.1

C
0.1 mm

Figure 112. Larva. A-D: postgenal cleft and hypostomium; A: S. (Aspathia) marquezi. B: S. (A.)
iriartei. C: S. (A.) metallicum. D: S. A(.) putre. E-F: S. (A.) putre; E: hypostomium anterior edge; F:
general structure of larva. G-H: S. (A.) wygoi; G: postgenal cleft and hypostomium; H:
frontoclypeus.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 517

42 primary rays. Thoracic proleg sclerite with 27-28 teeth. Anal sclerite with sparse hair
and few scales; anal ring with 80 rows of 13-14 hooks each; rectal papillae with 13-15
diverticula each, one larger than others on each lobe (Fig. 111 K).
Distribution: Mexico: Distrito Federal.
Bionomics: Unknown.
Discussion: This species is similar to S. parnassum MALLOCH, based on the clypeal
and thoracic rugosity of the pupa. The gill morphology is similar to that of S. hechti but S.
hechti has 3 main branches.

Simulium (Hemicnetha) ENDERLEIN (Figs. 113-117; 151.)


Hemicnetha ENDERLEIN, 1934a: 190 (as genus).
Type-species: Simulium paynei (VARGAS) 1942a: 246, new name for Hemicnetha mexicana ENDERLEIN,
1934a: 190.
Dyarella VARGAS et al., 1946: 105, as subgenus of Simulium. Type-species: Simulium mexicanum
BELLARDI, 1862: 6.
Female: Species medium-sized to large, wing length 2.6-4.5 mm. Frons relatively
wide, convergent, with gray pollinosity. Fronto-ocular triangle deep. Scutum brown to
blackish, pollinose, frequently with 1+1 submedian longitudinal grayish stripes appearing
as light lyre shape; tonality of vittae changing with different illumination; pilosity
homogeneous to grouped and simulating scales. Abdomen brownish. Cibarium basally
smooth, medially with well sclerotized cornuae, with or without small spiculae, as also
central area which sometimes appear over pharyngeal membrane. Sensory vesicle of
maxillary palp big, frequently half of palpomere length. Sc with hair, sometimes reduced
only to basal area. Basal sector of R generally without hair. Legs without petaloid hair.
Calcipala well developed; claws with big subbasal tooth. Eighth sternite with median
depression. Gonapophysis well developed; frequently 2 times longer than wide, subtrian-
glular to subtrapezoidal with abundant microtrichiae and internal edge well sclerotized;
big paraprocts subquadrangular, longer than wide, with distal edge straight to slightly
curved or rounded; cerci mostly with distal edge flattened. Genital fork robust with
anterior conical acuminate apodemes on lateral branches. Spermatheca subtrapezoidal
with light neck connecting with spermathecal duct.
Male: Coloration as in female but darker. Gonocoxite wider than high; gonostylus
generally 2 times longer than gonocoxite; gonostylus flattened with median lateral
protuberances, generally 2 times longer than wide, rounded apically and with small subapical
spur. Ventral plate wider than high with keel projected apically as median process to about as
long as wide or flattened laterally. Endoparameres with big bases and strong teeth.
Pupa: Cocoon with compact weave, generally with anterior edge elevated, shoe
shaped, sometimes with festoons protecting gills. Gills relatively thick, branching close to
base with short basal trunk, filaments in number 6-90, externally frequently with striated
annulations and spiculae, with rounded to acute apices. Frontoclypeus very wide in males
with basal reinforcement over facial trichomes. Frontoclypeus and exposed portion of
thorax generally without tubercles, but present and very small on facial area and posterior
edge of thorax. Cephalic and thoracic trichomes small and generally single in number,
518 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

with 2+2 frontal, 1+1 facial and 5+5 or 6+6 thoracic. Distal abdominal tergites without
spine comb and no terminal spur.
Larva: Body wider distally and flattened posterodorsally. Cephalic apotome with
punctiform spots median and sublateral generally not evident. Hypostomium generally with
frontal edge curved, with median tooth longer than lateral teeth. Mandible with anterior
internal teeth big and showing several rows of teeth; marginal teeth in number of 1-2 with
second tooth very small. Antenna with medial article bigger than proximal and distal. Anal ring
with 200-500 rows with 22-73 hooks each. Rectal papillae with 5-41 diverticulae on each lobe.
Simulium paynei was created by VARGAS for Hemicnetha mexicana ENDERLEIN because
the latter corresponded to the genus Simulium and is a homonym of Simulium mexicanum
BELLARDI, which also is a Simulium (Hemicnetha) species. Simulium (Hemicnetha) consists of
23 species in 4 species groups (COSCARÓN, 1987; CROSSKEY & HOWARD, 1997, 2004) plus
another one not included. The subgenus is widely distributed in Central America, reaching
the Nearctic Region with 4 species and South America with 7 species.
The subgenus Simulium (Hemicnetha) is morphologically similar to some other
Simulium taxa, showing relationships not only with the Neotropical fauna, including
Hearlea, Trichodagmia and Thyrsopelma, but also the Afrotropical fauna such as Freemaniellum,
Anasolen and Xenosimulium (MIRANDA ESQUIVEL & COSCARÓN, 2001, 2004), suggesting
Gondwanian ancestors.
There are two species from northern Venezuela, Simulium oviedoi and Simulium rivasi
that were included in Hemicnetha (RAMÍREZ PÉREZ, 1971). These species have common
characters with Hemicnetha, such as the smooth cibarium, deep fronto-ocular triangle,
absence of hair on the basal sector of R, shape of the pupal cocoon and gill and
reinforcement on the frontoclypeal base, characters that also are similar in Hearlea.
Simulium oviedoi and Simulium rivasi have characters that differ from the known HEMIC-
NETHA species groups, such as the gonapophysis acute apically and about as long as
wide at the base (Fig. 118 C), subtriangular paraprocts (Fig. 118 C), gonostylus shorter
than the gonocoxite, with slight distal curvature (Figs. 118 F, H), ventral plate subtriangu-
lar, without a median carina (Figs. 118 E, G) (closer to Ectemnaspis species) and larval body
with dorsal curvature subterminally (Fig. 119 H ), which differs from the characteristic flat
shape typical of Hemicnetha. These differences from Hemicnetha are considerable, but this
subgenus offers the closest fit. In order to facilitate the arrangement of the Simulium
Neotropical species, the OVIEDOI species group is included in Hemicnetha.

The Simulium (HEMICNETHA) species groups are characterized as follows:


PAYNEI species group: gonapophysis 2 times longer than wide, blunt apex with
abundant microtrichiae and not sclerotized internally (Fig. 113 C); paraproct subrectangu-
lar, with anterior edge not concave. Gonostylus 2.1-2.2 times longer than wide at base,
lateral sides with slight protuberances (Fig. 114 A); ventral plate with moderate body
length and thick median process with abundant hair (Fig. 114 B). Cocoon well elevated
anteriorly with collar of festoons (Figs. 115 A-D); frontoclypeus and exposed portion of
thorax without tubercles, only diminutive tubercles on facial area; thorax dorsally generally
with gelatinous cover of variable disposition according to species (Figs. 115 L-N).
Postgenal bridge about as long as hypostomium. Gills with 8 branches. Hypostomium
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 519

frontal edge curved (Fig. 117 N). Anal ring with about 220-360 rows of 40 hook each:
Simulium paynei, Simulium rubrithorax, Simulium biuxinisa, Simulium virgatum, Simulium bricenoi.
BRACHYCLADUM: Gonapophysis subtriangular, acute apex, nearly 2 times longer
than width with sparse microtrichiae, moderate number of hairs and well sclerotized along
inner margin (Fig. 113 Q); paraprocts membranous without microtrichiae and low number
of hairs, apex rounded with small concavity posteriorly (Fig. 113 M). Cibarium with
median area bearing small teeth (Fig. 113 J). Gonostylus relatively short with lateral
protuberances, ratio of length/width about 1.5. Cocoon with aperture elevated, without
festoons; frontoclypeus and thorax without tubercles, except on facial area; gill with 6-15
branches. Hypostomium anterior edge straight with median teeth generally of same length
or longer than lateral teeth (Fig. 117 Q). Postgenal bridge longer than hypostomium. Anal
ring with 134-190 rows with 25-28 hooks each. Simulium brachycladum, Simulium pulverulen-
tum, Simulium cristalinum, Simulium guerrerense, Simulium lobatoi, Simulium solarii.
MEXICANUM: Large variation in characters. Gonapophysis about as long as wide at
base, up to 2 times longer than wide, distally acute to blunt (Figs. 113 K, V). Paraproct
subrectangular, with abundant microtrichia and no anterior concavity (Fig. 113 R). Gonostylus
about 3 times longer than width with slight lateral protuberances (Fig. 114 I); ventral plate
about as long as wide, without median process but flattened laterally and with high keel (Fig.
114 J). Cocoon with aperture moderately to well elevated, no festoons; frontoclypeus and
thorax with or without tubercles. Gill with 8-90 branches. Postgenal bridge frequently shorter
than hypostomium; hypostomium front slightly curved with median teeth higher than lateral
teeth. Postgenal cleft apex sharp. Anal ring with 200-550 rows with 27-55 hooks. Simulium
mexicanum, Simulium seriatum, Simulium smarti, Simulium freemani, Simulium earlei, Simulium keenani,
Simulium yepocapense, Simulium hinmani, Simulium hieroglyphicum.
OVIEDOI: Eighth sternite well sclerotized on the center; gonapophysis subtriangu-
lar, relatively short with acute apex (Fig. 118 C) ratio of length/width at base = 0.7-0.9.
Paraprocts subtriangular, slightly shorter than wide at base (Figs. 118 A, B). Genital fork
with anterolateral process strong and well sclerotized (Fig. 118 D). Male ratio of length/
width of hind basitarsus = 4. Gonostylus shorter than gonocoxite subcylindrical with
slight distal curvature and apical spur; ventral plate about as long as wide at base without
any keel or lateral entrances (Figs. 118 E, G); endoparameres with strong teeth. Cocoon
shoe shaped, relatively low anteriorly with reinforced anterior edge (Fig. 119 A).
Frontoclypeus reinforced basally and very wide in males, nearly same shape as female,
frontoclypeus and thorax with abundant granulose tubercles (Figs. 119 B, E), frequently
very acute (Fig. 119 D) and multibranched trichomes (Fig. 119 G). Gill with 6 thick
relatively short branches emerging form short and thick trunk (Figs. 119 C, F). Abdomen
with apical spur small. Larval body with slight dorsal subterminal curvature and without
ventral tubercles (Fig. 119 H). Cephalic apotome with positive spots. Antennae short and
robust, only reaching half length of cephalic fan stem; medial article longer than others.
Cephalic fan with 20-25 rays with thick teeth all of similar length (Fig. 119 J). Mandible
with 2 marginal teeth of similar size (Fig. 119 K). Postgenal bridge as long as
hypostomium length. Lateral sclerite of thoracic proleg with 32-45 teeth (Fig. 119 N).
Anal sclerite with abundant hair, but no scales. Anal ring with 100-120 rows with 17-18
hooks each. Rectal papillae with 20-26 diverticula on each lobe.
520 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

KEYS TO SIMULIUM (HEMICNETHA) SPECIES

Females
1 Gonapophysis shorter than width at base (Fig. 118 C); paraprocts subtriangular
and apically blunt (Figs. 118 A, B) .............................Simulium oviedoi, Simulium rivasi
– Gonapophysis longer than width at base (Figs. 113 C, K, Q, T, V); paraprocts
subrectangular and apically rounded or curved (Figs. 113 C, F, L, M, R, U) ....... 2
2(1) Scutum without evident longitudinal stripes; gonapophysis subtrapezoidal, with
blunt or rounded apex, relatively short (Fig. 113 Q) ratio of length/width at base
= 1.0-1.1 ............................................................................................................................... 3
– Scutum with 1+1 longitudinal light gray submedian stripes, generally joined
posteriorly, forming lyre shape (Figs. 113 D, G), with different illumination,
color changes from silvery to blackish; gonapophysis generally subtriangular
and acuminate apically (Figs. 113 C, K.); ratio of length/width at base = 1.3-3.0
................................................................................................................................................ 4
3(2) Wing length 4.0-4.4 mm; basal portion of cibarium with spiculae on median
portion ..........................................................................................................Simulium smarti
– Wing length 3.1-3.4 mm; basal portion of cibarium without spiculae on median
portion (Fig. 113 B) .......................................... Simulium mexicanum, Simulium seriatum
4(2) Gonapophysis subtrapezoidal, with blunt apex (as Fig. 113 Q) .............................. 5
– Gonapophysis subtrapezoidal, with acute (Fig. 113 K) to blunt apex (Fig. 113 C)
................................................................................................................................................ 7
5(4) Gonapophysis relatively short, ratio of length/width at base = 1.0; paraproct with
apical edge mostly straight ................................................................... Simulium freemani
– Gonapophysis relatively long, ratio of length/width at base = 1.8-2.7; paraproct
with apical edge curved .................................................................................................... 6
6(5) Scutum hair arranged homogeneously; ratio of length of palpomere/sensory
vesicle = 2.3 ..............................................................................................Simulium keenani
– Scutum hair arranged in groups; ratio of length of palpomere/sensory vesicle =
1.9 ..................................................................................................... Simulium hieroglyphicum
7(4) Gonapophysis with apex acute (Figs. 113 K, V) ......................................................... 8
– Gonapophysis with apex blunt (Fig. 113 C) ............................................................... 15
8(7) Paraproct membranous on posterior 3/4, without microtrichia and with sparse
setae only on edge .............................................................................. Simulium cristalinum
– Paraproct with variable sclerotization on posterior 3/4, but covered by
microtrichia and relative abundance of setae (Figs. 113 L, M) ................................ 9
9(8) Small species, wing length generally less than 2.8 mm; ratio of palpomere length/
sensory vesicle = 1.6-1.8 ................................................................................................ 10
– Medium to big species, wing length 2.8-4.3; ratio of palpomere length/sensory
vesicle = 2.2-2.5 ................................................................................................................ 13
10(9) Sc with hair at least on basal 3/4 of vein ................................................................... 11
– Sc with hair only on basal 1/4 of vein ........................................................................ 12
11(10) Scutum with submedian vittae joining posteriorly, forming lyre shape (Fig. 113 G);
wing length 2.8-3.3 mm ................................................................ Simulium brachycladum
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 521

– Scutum with submedian vittae extending only to 2/3 of scutum, not joining
posteriorly; wing length 2.4-2.9 mm ..................................................... Simulium lobatoi
12(10) Scutum dark brownish, without ornamentation changing in different illumination
....................................................................... Simulium pulverulentum, Simulium guerrerense
– Scutum brown orange, with submedian vittae forming lyre shape in some
illumination .................................................................................................. Simulium solarii
13(9) Big species, wing length 4.0-4.3 mm ...................................................Simulium hinmani
– Medium-sized species, wing length 2.4-2.9 mm ........................................................ 14
14(13) Sc with hair all along vein ......................................................................... Simulium earlei
– Sc with hair only on basal half of vein ......................................... Simulium yepocapense
15(7) Without hair on Sc; ratio of gonapophysis length/width = 2.2 ..........................
.................................................................................................................... Simulium bricenoi
– With hair on Sc; ratio of gonapophysis length/width = 1.4-1.6 ........................... 16
16(15) Gonapophysis with inner edge bearing microtrichia ..................... Simulium biuxinisa
– Gonapophysis with inner edge not bearing microtrichia ............................................
................................................. Simulium paynei, Simulium rubrithorax, Simulium virgatum

Males
Not included: Simulium biuxinisa (unknown) and S. keenani (lack of characters).

1 Gonostylus shorter than gonocoxite (Figs. 118 F, H); ventral plate subtriangular
and without carina (Figs. 118 E, G) ...........................Simulium oviedoi, Simulium rivasi
– Gonostylus longer than gonocoxite (Figs. 114 A, E, I, M, N); ventral plate not
subtriangular and with carina (Figs. 114 B, C, F, H, J, L, O) .................................... 2
2(1) Ventral plate with strong median distal projection (Figs. 114 B, C, F, L) .............. 3
– Ventral plate without strong median distal projection (Figs. 114 J, O) ................ 10
3(2) Ratio of length of gonostylus/gonocoxite = 1.4-1.8 ................................................ 4
– Ratio of length of gonostylus/gonocoxite = 1.9-2.2 ................................................ 6
4(3) Ratio of gonostylus length/width = 1.3-2.0 ................................................................ 5
– Ratio of gonostylus length/width = 2.4-2.9 ............................... Simulium rubrithorax
5(4) Ventral plate body without 1+1 concavities on posterior edge (Fig. 114 F), ratio of
gonostylus length/width = 1.7 .................................................... Simulium brachycladum
– Ventral plate body with 1+1 deep concavities on posterior edge, ratio of
gonostylus length/width = 1.8-2.1 ........................... Simulium lobatoi, Simulium solarii
6(3) Ratio of hind basitarsus length/width = 3.8-5.6 ........................................................ 7
– Ratio of hind basitarsus length/width = 5.7-7.2 ........................................................ 9
7(6) Gonostylus with slight outer lateral protuberances; ratio of length of gonostylus/
gonocoxite = 1.6 ........................................................................................ Simulium paynei
– Gonostylus with well-developed outer lateral protuberances; ratio of length of
gonostylus/gonocoxite = 1.8-1.9 .....................................................................................
8(7) Ventral plate body with 1+1 submedian concavities posteriorly (Fig. 114 C);
scutum with dark pollinose stripes not reaching anterior edge; ratio of length/
width of hind basitarsus = 5.6 ............................................................ Simulium virgatum
522 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

– Ventral plate body straight posteriorly without submedian concavities; scutum


with dark pollinose stripes reaching anterior edge; ratio of length/width of hind
basitarsus = 3.8-4.0 .................................................................................. Simulium briceroi
9(6) Ratio of hind basitarsus length/width = 5.7 ........................... Simulium pulverulentum
– Ratio of hind basitarsus length/width = 7.2 ............................... Simulium cristalinum
10(2) Ventral plate body narrow, flattened laterally, appearing as axe in lateral view ......... 11
– Ventral plate body not narrow or flattened laterally ................................................ 13
11(10) Ratio of length of gonostylus/gonocoxite = 2.0; ratio of gonostylus length/
width = 2.3 ..................................................................................... Simulium hieroglyphicum
– Ratio of length of gonostylus/gonocoxite = 2.2-2.3; ratio of gonostylus length/
width = 2.4-2.9 ................................................................................................................. 12
12(11) Ratio of hind basitarsus length/width = 3.3 .......................................Simulium smarti
– Ratio of hind basitarsus length/width = 3.8 ................................... Simulium freemani
13(10) Ratio of hind basitarsus length/width = 7.2 ................................Simulium guerrerense
– Ratio of hind basitarsus length/width = 3.1-5.0 ...................................................... 14
14(13) Wing length 4.2-4.4 mm; ratio of hind basitarsus length/width = 5.0 ....................
.....................................................................................................................Simulium hinmani
– Wing length 2.7-3.4 mm; ratio of hind basitarsus length/width = 3.1-4.8 ......... 15
15(14) Ratio of gonostylus length/width 2.3-2.5 .................................................................. 16
– Ratio of gonostylus length/width 1.9/2.1 ................................................................. 17
16(15) Wing length 2.7-2.9 mm ....................................................................... Simulium seriatum
– Wing length 3.0-3.3 mm ................................................................... Simulium mexicanum
17(15) Wing length 3.1 mm; ratio of gonostylus length/width = 2.1 ......... Simulium earlei
– Wing length 3.4 mm; ratio of gonostylus length/width = 1.9 Simulium yepocapense

Pupae
1 Cocoon with festoons edging entire aperture (Figs. 115 A-D) ................................ 2
– Cocoon without festoons edging entire aperture or festoons very reduced (Figs.
115 E-J) ................................................................................................................................ 6
2(2) Frontoclypeus and thorax without tubercles; thorax wrinkled (Fig. 116 E) ...........
...................................................................................................................... Simulium bricenoi
– Frontoclypeus and thorax with tubercles (Fig. 115 Q); thorax smooth or wrinkled
................................................................................................................................................ 3
3(2) Festoons of anterior edge of aperture joined with thin weave; gills not surpassing
height of festoons (Fig. 115 B) .......................................................... Simulium biuxinisa
– Festoons of anterior edge of aperture free, without intermixed weave; gills
surpassing or not height of festoons ............................................................................. 4
4(3) Without gelatinous material on thoracic dorsum ............................ Simulium virgatum
– With thick gelatinous material on thoracic dorsum (Figs. 115 L-N) ...................... 5
5(4) Thoracic thick gelatinous material with lateral entrances appearing as 4 branches
(Fig. 115 M); frontoclypeus uniformly light brown ............................ Simulium paynei
– Thoracic thick gelatinous material with entrance only on median longitudinal line (Fig.
115 L); frontoclypeus darker on anterior area (Fig. 115 O) .......... Simulium rubrithorax
6(1) Gill with 6 branches .......................................................................................................... 7
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 523

– Gill with 8 or more branches ........................................................................................ 10


7(6) Cocoon very elevated anteriorly (Fig. 115 E); frontoclypeus and exposed portion
of thorax without tubercles or with only sparse tubercles ... Simulium brachycladum
– Cocoon medially elevated anteriorly (Figs. 115 F; 119 A); frontoclypeus and
exposed portion of thorax with abundant tubercles (Figs. 119 B, E) or sparse and
very dark with much sclerotization (Fig. 115 S) .......................................................... 8
8(7) Gills relatively short (Fig. 116 G), length 0.7-0.8 mm; frontoclypeus with tubercles
only on base (Fig. 115 S) .................................................................. Simulium cristalinum
– Gills relatively long (Figs. 119 C, F), length 1.4-3.0 mm; frontoclypeus with
abundant tubercles (Figs. 119 B, E) ............................................................................... 9
9(8) Frontoclypeal and thoracic tubercles acuminate, like spines (Fig. 119 D); gill
length 1.4-1.7 mm ..................................................................................... Simulium oviedoi
– Frontoclypeal and thoracic tubercles not acuminate, granulose (Fig. 119 G); gill
length 2.8-3.0 mm ....................................................................................... Simulium rivasi
10(6) Gill with 8 branches ........................................................................................................ 11
– Gill with more than 10 branches .................................................................................. 12
11(10) Frontoclypeus with moderate abundance of tubercles on frontal area, gills free;
gill length 2.0-2.1 mm ........................................................................... Simulium freemani
– Frontoclypeus without tubercles on frontal area, gills protected by cocoon; gill
length 0.9-1.8 mm ..................................................................................... Simulium lobatoi
12(10) Gill with 10 branches ...................................................................................................... 13
– Gill with more than 10 branches .................................................................................. 15
13(12) Cocoon not covering gills ............................................................ Simulium pulverulentum
– Cocoon covering gills ...................................................................................................... 14
14(13) Cocoon aperture cylindrical with same height anteriorly and posteriorly; cocoon
length basally 2.3-2.7 mm, gills 0.8-1.0 mm; thoracic trichomes single ...................
................................................................................................................ Simulium guerrerense
– Cocoon aperture more elevated anteriorly than posteriorly; cocoon length basally
3.5-4.0 mm, gills 1.4-1.5 mm; thoracic trichomes with 2-5 branches .......................
................................................................................................................ Simulium yepocapense
15(12) Gill with 12 branches ...................................................................................................... 16
– Gill with more than 12 branches .................................................................................. 17
16(15) Exposed portion of thorax with tubercles arranged homogeneously (Fig. 116 H);
frontoclypeus with abundant tubercles ......................................... Simulium mexicanum
– Exposed portion of thorax arranged in 2+2 longitudinal stripes joined anteriorly
(Figs. 116 I-J); frontoclypeus without abundant tubercles or with very sparse
tubercles (Fig. 115 T) ............................................................................ Simulium seriatum
17(15) Gill with 15 branches ..................................................... Simulium solarii, Simulium earlei
– Gill with 16 or more branches ...................................................................................... 18
18(17) Gill with 16 branches .................................................. Simulium earlei, Simulium keenani
– Gill with 18 or more branches ...................................................................................... 19
19(18) Gill with 18 branches; thorax not dorsally wrinkled .......................... Simulium smarti
– Gill with 60-90 branches (Fig. 116 L); thorax dorsally wrinkled ...............................
........................................................................................................... Simulium hieroglyphicum
524 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Larvae
Not included: Simulium freemani, S. keenani and S. lobatoi.

1 Postgenal bridge longer than hypostomium length (Fig. 117 P) .............................. 2


– Postgenal bridge shorter or same as hypostomium length ....................................... 7
2(1) Hypostomium frontal edge almost straight, with median tooth at same level as
lateral teeth; postgenal cleft arc shaped apically (Fig. 117 P) ................................... 3
– Hypostomium frontal edge convex, with median tooth extended beyond level of
lateral teeth (Fig. 117 N); postgenal cleft with arc to acute shape apically ........... 6
3(2) Anal ring with about 134 rows of hooks; larva maximum length 4.5-4.8 mm ......
............................................................................................................ Simulium pulverulentum
– Anal ring with about 160-220 rows of hooks; larva maximum length 6.0-9.0 mm
................................................................................................................................................ 4
4(3) Cephalic apotome with negative spots (Fig. 117 F); rectal papillae with 5
diverticula on each lobe ................................................................ Simulium brachycladum
– Cephalic apotome with positive spots; rectal papillae with 6-9 diverticula on each
lobe ........................................................................................................................................ 5
5(4) Anal ring with 190-220 rows, and 28-30 hooks each; maximum larva length 9.0
mm ................................................................................................................. Simulium earlei
– Anal ring with 160 rows, and 22 hooks each; maximum larva length 5.8 mm .......
.................................................................................................................Simulium guerrerense
6(2) Postgenal cleft subtriangular with acute apex; anal ring with 500-550 rows of
hooks; maximum larva length about 13.2 mm ....................... Simulium hieroglyphicum
– Postgenal cleft arc shaped with blunt apex; anal ring with 225 rows of hooks;
maximum larva length about 8.0 mm .................................................Simulium hinmani
7(1) Maximum larva length 10.5-12.6 mm; anal ring with 280-455 rows of hooks ........ 8
– Maximum larva length 6.0-9.5 mm; anal ring with 100-320 rows of hooks ....... 10
8(7) Postgenal bridge as long as hypostomium length (Fig. 117 O) ........ Simulium biuxinisa
– Postgenal bridge shorter than hypostomium length (Figs. 117 N, R) .................... 9
9(8) Anal ring with 450-460 rows of hooks; rectal papillae with 17-20 diverticula on
each lobe ......................................................................................................Simulium smarti
– Anal ring with 280-346 rows of hooks; rectal papillae with 9-15 diverticula on
each lobe ................................................................ Simulium paynei, Simulium rubrithorax
10(7) Anal ring with 100-170 rows of hooks ....................................................................... 11
– Anal ring with 210-320 rows of hooks ....................................................................... 13
11(10) Antennae with thick articles (Fig. 119 I); second marginal tooth as long as first
(Fig. 119 K) ...................................................................... Simulium oviedo, Simulium rivasi
– Antennae with articles relatively thin; second marginal tooth less than half as long
as first marginal tooth (Fig. 117 L) ............................................................................... 12
12(11) Maximum larva length 5.8-6.3 mm; cephalic apotome with negative spots (Fig.
117 G); postgenal bridge as long as hypostomium length (Fig. 117 Q); ratio of
proximal, medial and distal antennal articles = 1:1.6-1.7:1.0 ................................
................................................................................................................ Simulium cristalinum
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 525

– Maximum larva length 8.2-8.9 mm; cephalic apotome with positive spots;
postgenal bridge shorter than hypostomium length; ratio of proximal, medial and
distal antennal articles = 1: 1.2:0.8 ......................................................... Simulium solarii
13(10) Postgenal bridge as long as hypostomium length (Fig. 117 N) .............................. 14
– Postgenal bridge shorter than hypostomium length (Fig. 117 R) .......................... 15
14(13) Cephalic apotome with positive spots; rectal papillae with 10-17 diverticula on
each lobe .............................................................................................. Simulium mexicanum
– Cephalic apotome with negative spots; rectal papillae with 7-8 diverticula on each
lobe ............................................................................................................ Simulium virgatum
15(13) Mandible with only 1 big marginal tooth (Fig. 117 M); maximum larva length 6.5-
7.0 mm ...................................................................................................... Simulium seriatum
– Mandible with 1 big marginal tooth and another small tooth; maximum larva
length 7.1-10.7 mm .......................................................................................................... 16
16(15) Maximum larva length 9.1-10.7 mm; ratio of proximal, medial and distal antennal
articles = 1: 1.7:1.2 ................................................................................... Simulium bricenoi
– Maximum larva length 7.0 mm; ratio of proximal, medial and distal antennal
articles = 1:1.9-2:1.4-1.5 ................................................................... Simulium yepocapense

CLAVES DE ESPECIES DE S. (HEMICNETHA)


Hembras
1 Gonapófisis más cortas que anchas en la base (Fig. 118 C); paraproctos
subtriangulares y apicalmente romos (Figs. 118 A, B) .................................................
............................................................................................Simulium oviedoi, Simulium rivasi
– Gonapófisis más largas que anchas en la base (Figs. 113 C, K, Q, T, V);
paraproctos subrectangulares y apicalmente redondeados o curvados (Figs. 113 C,
F, L, M, R, U) ...................................................................................................................... 2
2(1) Escudo sin bandas longitudinales bien evidentes; gonapófisis subtrapezoidales
con ápice romo o redondeado y relativamente cortas (Fig. 113 Q); relación largo/
ancho en la base = 1,0-1,1 ............................................................................................... 3
– Escudo con 1+1 bandas submedianas gris claro, generalmente unidas
posteriormente en forma de lira (Figs. 113 D, G), con diferente iluminación
cambia el plateado por coloración negruzca; gonapófisis generalmente con forma
subtriangular y aguzadas apicalmente (Figs. 113 C, K); relación del largo/ancho en
la base = 1,3-3,0 ................................................................................................................. 4
3(2) Largo ala 4,4 mm; porción basal del cibario con espículas sobre la porción
mediana ........................................................................................................Simulium smarti
– Largo ala 3,1-3,4 mm; porción basal del cibario sin espículas sobre la porción
mediana (Fig. 113 B) ........................................ Simulium mexicanum, Simulium seriatum
4(2) Gonapófisis subtrapezoidal con ápice romo ( como en Fig. 113 Q) ...................... 5
– Gonapófisis subtrapezoidal con ápice agudo (Fig. 113 K) hasta con ápice romo
(Fig. 113 C) .......................................................................................................................... 7
5(4) Gonapófisis relativamente cortas; relación del largo/ancho en la base = 1,0; borde
apical del paraprocto mayormente recto ........................................... Simulium freemani
526 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

– Gonapófisis relativamente largas; relación del largo/ancho en la base = 1,8-2,7;


borde apical del paraprocto curvo ................................................................................. 6
6(5) Pelos del escudo dispuestos homogéneamente; relación largo palpómero/vesícula
sensorial = 2,3 ...........................................................................................Simulium keenani
– Pelos del escudo dispuestos en grupos; relación largo palpómero/vesícula
sensorial = 1,9 ................................................................................ Simulium hieroglyphicum
7(4) Gonapófisis con ápice aguzado (Figs. 113 K, V) ........................................................ 8
– Gonapófisis con ápice romo (Fig. 113 C) ................................................................... 15
8(7) Paraprocto membranoso sobre la 3/4 porción posterior, sin microtriquias y
escasas cerdas solo sobre el borde ................................................. Simulium cristalinum
– Paraprocto con variable grado de esclerotización sobre la 3/4 porción posterior,
pero cubierta por microtriquias y abundantes cerdas (Figs. 113 L, M) .................. 9
9(8) Especies pequeñas; largo ala generalmente por debajo de 2,8 mm; relación del
largo del palpómero/vesícula sensorial = 1,6-1,8 ..................................................... 10
– Especies medianas a grandes; largo ala 2,8-4,3; relación del largo del palpómero/
vesícula sensorial 2,2-2,5 ................................................................................................. 13
10(8) Sc con pelos por lo menos en el 3/4 basal de la vena ............................................. 11
– Sc con pelos solo en el 1/ 4 basal de la vena ............................................................ 12
11(10) Escudo con 1+1 banda submediana uniéndose posteriormente dando aspecto de
lira (Fig. 113 G); largo ala 2,8-3,3 mm ...................................... Simulium brachycladum
– Escudo con 1+1 banda submediana alcanzando solamente los 2/3 del escudo; no
uniéndose posteriormente; largo ala 2,4-2,9 mm ............................... Simulium lobatoi
12(10) Escudo castaño oscuro, sin cambio de ornamentación con diferente iluminación
........................................................................ Simulium pulverulentum, Simulium guerrerense
– Escudo castaño naranja, mostrando con alguna iluminación las bandas
submedianas en forma de lira ................................................................. Simulium solarii
13(9) Especies grandes, largo ala 4,0-4,3 mm .............................................Simulium hinmani
– Especies medianas, largo ala 2,4-2,9 mm .................................................................... 14
14(13) Sc con pelos todo a lo largo de la vena .................................................. Simulium earlei
– Sc con pelos solo en la mitad basal ................................................ Simulium yepocapense
15(7) Sin pelos sobre la Sc; relación largo/ancho en la base de gonapófisis = 2,2 .........
...................................................................................................................... Simulium bricenoi
– Con pelos sobre la Sc; relación largo/ancho en la base de gonapófisis = 1,4-1,6
.............................................................................................................................................. 16
16(15) Borde interno de gonapófisis con microtriquias ............................ Simulium biuxinisa
– Borde interno de gonapófisis sin microtriquias ............................................................
................................................. Simulium paynei, Simulium rubrithorax, Simulium virgatum

Machos
Simulium biuxinisa y S. keenani no incluidas por desconocimiento de caracteres.

1 Gonostilo más corto que el gonocoxito (Figs. 118 F, H); placa ventral
subtriangular y sin carena (Fig. 118 G) .....................Simulium oviedoi, Simulium rivasi
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 527

– Gonostilo más largo que el gonocoxito (Figs. 114 A, E, I, M, N); placa ventral no
subtriangular y con carena (Figs. 114 B, C, F, H, J, L, O) ......................................... 2
2(1) Placa ventral con fuerte prolongación medio distal (Figs. 114 B, C, F, L) ............ 3
– Placa ventral sin fuerte prolongación medio distal (Figs. 114 J, O) ...................... 10
3(2) Relación largo gonostilo/gonocoxito = 1,4-1,8 .......................................................... 4
– Relación largo gonostilo/gonocoxito = 1,9-2,2 .......................................................... 6
4(3) Relación largo/ancho del gonostilo = 1,3-2,0 ............................................................. 5
– Relación largo/ancho del gonostilo = 2,4-2,9 ............................ Simulium rubrithorax
5(4) Cuerpo de placa ventral sin 1+1 concavidad sobre el borde posterior (Fig. 114 F):
relación largo/ancho del gonostilo = 1,7 ................................. Simulium brachycladum
– Cuerpo de la placa ventral con 1+1 concavidad profunda sobre el borde posterior;
relación largo/ancho del gonostilo = 1,8-2,1 ........ Simulium lobatoi, Simulium solarii
6(3) Relación largo/ancho de basitarso posterior 3,8-5,6 .................................................. 7
– Relación largo/ancho de basitarso posterior 5,7-7,2 .................................................. 9
7(6) Gonostilo con escasas protuberancias látero-externas; relación largo del
gonostilo/gonocoxito = 1,6 ..................................................................... Simulium paynei
– Gonostilo con protuberancias latero-externas bien desarrolladas; relación largo
del gonostilo/gonocoxito = 1,8-1,9 ............................................................................... 8
8(7) Cuerpo de la placa ventral con 1+1 concavidad posterior submediana (Fig. 114 C);
bandas polinosas oscuras del escudo no llegan al borde anterior; relación largo/
ancho del basitarso posterior = 5,6 .................................................... Simulium virgatum
– Cuerpo de la placa ventral recto posteriormente, sin concavidad submediana;
bandas polinosas oscuras del escudo llegan al borde anterior; relación largo/
ancho del basitarso posterior = 3,8-4,0 ............................................... Simulium bricenoi
9(6) Relación largo/ancho del basitarso posterior = 5,7 ............... Simulium pulverulentum
– Relación largo/ancho del basitarso posterior = 7,2 ................... Simulium cristalinum
10(2) Cuerpo de placa ventral angosto aplastado lateralmente semejando un hacha
antigua ................................................................................................................................ 11
– Cuerpo de placa ventral no angosto, ni aplastado lateralmente ............................. 13
11(10) Relación del largo gonostilo/gonocoxito = 2,0; relación largo/ancho de gonostilo
= 2,3 ................................................................................................. Simulium hieroglyphicum
– Relación del largo gonostilo/gonocoxito = 2,2-2,3; relación largo/ancho del
gonostilo = 2,4-2,9 ........................................................................................................... 12
12(11) Relación largo/ancho del basitarso posterior = 3,3 ...........................Simulium smarti
– Relación largo/ancho del basitarso posterior = 3,8 ....................... Simulium freemani
13(10) Relación largo/ancho del basitarso posterior = 7,2 ....................Simulium guerrerense
– Relación largo/ancho del basitarso posterior = 3,1-5,0 .......................................... 14
14(13) Largo ala 4,2-4,4 mm; relación largo/ancho del basitarso posterior = 5,0 .............
.....................................................................................................................Simulium hinmani
– Largo ala 2,7-3,4 mm; relación largo/ancho del basitarso posterior = 3,1-4,8 ......... 15
15(14) Relación largo/ancho del gonostilo 2,3-2,5 ............................................................... 16
– Relación largo/ancho del gonostilo 1,9-2,1 ............................................................... 17
16(15) Largo ala 2,7-2,9 mm ............................................................................ Simulium seriatum
– Largo ala 3,0-3,3 mm ........................................................................ Simulium mexicanum
528 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

17(15) Largo ala 3,1 mm ; relación gonostilo largo/ancho = 2,1 ................. Simulium earlei
– Largo ala 3,4 mm; relación gonostilo largo/ancho = 1,9 ......... Simulium yepocapense

Pupas
1 Capullo con festones bordeando totalmente la abertura (Figs. 115 A-D) ............. 2
– Capullo sin festones bordeando totalmente la abertura o muy reducidos (Figs. 115
E-J) ........................................................................................................................................ 6
2(1) Frontoclípeo y tórax sin tubérculos, tórax rugoso (Fig. 116 E) ........ Simulium bricenoi
– Frontoclípeo y tórax con tubérculos (Fig. 115 Q); tórax liso o rugoso ................. 3
3(2) Festones del borde anterior de la apertura unidos por un tejido fino; branquias no
sobrepasando la altura de los festones (Fig. 115 B) ...................... Simulium biuxinisa
– Festones del borde anterior de la apertura libres, sin tejido entremezclado;
branquias sobrepasando o no la altura de los festones .............................................. 4
4(3) Sin membrana gelatinosa sobre el dorso del tórax ......................... Simulium virgatum
– Con gruesa membrana gelatinosa sobre el dorso del tórax (Figs. 115 L-N) ........ 5
5(4) Tórax con gruesa membrana gelatinosa con entradas laterales presentando como
4 ramas (Fig. 115 M); frontoclípeo castaño claro homogéneo ......... Simulium paynei
– Tórax con gruesa membrana gelatinosa con una sola entrada sobre la línea media
longitudinal (Fig. 115 L); frontoclípeo más oscuro sobre el área anterior (Fig. 115
O) .......................................................................................................... Simulium rubrithorax
6(1) Branquias con 6 ramas ...................................................................................................... 7
– Branquias con 8 ó más ramas ........................................................................................ 10
7(6) Capullo muy elevado anteriormente (Fig. 115 E); frontoclípeo y porción expuesta
del tórax sin o con escasos tubérculos ...................................... Simulium brachycladum
– Capullo medianamente elevado anteriormente (Figs. 115 F; 119 A); frontoclípeo y
porción expuesta del tórax con abundantes tubérculos (Figs. 119 B, E) o muy
escasos oscuros, con gran esclerotización (Fig. 115 S) .............................................. 8
8(7) Branquias relativamente cortas (Fig. 116 G); largo 0,7-0,8 mm; frontoclípeo con
tubérculos solo en la base (Fig. 115 S) .......................................... Simulium cristalinum
– Branquias relativamente largas (Figs. 119 C, F); largo 1,4-3,0 mm; frontoclípeo
con abundantes tubérculos (Figs. 119 B, E) ................................................................. 9
9(8) Frontoclípeo y tórax con tubérculos aguzados como espinas (Fig. 119 D); largo
branquia 1,4-1,7 mm ................................................................................ Simulium oviedoi
– Frontoclípeo y tórax con tubérculos no aguzados y granulosos (Fig. 119 G); largo
branquia 2,8-3,0 mm .................................................................................. Simulium rivasi
10(6) Branquia con 8 ramas ...................................................................................................... 11
– Branquia con 10 o más ramas ....................................................................................... 12
11(10) Frontoclípeo con tubérculos de área frontal en moderada abundancia; branquias
libres; largo branquia 2,0-2,1 mm ....................................................... Simulium freemani
– Frontoclípeo sin tubérculos en área frontal; branquias protegidas por capullo;
largo branquias 0,9-1,8 mm .................................................................... Simulium lobatoi
12(10) Branquias con 10 ramas .................................................................................................. 13
– Branquias con más de 10 ramas ................................................................................... 15
13(12) Capullo no cubre las branquias ................................................... Simulium pulverulentum
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 529

– Capullo cubriendo las branquias ................................................................................... 14


14(13) Abertura del capullo en forma cilíndrica con la misma altitud anterior y posterior;
largo capullo en la base 2,3-2,7 mm; branquias 0,8-1,0 mm; tricomas torácicos
simples ...................................................................................................Simulium guerrerense
– Abertura del capullo más elevada anterior que posteriormente; largo capullo en la
base 3,5-4,0 mm; branquias 1,4-1,5 mm; tricomas torácicos con 2-5 ramas ..........
................................................................................................................ Simulium yepocapense
15(12) Branquia con 12 ramas ................................................................................................... 16
– Branquia con más de 12 ramas ..................................................................................... 17
16(15) Porción expuesta del tórax con tubérculos distribuidos homogéneamente (Fig.
116 H); frontoclípeo con abundantes tubérculos ........................ Simulium mexicanum
– Porción expuesta del tórax con tubérculos dispuestos en 2+2 bandas
longitudinales unidas anteriormente (Figs. 116 I, J); frontoclípeo sin abundantes
tubérculos o muy escasos (Fig. 115 T) .............................................. Simulium seriatum
17(15) Branquia con 15 ramas .................................................. Simulium solarii, Simulium earlei
– Branquia con 16 o más ramas ....................................................................................... 18
18(17) Branquia con 16 ramas ............................................... Simulium earlei, Simulium keenani
– Branquia con 18 o más ramas ....................................................................................... 19
19(18) Branquia con 18 ramas .............................................................................Simulium smarti
– Branquia con 60-90 ramas (Fig. 116 L); tórax dorsalmente rugoso ..........................
........................................................................................................... Simulium hieroglyphicum

Larvas
No incluidas: Simulium freemani, S. keenani y S. lobatoi.

1 Puente postgenal más largo que el hipostomio (Fig. 117 P) .................................... 2


– Puente postgenal más corto que el hipostomio .......................................................... 7
2(1) Borde frontal del hipostomio casi recto, con diente mediano al mismo nivel que el
diente lateral; ápice de hendidura postgenal con forma de arco (Fig. 117 P) ....... 3
– Borde frontal del hipostomio convexo, con diente mediano sobre el nivel del
diente lateral (Fig. 117 N); ápice de abertura postgenal con forma de arco o
aguzado ................................................................................................................................. 6
3(2) Anillo anal con alrededor de 134 hileras de ganchos; máximo largo de larva 4,5-
4,8 mm .............................................................................................. Simulium pulverulentum
Anillo anal con 160-220 hileras de ganchos; máximo largo de larva 6,0-9,0 mm
................................................................................................................................................ 4
4(3) Apotoma cefálico con manchas negativas (Fig. 117 F); papilas rectales con 5
divertículos en cada lóbulo .......................................................... Simulium brachycladum
– Apotoma cefálico con manchas positivas; papilas rectales con 6-9 divertículos en
cada lóbulo .......................................................................................................................... 5
5(4) Anillo anal con 190-220 hileras y con 28-30 ganchos en cada una; máximo largo
de larva 9,0 mm ........................................................................................... Simulium earlei
– Anillo anal con 160 hileras y con 22 ganchos en cada una; máximo largo de larva
5,8 mm ...................................................................................................Simulium guerrerense
530 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

6(2) Hendidura postgenal subtriangular, con ápice aguzado; anillo anal con 500-550
hileras de ganchos; máximo largo de larva alrededor de 13,2 mm ...........................
........................................................................................................... Simulium hieroglyphicum
– Hendidura postgenal en forma de arco con ápice romo; anillo anal con 225 hileras
de ganchos; máximo largo de larva alrededor de 8,0 mm ............. Simulium hinmani
7(1) Máximo largo de larva 10,5-12,6 mm; anillo anal con 280-455 hileras de ganchos
................................................................................................................................................ 8
– Máximo largo de larva 6,0-9,5 mm; anillo anal con 100-320 hileras de ganchos
.............................................................................................................................................. 10
8(7) Puente postgenal tan largo como el largo del hipostomio (Fig. 117 O) ..................
................................................................................................................... Simulium biuxinisa
– Puente postgenal más corto que el largo del hipostomio (Figs. 117 N, R) ........... 9
9(8) Anillo anal con 450-460 hileras de ganchos; papilas rectales con 17-20 divertículos
sobre cada lóbulo .......................................................................................Simulium smarti
– Anillo anal con 280-346 hileras de ganchos; papilas rectales con 9-15 divertículos
sobre cada lóbulo ................................................. Simulium paynei, Simulium rubrithorax
10(7) Anillo anal con 100-170 hileras de ganchos ............................................................... 11
– Anillo anal con 210-320 hileras de ganchos ............................................................... 13
11(10) Antena con artejos gruesos (Fig. 119 I); segundo diente marginal tan largo como
el primero (Fig. 119 K) .................................................Simulium oviedoi, Simulium rivasi
– Antena con artejos relativamente finos; segundo diente marginal menos de la
mitad del primer diente marginal (Fig. 117 L) ........................................................... 12
12(11) Máximo largo de larva 5,8-6,3 mm; apotoma cefálico con manchas negativas
(Fig. 117 G); puente postgenal tan largo como el hipostomio (Fig. 117 Q);
relación de los artejos antenales proximal, mediano y distal = 1:1,6-1,7:1,0 .......
................................................................................................................ Simulium cristalinum
– Máximo largo de larva 8,2-8,9 mm; apotoma cefálico con manchas positivas;
puente postgenal más corto que el hipostomio; relación de los artejos antenales
proximal, mediano y distal = 1:1,2:0,8 .................................................. Simulium solarii
13(10) Puente postgenal tan largo como el hipostomio (Fig. 117 N) ............................... 14
– Puente postgenal más corto que el hipostomio (Fig. 117 R) ................................. 15
14(13) Apotoma cefálico con manchas positivas; papilas rectales con 10-17 divertículos
sobre cada lóbulo ............................................................................... Simulium mexicanum
– Apotoma cefálico con manchas negativas; papilas rectales con 7-8 divertículos en
cada lóbulo .............................................................................................. Simulium virgatum
15(13) Mandíbula con solo un diente marginal grande (Fig. 117 M); máximo largo de
larva 6,5-7,0 mm ..................................................................................... Simulium seriatum
– Mandíbula con un gran diente marginal y otro pequeño; máximo largo de larva
7,1-10,7 mm ...................................................................................................................... 16
16(15) Máximo largo de larva 9,1-10,7 mm; relación de artejos antenales proximal,
mediano y distal = 1: 1,7:1,2 .................................................................. Simulium bricenoi
– Máximo largo de larva 7,0 mm; relación de artejos antenales proximal, mediano y
distal = 1:1,9-2,1:1,4-1,5 ................................................................... Simulium yepocapense
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 531

PAYNEI species group

Simulium (Hemicnetha) paynei VARGAS (Figs. 113 D; 115 M, P.)


Simulium (Eusimulium) paynei VARGAS, 1942a: 246 (n. nom. for Simulium mexicanum ENDERLEIN, 1934a:
190-191, homonym of Simulium mexicanum BELLARDI, 1862: 6).
Simulium (Hemicnetha) paynei VARGAS & DÍAZ NÁJERA, 1957b: 175, 176, 178, 179, 232, 240, 241, 322;
WYGODZINSKY, 1958: 134-140; COSCARÓN, 1987: 36; IBÁÑEZ-BERNAL, 1992: 212-228; COS-
CARÓN ARIAS, 2003: 200, 205-208, 213; ADLER et al., 2004: 372; CROSSKEY & HOWARD, 1997: 38;
2004: 30.
Hemicnetha paynei PY-DANIEL & MOREIRA SAMPAIO, 1995: 118.
Simuloium (Dyarella) virgatum: VARGAS et al., 1946: 111; DALMAT, 1955: 169-170.
Simulium mathesoni VARGAS, 1943b: 360-363.
Simulium bilimekae SMART, 1944: 132.
Simulium (Dyarella) acatenangoensis DALMAT, 1951: 31-38.
Simulium (Hemicnetha) conviti RAMÍREZ PÉREZ & VULCANO, 1973: 376-379.
?Simulium mutucuna NUNES DE MELLO & VIEIRA DA SILVA, 1974: 69-71.
Female: Wing length 3.6-3.9mm. Frons and clypeus brownish with grayish pruinos-
ity. Scutum light brown with 1+1 submedian brown-reddish lyre-shaped stripes (Fig. 113
D) that with different light position become thicker and darker; humeral area lighter;
pilosity decumbent, pale gray with hair arranged in groups not well defined. Scutellum and
metanotum brown, the latter with velvety aspect. Pleurae brown, grayish pollinose. Legs
light grayish brown with darker areas brown-blackish. Abdomen dark brown. Cibarium
with small serrations on lateral cornuae, wing chaetotaxy and genitalia similar to those of
S. rubrithorax. Ratio of palpomere/sensorial vesicle 1.5-1.7.
Male: 3.1-3.6 mm. Scutum dark brown to reddish brown with 1+1 silvery
submedian stripes joining posterior gray pollinose area, showing variation in accordance
with light position. Legs as in female but more darkened; ratio of length/width of hind
basitarsus = 4.3-4.4. Abdomen dark brown. Genitalia as in S. rubrithorax.
Pupa: Cocoon and gill shape as in S. rubrithorax. Length basally about 3.7 mm, gills
2.5 mm. Frontoclypeus with abundant platelets on facial area (Fig. 115 P). Thorax dark
brown dorsally, smooth on exposed portion and with small acuminate tubercles, relatively
abundant on posterior portion. Thorax dorsally covered by gelatinous substance, with
lateral entrances appearing as 4 branches (Fig. 115 M).
Larva: Maximum length 10.4 mm. Cephalic apotome slightly darkened on basal 2/3,
with positive spots on median longitudinal area and 1+1 sublateral spots. Ratio of proximal,
medial and distal antennal articles = 1:1-1.2:0.9-1.1. Cephalic fan with 55-65 rays.
Hypostomium with 7-8 lateral setae on each side. Postgenal bridge slightly shorter than
hypostomium length. Lateral sclerite of thoracic proleg with 54-63 teeth. Anal ring with 326-
344 rows with 47-57 hooks each. Rectal papillae with 9-15 diverticula on each lobe.
Distribution: Colombia: Cundinamarca; Ecuador: Pichincha; Guatemala: Chi-
maltenango, Huehuetenango, Sacatepiquez, Santa Rosa, Totonicapan; Mexico: Chiapas,
Colima, Durango, Guerrero, Jalisco, Mexico, Michoacan, Morelos, Nuevo León, Oaxaca,
Potosí, Puebla, San Luis, Sinaloa, Veracruz; Peru: Lima, Junin; Venezuela: Aragua,
532 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

A D G H

B O

J
V

P
C

T
N

M
L
U Q

K
F

R E
Figure 113. Female. A-C: S. (Hemicnetha) rubrithorax; A: scutum; B: cibarium, C: genitalia ventral view,
showing eighth sternite, gonapophysis, cercus and paraproct (at left cercus and paraproct mounted on
slide), genital fork and spermatheca. D: S. (H.) paynei, scutum. E-F: S. (H.) biuxinisa; E: palpomere and
sensory vesicle; F: cercus and paraproct. G-L: S. (H.) brachycladum; G: scutum with frontal illumination,
H: scutum with posterior illumination; I: fronto-ocular triangle; J: cibarium; K: eighth sternite and
gonapophysis; L: cercus and paraproct. M: S. (H.) pulverulentum, cercus and paraproct. N: S. (H.)
cristalinum, scutum hairs. O-S: S. (H.) seriatum; O: cibarium; P: claw; Q: eighth sternite and
gonapophysis; R: cercus and paraproct; S: genital fork. T-U: S. (H.) smarti; T: eighth sternite and
gonapophysis; U: cercus and paraproct. V: S. (H.) yepocapense, eighth sternite and gonapophysis.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 533

Carabobo, Federal District, Guarico, Mérida, Táchira, Nueva Esparta; USA: Arizona,
Oklahoma, Texas.
Bionomics: Simulium paynei breeds in small and torrential creeks, primarily attached
to rocks in water of 10-25 °C. Females bite cows and horses (DALMAT, 1955).
Discussion: The closest species are S. rubrithorax and S. biuxinisa; the differences are
explained under each of those species. Simulium bricenoi shows great similarity, but the
wrinkled thorax of the pupa allows separation.

Simulium (Hemicnetha) rubrithorax


Figs. 113 A-C; 114 A, B; 115 A, L, O; 116 A; 117 A, D, H, K, N.
Simulium rubrithorax LUTZ, 1909: 132; 1910: 221-224; MAIA-HERZOG et al., 1984: 342-354.
Simulium (Hemicnetha) rubrithorax: COSCARÓN, 1987: 36; PY-DANIEL 1989a: 255; CROSSKEY & HOWARD,
1997: 38; 2004: 31.
Simulium magnum LANE & PORTO, 1940: 192 (major, incorrect spelling).
Eusimulium rubrithorax LANE & VULCANO, 1943: 436.
Hemicnetha rubrithorax: PY-DANIEL & MOREIRA SAMPAIO, 1995: 118; STRIEDER, 2004: 130-131.
Female: Wing length 3.0-4.0 mm. Scutum brown-orange to reddish brown with
posterior 1/3 dark grayish, anteriorly with 1+1 submedian silver stripes light grayish,
wider anteriorly, and lateral rim grayish silvery (Fig. 113 A); with changing light position,
submedian light vittae extend posteriorly in lyre shape as in S. paynei but slightly darkened.
Scutum pilosity silvery to tin colored, not homogeneously distributed. Scutellum and
metanotum brown-grayish, the latter velvety. Pleurae dark brown with grayish pruinosity.
Legs yellowish with apical portion of segments darker. Abdomen yellowish brown with
dark spots on tergal plates I-V; tergites VI-VIII shinny. Cibarium with central area smooth
without spiculae, cornuae well sclerotized (Fig. 113 B). Ratio of palpomere length/palp
sensory vesicle = 1.9. Basal portion of R without hair. Eighth sternite with about 10 setae
on each side; gonapophysis about 2 times longer than wide at base, blunt apically;
paraproct wide, subquadrate, with abundant microtrichia (Fig. 113 C); genital fork with
1+1 robust subcylindrical posterior sublateral process.
Male: Wing length 3.0-3.5 mm. Scutum yellowish brown to blackish, with anterior
1/3 silvery gray, with gray pruinosity edge laterally and posteriorly; with different
illumination, submedian light vittae show lyre shape. Metanotum as in female. Abdomen
with tergal plates velvety brown-blackish; silvery pruinose on anterior edge and 1+1 big
lateral spots on tergite II and 1+1 lateral spots on tergites VI-VII. Ratio of length/width
of hind basitarsus = 4.1. Gonostylus subcylindrical, slightly acuminate distally and with
weak apical spur (Fig. 114 A); ratio of length of gonostylus/gonocoxite = 1.6; ratio of
length/width of gonostylus = 2.4-2.9. Ventral plate with robust median process with
parallel sides (Fig. 114 B); median process subrectangular; endoparameres with abundant
and strong hooks; aedeagal membrane with abundant spiculae.
Pupa: Cocoon shoe shaped with collar of festoons joined apically, protecting gills
and sometimes larval exuviae (Fig. 115 A), length basally 3.5-4.0 mm, dorsally about 5.0
mm; gills 2.5-3.1 mm. Frontoclypeus and thorax with small tubercles, only present on
facial area of frontoclypeus (Fig. 115 O) and on posterior edge of thorax; trichomes small
534 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

D K

B
F

N J

C L H O

Figure 114. Male. A-B: S. (Hemicnetha) rubrithorax; A: genitalia ventral view showing gonocoxite,
gonostylus, ventral plate and endoparameres; B: ventral plate. C: S. (H.) virgatum, ventral plate. D-F:
S. (H.) brachycladum; D: scutum; E: gonocoxite and gonostylus ventral view; F: ventral plate. G-H: S.
(H.) cristalinum; G: gonostylus dorsal view; H: ventral plate. I-K: S. (H.) seriatum; I: gonocoxite and
gonostylus dorsal view; J: ventral plate; K: endoparameres and median sclerite. L: S. (H.)
pulverulentum, ventral plate. M: S. (H.) smarti, gonocoxite and gonostylus dorsal view. N-O: S. (H.)
earlei; N: gonocoxite and gonostylus ventral view; O: ventral plate.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 535

and single. Dorsum of thorax covered by thick gelatinous material only separated on
median line (Fig. 115 L) (material is destroyed by potash). Gill with 8 filamentous
branches, smooth with blunt apex and bifurcated close to base (Fig. 116 A).
Larva: Maximum length 11.0-12.5 mm. Aspect as in figure 117 A; color dark
greenish gray. Head dark brown, except anterolateral portion of frontoclypeus; cephalic
apotome with small dark spots on midline and mediolaterally (Fig. 117 D). Antennae
about as long as cephalic fan stem; shape as in figure 117 H; ratio of proximal, medial and
distal articles = 1:1.2-1.7:1-1.1. Palps well sclerotized. Mandibular teeth as in figure 117 K,
with big marginal tooth generally accompanied by another small tooth. Hypostomium
with curved edge and median tooth higher than lateral teeth (Fig. 117 N); lateral setae 8-
12; postgenal cleft deep subpentagonal, medially acuminate; postgenal bridge slightly
shorter than hypostomium length. Cephalic fan with about 70 rays. Lateral sclerite of
thoracic proleg with about 110 teeth arranged in groups in 1 or 2 rows. Anal sclerite well
sclerotized with hair, but not scales. Anal ring with about 320 rows of 40-60 hooks each.
Rectal papillae with about 10 diverticula on each lobe.
Distribution: Argentina: Misiones; Bolivia: Cochabamba; Brazil: Bahia, Goiás,
Minas Gerais, Rio Grande do Sul, Rio de Janeiro, Roraima, São Paulo; Guyana;
Venezuela: Aragua, Merida, Miranda Yaracuy.
Bionomics: This species breeds in very fast streams that are clear to slightly muddy.
The larvae and pupae are attached to rocks and stones in large groups covering a great
portion of the surface. Females bite horses ( LANE & PORTO, 1940; as magnum).
Discussion: Simulium paynei is the closest species, but S. paynei has a darker male thorax, a
little narrower gonostylus, and the gelatinous material of the pupal thorax has lateral entrances
appearing as 4 branches, which is very easy to see in material maintained in alcohol. Simulium
biuxinisa is differentiated by the presence in the pupa of abundant tubercles on the exposed
portion of the thorax, the gill totally protected by the cocoon, and the gelatinous material on
the thorax leaving free the median area; in the female by the larger sensory vesicle of the palp;
and in the larva by the postgenal bridge as long as the hypostomium height. Simulium bricenoi is
close but can be separated by the wrinkled surface of the thoracic pupal dorsum. Simulium
virgatum can be differentiated by the male gonostylus with a narrower median process. The
scarce differences with S. paynei, S. biuxinisa and S. virgatum suggest the need to do cytological
and molecular studies in order to resolve species identities.

Simulium (Hemicnetha) biuxinisa COSCARÓN & IBÁÑEZ-BERNAL


(Figs. 113 E, F; 115 B, N, Q; 116 C, D; 117 O.)
Simulium (Hemicnetha) biuxinisa COSCARÓN & IBÁÑEZ-BERNAL, 1993: 64; CROSSKEY & HOWARD, 1997:
38; 2004: 31.
Female (pharate): General coloration brown grayish similar to Simulium paynei.
Scutum with wide median and 1+1 sublateral reddish brown vittae separated by 1+1
narrow submedian light grayish stripes in lyre shape. Scutellum and metanotum brown
grayish, metanotum velvety. Pleurae dark brown. Abdomen brown-yellow grayish. Legs
yellowish with apex of femora, tibiae and tarsomere blackish. Fronto-ocular triangle deep;
frons with abundant long hair. Sensory vesicle of maxillary palp surpassing half length of
536 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

A D
C
B
E

H L

G I

F
J N

O U
Q
S

P
R T

Figure 115. A-K: Pupa in lateral view; A: S. (Hemicnetha) rubrithorax; B: S. (H.) biuxinisa; C: S. (H.)
virgatum; D: S. (H.) bricenoi; E: S. (H.) brachycladum; F: S. (H.) cristalinum; G: S. (H.) pulverulentum; H: S.
(H.) mexicanum; I: S. (H.) seriatum; J: S. (H.) smarti; K: S. (H.) earlei. L-N: Pupa without cocoon
showing gelatinous mass over scutum; L: S. (H.) rubrithorax; M: S. (H.) paynei; N: S. (H.) biuxinisa. O-
U: Frontoclypeus; O: S. (H.) rubrithorax; P: S. (H.) paynei; Q: S. (H.) biuxinisa; R: S. (H.) brachycladum;
S: S. (H.) cristalinum; T: S. (H.) seriatum; U: S. (H.) smarti.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 537

palpomere (Fig. 113 E). Sc with 7-9 hairs. Cibarium and gonapophysis similar to those of
S. rubrithorax. Paraproct slightly wider than length, wider than in S. rubrithorax (Fig. 113 F).
Male: Unknown.
Pupa: Cocoon with collar of 10-12 festoons not joined apically and covering gills (Fig.
115 B). Length basally 4.5-4.8 mm dorsally 6.2-6.3 mm, gills 2.2-2.5mm. Frontoclypeus with
abundant rounded platelets (Fig. 115 Q). Exposed portion of thorax with abundant
tubercles, wrinkled on dorsum, acuminate in those laterally and posteriorly positioned (Fig.
116 D); gelatinous material with 4 branches (Fig. 115 N). Gill with 8 branches (Fig. 116 C).
Larva: Maximum length 11.4-12.6 mm. Cephalic apotome dark brown, mostly
uniform in coloration, with small negative spots midlongitudinally and 1+1 median
sublaterally and subbasally. Postgenal bridge about same length as hypostomium (Fig.117
O). Antenna surpassing apex of cephalic fan stem; ratio of proximal, medial and distal
articles 1:1.5-1.6:1-1.1. Mandible with 1 well-developed marginal tooth sometimes with
additional very small teeth. Cephalic fan with 56-59 rays. Lateral sclerite of thoracic proleg
with 52-80 teeth arranged in 22-26 groups. Anal ring with 350-362 rows, with 39-42 hooks
each. Anal sclerite with hair, but not scales. Rectal papillae with 15-17 diverticula on each
lobe. No hair on body integument.
Distribution: Mexico: Oaxaca.
Bionomics: This species breeds in torrential creeks with fast, crystalline and
relatively cold water, attached to rocks, in mountainous areas with mesophilic vegetation.
Female feeding habits are unknown.
Discussion: The species is very close to S. paynei and S. rubrithorax. Differences were
treated under those species. Simulium bricenoi is similar, but can be separated by the
wrinkled dorsal portion of the thorax.

Simulium (Hemicnetha) virgatum COQUILLETT (Figs. 114 C; 115 C; 116 B.)


Simulium virgatum COQUILLETT, 1902: 97.
Simulium (Dyarella) virgatum: VARGAS et al., 1946: 111; DALMAT, 1955: 194-199.
Simulium (Hemicnetha) virgatum: VARGAS & DÍAZ NÁJERA, 1957b: 154, COSCARÓN, 1987: 36; PETERSON
et al., 1988: 84; PETERSON & KONDRATIEFF, 1995: 34-35; CROSSKEY & HOWARD, 1997: 38; 2004:
31; IBÁÑEZ-BERNAL, 1992: 228-244; ADLER et al., 2004: 374; SHELLEY et al., 2002: 148-153.
Simulium cinereum BELLARDI, 1859: 13.
Simulium tephrodes SPEISER, 1904: 148.
Simulium hippovorum MALLOCH, 1914: 28-29.
Simulium rubicundulum KNAB, 1915b: 178-179.
Simulium virgatum chiapanense HOFFMAN, 1930b: 293.
Female: Wing length 3-3.7. General coloration brownish. Scutum brown-grayish with
recumbent golden to blackish pilosity. Ornamentation similar to that of S. rubrithorax.
Changing light direction shows 1+1 submedian vittae wider anteriorly with lyre shape, dark
brown to silvery, edged by median and lateral areas gray pruinose to brownish. Scutellum
and metanotum dark brown to black with grayish pruinosity. Pleura gray to blackish with
grey pruinosity. Legs light yellowish grey with dark brown areas on apex of tibiae, femora
and tarsomeres. Abdomen brownish with tergites I-V velvety black; VI-IX shiny black.
538 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

A C D

B E

I H
F

J K M

Figure 116. A-M: Pupal gills and portion of thorax dorsally; A: S. (Hemicnetha) rubrithorax; B: S. (H.)
virgatum; C-D: S. (H.) biuxinisa; E: S. (H.) bricenoi; F: S. (H.) brachycladum; G: S. (H.) cristalinum; H: S.
(H.) mexicanum; I-J: S. (H.) seriatum, K: S. (H. ) smarti; L: S. (H.) hieroglyphicum (from PETERSON et al.,
1988); M: S. (H.) earlei.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 539

Genitalia as in S. rubrithorax but gonapophysis more rounded apically. Cibarium smooth with
small spiculae centrally. Basal sector of R bare; Sc with 10-16 hairs. Sensory vesicle of
maxillary palp surpassing half of palpomere length; ratio of palpomere/vesicle length = 1.8.
Male: Wing length 2.5-3.5mm. General coloration brownish black. Scutum yellowish
brown with ornamentation as in Simulium rubrithorax, changing coloration in accordance
with light position. Legs as in female. Abdominal segments I-IX velvety brown with 1+1
silvery pruinose spots on tergites II-VI. Genitalia as in S. rubrithorax but ventral plate with
median process thinner (Fig. 114 C).
Pupa: Cocoon with collar, festoons joined on superior edge and covering most of
gills (Fig. 115 C). Basal length 4.5-6.0 mm, dorsally 4.0-5.0 mm, height considering
festoons 2.4 mm, gills 1.9-2.3 mm. Frontoclypeus with abundant rounded to pointed
tubercles. Cephalic and thoracic trichomes single and weakly developed. Gill with 8
branches, showing well-defined basal trunk and very short primary branches (Fig. 116 B);
branches with minute striations on surface and rounded apex.
Larva (based on IBÁÑEZ-BERNAL, 1992): Maximum length about 9.0 mm. Cephalic
apotome ornamentation similar to that of rubrithorax but with median longitudinal and sublateral
median and basal spots negatives. Ratio of proximal, medial and distal antennal articles =
1:1.2:0.9; cephalic fan with about 45 rays. Hypostomium and postgenal cleft as in S. rubrithorax.
Anal ring with 220-255 rows of hooks. Rectal papillae with 7-8 diverticula on each lobe.
Distribution: Mexico: Colima, Chiapas, Chihuahua, Durango, Michoacán, Morelos,
Oaxaca, Tabasco, Veracruz; USA: Arizona, Colorado, New Mexico, South Dakota, Texas,
Utah, Wyoming.
Bionomics: This species breeds in streams at 900-2500 m, on rocks and aquatic
vegetation. Females bite livestock and domestic animals.
Discussion: Simulium virgatum is very similar to other species of the Simulium
PAYNEI species group. Differences are few and often can be considered population
variations. SHELLEY et al. (2002) gave a detailed explanation of the synonymical history and
the possibilities of four different cytotypes that MUHAMMAD (1988) described, but
according to PETERSON & KONDRATIEFF (1995), might correspond to different species
because of incorrect determination of the material. The differences are not consistent
enough to justify that the Neotropical material of S. virgatum is conspecific with S. paynei
and S. rubrithorax (SHELLEY et al., 2002). ADLER et al. (2004) recognized two species in
North American material formerly considered S. virgatum: S. hippovorum and S. virgatum
sensu stricto. A cytological and molecular study based on morphologically well-identified
material of the species in the S. PAYNEI species group is needed.

Simulium (Hemicnetha) bricenoi VARGAS , M ARTÍNEZ P ALACIOS & D ÍAZ N ÁJERA


(Figs. 115 D; 116 E.)
Simulium (Dyarella) bricenoi VARGAS et al., 1946: 115-118, 177.
Simulium (Hemicnetha) bricenoi: VARGAS & DÍAZ NÁJERA, 1957b: 174, 175, 177-179, 310; COSCARÓN, 1987:
36; IBÁÑEZ-BERNAL, 1992: 65-81; ADLER et al., 2004: 371; CROSSKEY & HOWARD, 1997: 38; 2004: 31.
Female: Wing length 4.0 mm. Frons clypeus dark brown with grayish pollinosity. Scutum
brown-grayish with hair light golden; in frontal illumination, with 1 median and 1+1 sublateral
540 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

grayish vittae, margined by dark brown 1+1 submedian lyre-shaped and thin lateral stripes;
with changing light position, grayish vittae become blackish, and thin marginal stripes become
light gray and wider anteriorly. Scutellum light brown. Legs yellowish with dark brown on apex
of tibiae, femora and tarsomeres. Abdomen brownish with tergites III-V lighter, VI-VIII shiny.
Cibarium, claws and genitalia as in S. rubrithorax, with gonapophysis slightly narrower, and ratio
of length/width 2.2. Sensory vesicle of maxillary palp nearly half of palpomere length, ratio
of length of palpomere/sensory vesicle = 1.8. Sc without hair.
Male: Wing length about 3.7 mm. Scutum blackish with ornamentation as in
Simulium paynei. Abdomen dark brown, II-VIII with 1+1 lateral pruinose silvery spots.
Ratio of length/width of hind basitarsus = 3.8-4.0. Genitalia as in S. rubrithorax but
gonostylus more pronounced with outer subapical curvature and narrower median
process of basal plate.
Pupa: Cocoon with collar of festoons joined distally (Fig. 115 D); length basally 4.5
mm, height 3.0 mm, gills 1.8 mm. Frontoclypeus as in S. rubrithorax, with platelets only on
basal area. Exposed portion of thorax smooth, but with rugosity as shown in figure 116 E,
and small tubercles laterally and posteriorly. Cephalic and thoracic trichomes single and
diminutive. Gill with 8 branches arising from short basal trunk as in S. rubrithorax.
Larva: Maximum length 9.1-10.7 mm. Cephalic apotome with positive spots arranged
discontinuously on midlongitudinal line, 1+1 sublateral and 1+1 latero-basal spots. Ratio of
proximal, medial and distal antennal articles = 1:1.8:1-1.3. Cephalic fan with 45-58 rays.
Hypostomium and postgenal cleft as in S. rubrithorax. Anal ring with 260-270 rows with 55-
60 hooks on each row. Rectal papillae with 10-18 diverticula on each lobe.
Distribution: Mexico: Distrito Federal, Durango, Mexico, Morelos, Veracruz; USA:
Arizona, New Mexico.
Bionomics: The species breeds in clear and cold water of mountainous creeks 1-3m wide,
at altitudes over 1700 m, with sandy and rocky bottoms. Female feeding habits are unknown.
Discussion: Simulium bricenoi is very close to S. rubrithorax, S. paynei and S. biuxinisa in
the adult ornamentation and general aspects of the pupa, but S. bricenoi has unique
rugosity on the dorsal portion of the thorax. Simulium hieroglyphicum has similar rugosity on
the thorax, but has 60-90 gill branches.

BRACHYCLADUM species group

Simulium brachycladum LUTZ & PINTO (Figs. 113 G-L; 114 D-F; 115 E, R; 116 F; 117 F, J, P.)
Simulium brachycladum LUTZ & PINTO, in PINTO, 1931: 690; MAIA-HERZOG et al., 1984: 341-350, 353-354.
Simulium (Hemicnetha) brachycladum COSCARÓN, 1987: 36; 1991: 274-275; CROSSKEY & HOWARD, 1997:
37; 2004: 30.
Simulium brevibranchium LUTZ & MACHADO, 1915: 46 (nomen nudum).
Female: Wing length 2.8-3.3 mm. General coloration brownish gray. Scutum brown
with gray pollinosity and abundant pile light gray intermixed with blackish hair; in frontal
illumination, with 1+1 submedian silver, narrow lyre-shaped vittae (Fig. 113 G) that with
posterior light become darker especially anteriorly (Fig. 113 H). Scutellum and metanotum
brownish gray. Pleurae dark brown, grayish pollinose. Legs brownish yellow with apical
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 541

portion of segments dark brown. Abdomen brownish yellow with tergal plate of segments
III-IX dark brown. Fronto-ocular triangle deep (Fig. 113 I). Sensory vesicle of maxillary
palp big, ratio of palpomere/sensory vesicle length = 1.6. Cibarium with abundant small
spiculae on median area (Fig. 113 J). Sc with hair on basal 3/4 or more. Eighth sternite well
sclerotized with about 25 setae; gonapophysis subtriangular, acute distally, with internal edge
well sclerotized, with abundant hair and sparse microtrichia (Fig. 113 K); ratio of length/
width at base = 1.8. Paraprocts membranous and rounded apically (Fig. 113 L). Genital fork
well sclerotized and similar to that of S. rubrithorax in shape.
Male: Wing length 2.6-3.1. Scutum darker brown-grayish with pile tin colored. Ornamenta-
tion shape as in figure 114 D. Abdomen dark brown with 1+1 big silvery spots on tergite II, only
on anterior margin in tergites III-V and most developed laterally on VI-VII. Ratio of length/
width of hind basitarsus = 6. Gonostylus relatively wide with strong protuberances laterally on
median portion (Fig. 114 E); ratio of length/wide = 1.5; ratio of gonostylus/gonocoxite length =
1.7. Ventral plate with median process slightly wider distally (Fig. 114 F).
Pupa: Cocoon shoe shaped, leaving free gills or protecting only gill base (Fig. 115 E).
Length basally 3.0-4.0 mm, dorsally 2.7-3.8 mm; maximum length 3.8-4.5 mm; gills 1.5
mm. Frontoclypeus and exposed portion of thorax smooth with only some on facial area
and sparse acute tubercles on posterior area of thorax (Fig. 115 R). Cephalic and thoracic
trichomes short, single or bifid. Gill with 6 branches short and acute distally, arising from
short basal trunk (Fig. 116 F).
Larva: Maximum length 7.5-8.5 mm. Cephalic apotome darkened on basal 2/3 and
with negative spots (Fig. 117 F). Hypostomium with straight edge with lateral teeth at
same level as median teeth; postgenal cleft medially developed with postgenal bridge
longer than hypostomium length, postgenal cleft subpentagonal (Fig. 117 P); ratio of
postgenal bridge/hypostomium length = 1.2. Cephalic fan with about 45 rays. Antenna
with relatively thick articles (Fig. 117 J); ratio of proximal, media; and distal articles =
1:1.5-1.6:0.9-1. Mandible with 1 thin marginal tooth. Anal ring with 190 rows with 31-35
hooks each. Rectal papillae with 5 diverticula on each lobe.
Distribution: Brazil: Bahia, Minas Gerais, Pernambuco, São Paulo.
Bionomics: This species breeds in torrential creeks with a range of low to high
discharge, rocky bottoms and clear water, attached to stones and consolidated sediment.
Female feeding habits are unknown.
Discussion: A similar species is S. cristalinum whose differences are treated under
that species. Simulium pulverulentum is also very close but can be differentiated by the pupal
gill with 10 branches.

Simulium (Hemicnetha) cristalinum COSCARÓN & PY-DANIEL


(Figs. 113 N; 114 G, H; 115 F, S; 116 G; 117 B, G, L, Q, T, V.)
Simulium (Hemicnetha) cristalinum COSCARÓN & PY-DANIEL, 1989: 313-318; CROSSKEY & HOWARD,
1997: 37; 2004: 30.
Hemicnetha cristalina PY-DANIEL & SAMPAIO, 1995: 118.
Female (pharate): General coloration brownish. Scutum dark brown with gray
pollinosity and thin and thicker hair intermixed (Fig. 113 N) and arranged in groups.
542 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

C
A B 1 mm

F G
E

0.2 mm
D
H

N Q P

O R

M
J
T

S
m
m
0.2

K
U V
Figure 117. A-V: Larva. A-C: general aspect in lateral view; A: S. (Hemicnetha) rubrithorax; B: S. (H.)
cristalinum; C: S. (H.) seriatum. D: S. (H.) rubrithorax, head dorsal view. E-G: cephalic apotome; E: S.
(H.) seriatum; F: S. (H.) brachycladum; G: S. (H.) cristalinum. H-J: antenna; H: S. (H.) rubrithorax; I: S.
(H.) seriatum; J: S. (H.) brachycladum. K-M: mandibular teeth; K: S. (H.) rubrithorax; L: S. (H.)
cristalinum; M: S. (H.) seriatum. N-R: postgenal cleft, hypostomium and frontal portion of
hypostomium; N: S. (H.) rubrithorax; O: S. (H.) biuxinisa; P: S. (H.) brachycladum; Q: S. (H.) cristalinum;
R: S. (H.) seriatum; S: S. (H.) seriatum, lateral sclerite of thoracic proleg; T: S. (H.) cristalinum, anal
sclerite. U-V: rectal papillae; U: S. (H.) seriatum; V: S. (H.) cristalinum.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 543

Scutum in anterior illumination with 1+1 anterior submedian subtriangular silvery


pollinose spots, continued posteriorly as vittae with lyre shape; with changing light
position, vittae get thinner and darker, and lateral edges become silvery. Scutellum light
brown, metanotum brownish gray. Abdomen dark brown. Cibarium with small teeth on
central area. Sensory vesicle of maxillary palp shorter than half of palpomere length;
ratio of palpomere/sensory vesicle length = 1.8. Eighth sternite well sclerotized
medially with about 24-28 setae on each side. Gonapophysis subtriangular, acute
distally, shorter than in S. brachycladum. Paraproct membranous, rounded apically, with
sparse setae, no microtrichia.
Male (pharate): Coloration as in female. Ratio of length/width of posterior
basitarsus about 7.2. Gonostylus relatively short with moderate mediolateral inner and
outer protuberances (Fig. 114 G), ratio of length/width = 1.8. Ratio of length
gonostylus/gonocoxite = 2.1. Ventral plate with short and relatively thick median process
(Fig. 114 H).
Pupa: Cocoon shoe shaped, aperture lower anteriorly and covering most of gill (Fig.
115 F). Length basally 3.5-3.8 mm, dorsally 2.0-2.3 mm; gills 0.7-0.8 mm. Frontoclypeus
and exposed portion of thorax smooth, with small tubercles on facial and posterior area
of thorax; frontoclypeus darkened by strong sclerotization (Fig. 115 S); cephalic and
thoracic trichomes simple and small. Gill with 6 branches very short, with transverse
striations and slightly thin but not acute apices (Fig. 116 G).
Larva: Maximum length 5.8-6.3 mm. Body shape and coloration as in figure 117
B. Cephalic apotome with posterior 2/3 darkened with median longitudinal, sublater-
al median and sublateral posterior negative spots (Fig. 117 G). Antenna surpassing
length of cephalic fan stem, with ratio of proximal, medial and distal articles = 1:1.6-
1.7:1. Cephalic fan with 36-40 rays. Mandibular teeth as in figure 117 L. Hypostomi-
um frontal edge straight with median tooth longer than lateral teeth. Postgenal cleft
subtriangular, with postgenal bridge as long as hypostomium length (Fig. 117 Q).
Lateral plate of thoracic proleg sclerite with about 72 teeth. Anal sclerite with hair
and scales (Fig. 117 T). Anal ring with 156-170 rows with 25-29 hooks each. Rectal
papillae with 5-6 diverticula on each lobe (Fig. 117 V).
Distribution: Brazil: Roraima.
Bionomics: This species was collected in a medium-size, torrential creek, with rocky
bottom and clear, temperate water.
Discussion: The closest species is S. brachycladum. It can be differentiated by several
small characters: in the female by the smaller teeth on the median area of the cibarium
and sensory vesicle of the maxillary palp longer than half of the palpomere, in the male
by the shorter gonostylus with more developed lateral protuberances and the median
process of the ventral plate being thinner and more elongated, in the pupa by the larger
size and higher cocoon aperture and gill branches about two times longer, and in the larva
by the greater number of hook rows and the hypostomium lateral teeth longer than the
median tooth. It would be useful to have more material, and to do karyological and
molecular studies to evaluate the status of these species.
544 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Simulium (Hemicnetha) pulverulentum KNAB (Figs. 113 M; 114 L; 115 G.)


Simulium pulverulentum KNAB, 1914b: 177-178; FAIRCHILD, 1940: 716-717; VARGAS, 1943b: 363;
DALMAT, 1955: 190-194.
Simulium (Hemicnetha) pulverulentum VARGAS & DÍAZ NÁJERA, 1957b: 154, 174, 176-178, 180, 263, 324;
RAMÍREZ PÉREZ, 1983: 64-65; COSCARÓN, 1987: 37; IBÁÑEZ-BERNAL, 1992: 178-195; CROSSKEY
& HOWARD, 1997: 37; 2004: 30; SHELLEY et al., 2002: 146-148.
Female: Wing length 2.3-2.7. Scutum dark brown-grayish to blackish with gray
pollinosity and abundant gray groups of recumbent hair. Ornamentation similar to that of
S. brachycladum and changeable with light position. Scutellum dark brown and metanotum
blackish. Pleurae blackish with gray pollinosity. Legs brownish yellow with darkened areas
of segments dark brown. Abdomen dark brown with tergites III-V velvety and VI-IX
shiny; tergite II with 1+1 grayish spots. Cibarium with small teeth on median area. Ratio
of palpomere/sensory vesicle length = 2.1. Sc with sparse hair on basal half. Gonapophy-
sis subtriangular, acute apically, as in S. brachycladum but basally wider. Paraprocts
subquadrangular, membranous, with sparse hair and microtrichia (Fig. 113 M).
Male: Wing length 2.0-2.5 mm. Scutum dark brown to blackish with light gray
pollinosity and golden hair, showing changeable ornamentation with light position, but
very similar to S. brachycladum. Scutellum, metanotum and pleurae as in female. Abdomen
dark brown to blackish with silver pruinosity on 1+1 large areas of tergites II and VI, and
small areas on VII-VIII. Ratio of length/width of posterior basitarsus = 5.7. Gonostylus
with well-developed lateral inner and outer protuberances, similar to that of S. brachycla-
dum. Ratio of length/width of gonostylus = 1.8; ratio of length of gonostylus/
gonocoxite = 2. Ventral plate with elongate median process (Fig. 114 L).
Pupa: Cocoon shoe shaped, relatively low anteriorly leaving gills free (Fig. 115 G).
Length basally 2.5-3.5 mm, maximum length 4.1 mm; on dorsum 2-3mm. Frontoclypeus
and exposed portion of thorax smooth, except facial area of frontoclypeus and posterior
area of thorax with small tubercles. Cephalic and thoracic trichomes short and single. Gill
with 10 branches relatively short; filaments with superficial annulations, with spiculae and
pointed apices.
Larva: Maximum length 4.9-5.5 mm. Cephalic apotome with negative spots.
Antenna surpassing apex of cephalic fan; ratio of proximal, medial and distal antennal
articles = 1:1.1:0.6. Cephalic fan with about 44 rays. Hypostomium with straight edge and
median tooth longer than lateral teeth, postgenal cleft arc shaped. Postgenal bridge about
same length as hypostomium, anal ring with about 140 rows of 25-28 hooks each. Rectal
papillae with 6-10 diverticula on each lobe.
Distribution: Belize; Costa Rica; El Salvador; Guatemala; Honduras; Mexico:
Chiapas, Guerrero, Hidalgo, Michoacan, Morelos, Oaxaca, San Luis Potosi, Veracruz;
Panama; Venezuela.
Bionomics: This species breeds in creeks and rivers with fast currents, rocky
bottoms, and clear or moderately muddy, warm water (16-21 °C). Larvae and pupae are
found on aquatic vegetation. Females bite cows, mules and horses.
Discussion: The most similar species are S. brachycladum, S. cristalinum and S.
guerrerense. The first two species can be separated by their pupal gill with 6 branches.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 545

Simulium guerrerense has 10 branches and the apex of the postgenal cleft is arc shaped, but
it has a pupal cocoon with an elevated aperture covering the branches, a male ventral plate
with a shorter median process and a larva with positive spots on the cephalic apotome and
lateral teeth of the hypostomium longer than the median tooth. These small differences
can be intraspecific variations (SHELLEY et al., 2002).

Simulium (Hemicnetha) lobatoi LUNA DIAS, HERNÁNDEZ, MAIA-HERZOG & SHELLEY


Simulium lobatoi LUNA DIAS et al., 2004: 37-42.
(Based on original description).
Female: Wing length 2.4-2.9 mm. Scutum dark brown with recumbent whitish hair
interspersed with brown hair; in anterior illumination, with 1+1 submedian whitish vittae
as in S. rubrithorax but only on 2/3 of scutum length; humeri brown, silvery pruinose; in
posterior light, with median silver pruinose vittae and median dark brown line and 1+1
rounded anterior pruinose spots, as commonly seen in S. rubrithorax and S. brachycladum.
Scutellum and metanotum dark brown, metanotum silvery pruinose. Pleurae brown,
silvery pruinose. Basal section of R and Sc haired. Legs brown with apex of femur, tibiae
and tarsomeres dark brown. Abdomen dark brown with 1+1 lateral silver pruinose spots.
Cibarium smooth. Claws with subbasal tooth. Gonapophysis subtriangular, similar to that
of S. smarti but slightly narrower apically; paraproct subquadrangular, weakly sclerotized
on margin, membranous apically and with prominent hair.
Male: Wing length 2.0-2.7 mm. Scutum orange on anterior 1/3 and darker posteriorly,
with yellowish pile and semi-erect brown hair. With anterior illumination, vittae very similar
to those of S. brachycladum; with posterior light, vittae dark brown. Sc without hair. Abdomen
dark brown to black, with pruinosity on anterolateral margins of tergite II and 1+1 laterally
on tergites III-VIII. Gonostylus similar to that of S. cristalinum. Ratio of gonostylus/
gonocoxite = 1.7; ratio of gonostylus length/width = 1.8. Ventral plate with median process
moderately developed, similar to that of S. cristalinum, but more acuminate apically.
Pupa: Cocoon with aperture anteriorly elevated at same height as posteriorly, very
similar to that of S. seriatum, protecting branches. Length basally 3.9-6.0 mm, dorsally 2.5-
3.6 mm; gills 0.9-1.8 mm. Frontoclypeus with tubercles on facial region. Gill with 8
branches arising from short trunk, branches stout, pointed apically as in S. brachycladum,
edges weakly crenate and without spicules.
Larva: Unknown.
Distribution: Brazil: Goiás, Mato Grosso.
Bionomics: The species breeds in fast currents of small to medium (0.5-10 m) clear-
water streams, attached to rocks and dead leaves. Females bite humans.
Discussion: Females of S. rubrithorax and S. brachycladum have similar scutal coloration,
but in these species the 1+1 submedian vittae join posteriorly, whereas in S. lobatoi, they
diverge posteriorly. In those species, the gonapophyses and paraprocts are longer, and the
gonapophyses are more acute apically in S. brachycladum. The median process of the male
ventral plate is also more elongated; a similar species in this character is S. cristalinum. The
pupa gill of S. brachycladum has 6 branches. Simulium rubrithorax has a similar number of
branches that are longer and not pointed apically, and the cocoon has conspicuous festoons.
546 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Simulium (Hemicnetha) guerrerense VARGAS & DÍAZ NÁJERA


Simulium (Hemicnetha) guerrerense VARGAS & DÍAZ NÁJERA, 1956: 51-56; 1957b: 153, 174, 176, 177,
179, 180, 262, 316; COSCARÓN, 1987: 35-36; IBÁÑEZ-BERNAL, 1992: 111-127; CROSSKEY &
HOWARD, 1997: 38; 2004: 30.
Female: Wing length 2.5-2.7mm. Scutum brownish, with golden pile; with frontal
illumination, 1+1 submedian light gray pollinose stripes wider frontally and curved
posteriorly in shape of lyre, anterior portion connected laterally with thin marginal
concolorous stripes; thin blackish stripe along midline; with posterior illumination, 1 median
and 1+1 sublateral wide grayish vittae emarginated by 1+1 submedian and 1+1 lateral thin
dark grayish pollinose stripes. Scutum yellowish, metanotum brown. Legs yellowish with
most segments apically dark brown. Abdomen brown with tergite I light brown, tergite II
and 1+1 on V with whitish pruinosity, shiny. Sc with hair on basal 1/3. Cibarium with small
spiculae on median area; sensory vesicle of maxillary palp surpassing half of palpomere
length. Paraproct membranous, and with scarce microtrichia as in S. pulverulentum.
Male: Wing length 2.1-2.4 mm. Scutum dark brown to black with vittae as in female.
Scutellum, metanotum, legs and abdomen as in female. Ratio of length/width of hind basitarsus
= 4.2. Ratio of gonostylus/gonocoxite = 1.4; ratio of gonostylus length/width = 1.5. Gonostylus
shape as in S. cristalinum; ventral plate similar to that of S. cristalinum but wider distally.
Pupa: Cocoon shoe shaped, very high anteriorly covering gill. Length basally: 2.3-2.7
mm; anterior height 1.5-1.9 mm, gill length 0.8-1.0. Frontoclypeus and exposed portion of
thorax smooth, with small tubercles on facial area and posterior portion of thorax;
trichomes single and small. Gill with 10 branches emerging from short trunk.
Larva: Maximum length 5.5-6.0 mm. Cephalic apotome with positive spots. Antenna
as long as cephalic stem; ratio of proximal, medial and distal segments = 1:1.3:0.8.
Postgenal cleft rounded apically; postgenal bridge slightly longer than hypostomium
length. Cephalic fan with about 47 rays. Anal ring with about 160 rows with 22 hooks
each. Rectal papillae with 6 diverticula on each lobe.
Distribution: Mexico: Guerrero.
Bionomics: Unknown.
Discussion: Simulium pulverulentum has similar adult coloration and pupal gill, but it can be
separated by the elongate median process of the ventral plate and short anterior portion of the
cocoon that leaves free the gill branches. Studies with more material and using cytological and
molecular techniques would be useful in order to assure the identity of S. guerrerense.

Simulium (Hemicnetha) solarii STONE


Simulium (Dyarella) solarii STONE, 1948: 402-404.
Simulium (Hemicnetha) solarii: VARGAS & DÍAZ NÁJERA, 1957b: 175, 176, 178, 180, 263, 328;
COSCARÓN, 1987: 37; IBÁÑEZ-BERNAL, 1992: 212-228; ADLER et al., 2004: 373-374; CROSSKEY &
HOWARD; 1997: 37; 2004: 30.
(Based on IBÁÑEZ-BERNAL, 1992).
Female: Wing length 2.5-2.8 mm. Scutum brown-orange with grayish yellow
pollinosity, 1+1 submedian and sublateral longitudinal areas delimited by 1 dark and thin
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 547

stripe, 1+1 submedian wider anteriorly, and 1+1 laterally; with anterior illumination,
submedian darker vittae conspicuous as lyre shape. Scutellum and metanotum with
coloration similar to scutum. Legs yellowish with apex of femora, tibiae and tarsomeres
dark brown. Abdomen brown with gray pollinosity on posterior margin of segments III-
VI and velvety brown on tergal plates; VII-IX shiny. Cibarium with lateral margins of
cornuae and median area of pharyngeal membrane with spiculae. Sc with hair on basal
half. Gonapophysis acute distally; relatively short and wide medially.
Male: Wing length 2.3-2.5 mm. Scutum dark brown with hair light golden and
grayish pollinosity, 1+1 submedian and 1+1 sublateral areas delimited by 1+1 submedian
and 1+1 lateral thin and dark stripes. Scutellum and metanotum similar to scutum;
scutellum shiny. Legs as in female. Abdomen dark brown with silvery pruinose spots on
tergites, medially on tergite I, and 1+1 laterally on tergites II, V-VII. Ratio of length/
width of hind basitarsus = 4.8. Gonostylus very similar to that of S. brachycladum and S.
cristalinum. Ratio of length/width = 1.9-2.1; ratio of length gonostylus/gonocoxite = 2.0-
2.2. Ventral plate short with median elongated process as in S. brachycladum.
Pupa: Cocoon shoe shaped with anterior edge moderately lower leaving pupal gills
free; weave relatively thin, translucent. Length basally 3.0-3.5 mm, height of anterior
edge 1.0-1.7 mm. Frontoclypeus and exposed portion of thorax smooth, with tubercles
on facial area of frontoclypeus, and posterior area of thorax; cephalic and thoracic
trichomes single. Gill with 15 filamentous branches bearing transverse striations with
spiculae, and acute apex.
Larva: Maximum length 8.2-8.9 mm. Cephalic apotome with positive and negative
spots. Ratio of antennal articles proximal, medial and distal = 1:1.2:0.8. Ratio of
hypostomium/postgenal bridge = 1.4. Cephalic fan with about 62 rays. Anal ring with
about 164 rows of hooks.
Distribution: Mexico: Nueva Leon, San Luis Potosi, Tamaulipas; USA: Texas, New
Mexico.
Bionomics: Simulium solarii lives in torrential streams and rivers. Female feeding
habits are poorly known; they have been collected from horses.
Discussion: The female is very close to those of S. guerrerense and S. pulverulentum in
genitalia morphology, but in these species the scutum is darker. The male is externally
similar to that of S. virgatum but differs in the ventral plate, lacking laterodistal
protuberances and having a gonostylus with lower lateral curvature. The greater number
of gill branches differs from the condition in the aforementioned species. Some pupae of
S. earlei with 15 gill branches can produce confusion, but differences in genitalia, as
mentioned for that species, allow separation.

MEXICANUM species group

Simulium (Hemicnetha) mexicanum BELLARDI (Figs. 115 H; 116 H.)


Simulium mexicanum BELLARDI, 1862: 6; VARGAS, 1941a: 119-121; 1942a: 236-237.
Simulium (Hemicnetha) mexicanum: VARGAS & DÍAZ NÁJERA, 1957b: 174-177, 179, 236-237; RAMÍREZ
PÉREZ, 1983: 57-59, COSCARÓN, 1987: 36; COSCARÓN & COSCARÓN ARIAS, 2000a: 187; SHELLEY
548 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

et al., 2002: 144-145; IBÁÑEZ-BERNAL, 1992: 144-161; CROSSKEYS & HOWARD, 1997: 38; 2004:
30; HERNÁNDEZ & SHELLEY, 2005: 6.
Simulium (Dyarella) mexicanum: VARGAS et al., 1946: 164-179; DALMAT, 1955: 185-190.
Simulium aureopunctatum MALLOCH, 1914: 27.
Simulium placidum KNAB, 1915c: 281.
Simulium lugubre LUTZ & NUÑEZ TOVAR, 1928: 46.
Eusimulium turgidum HOFFMANN, 1930b: 293, 298-300, 303.
Female: Wing length 3.0-3.4mm. General coloration dark brownish. Head dark
brown, frons and clypeus brownish with grayish pollinosity; antennae and legs dark
brown; scape and pedicel yellowish brown. Scutum reddish brown to blackish pollinose
without change of ornamentation in different light positions, with grouped gray to golden
iridescent scales, depending on illumination. Scutellum and metanotum dark brown to
black; pleura dark grayish brown, pollinose. Legs light brownish, black on tarsomeres and
brown on apices of femora and tibiae. Abdomen blackish; tergite II with 1+1 submedian
silvery spots and posterior edge dark brown; III-V velvety, VI-IX shiny and waxy. Frons
convergent below, front-ocular triangle deep. Smooth cibarium base and cornuae
branches, without spiculae. Sensory vesicle of maxillary palp about 1/3 of palpomere
length. Basal sector of R without hair; Sc with hair all along vein. Ratio of length/width
of posterior basitarsus = 5.6; claw with strong subbasal tooth. Genitalia: paraproct robust,
subrectangular with abundant microtrichia, cercus rounded on outer edge; eighth sternite
with median transverse sclerotized spot, gonapophysis about as long as wide, subovoidal
with rounded apical edge. Genital fork with robust branches and distal apodemes.
Male: Wing length 3.0-3.3 mm. Coloration similar to that of female but darker;
scutum blackish brown, pollinose, without special ornamentation and golden pilosity.
Abdomen brown with 1+1 silvery, pollinose, sublateral spots on tergite II, IV-VII. Basal
sector of R without hair. Ratio of length/width of hind basitarsus = 3.1-3.8. Gonocoxite
subtrapezoidal, gonostylus about 2 times gonocoxite length, with slightly rounded lateral
projection and thin apical spur; ratio of length/width of gonostylus = 2.3-2.5; en-
doparameres with strong teeth; basal plate wider than high with median carina and
abundant setae.
Pupa: Cocoon shoe shaped, projecting anteriorly with anterior edge reinforced (Fig.
115 H); thick silk without visible threads. Length basally 3.5-4.0 mm, dorsally 2.6-3.4 mm,
gill branches length 2.0-2.3 mm. Frontoclypeus with few platelets mostly on basal edge
and laterally, reinforced basally with some wrinkles; trichomes single with 1+1 facial and
2+2 frontal. Thorax with exposed area homogeneously covered by tubercles (Fig. 116 H).
Gill with 12 filamentous branches, emerging from 3 short primary branches that originate
from short basal trunk. Filaments closely arranged in bunch (Fig. 116 H).
Larva (based on IBÁÑEZ-BERNAL, 1992): Maximum length 6.5-8.6 mm. Coloration
light brown grayish, darkened on dorsum and lighter ventrally. Cephalic apotome light
brown, slightly darkened at base, and with median longitudinal, 1+1 submedian and
subbasal positive spots. Antennae longer than base of cephalic fans; ratio of proximal,
medial and distal articles 1:1.1-1.3:0.5-.0.6. Postgenal cleft deep; ratio of hypostomium/
hypostomial bridge = 1:1.3-1.5. Cephalic fan with about 57-63 rays. Anal ring with about
320-340 rows, with 44-47 hooks each. Rectal papillae with 10-14 diverticula on each lobe.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 549

Distribution: Belize; Bolivia; Colombia; Costa Rica; Ecuador; El Salvador;


Guatemala; Honduras; Mexico: Chiapas, Oaxaca, Veracruz; Panama; Trinidad,
Tobago; Venezuela.
Bionomics: Simulium mexicanum breeds in small streams with warm water (16-21 °C),
attached to rocks and aquatic vegetation. It is a zoophilic species (DALMAT, 1955), biting
livestock and domestic birds, but not humans (SHELLEY et al., 2002).
Discussion: Simulium seriatum is the closest species. Differences are explained under
that species. The female of S. mexicanum is similar to that of S. smarti and the male to that
of S. smarti, S. freemani and S. bricenoi. Differences in the pupa allow identification.
Simulium smarti has 18 gill branches, S. bricenoi has rugosity on the exposed area of the
thorax and S. freemani has 8 gill branches. HERNÁNDEZ & SHELLEY (2005) synonymized S.
mexicanum and S. seriatum with S. tarsatum MACQUART, 1845, based on the female holotype
(not male as published) from Colombia, without locality information. As in Simulium
(Hemicnetha), there are several species with females that cannot be differentiated by
morphological characters. We prefer to maintain the validity of S. mexicanum and S.
seriatum, pending future studies of the other life stages from the same area.

Simulium (Hemicnetha) seriatum KNAB


(Figs. 113 O-S; 114 I-K; 115 I, T; 116 I, J; 117 C, E, I, M, R, S, U.)
Simulium seriatum KNAB, 1914b: 81.
Simulium (Hemicnetha) seriatum: COSCARÓN, 1987: 36; COSCARÓN & COSCARÓN ARIAS, 2000a: 182-187;
COSCARÓN ARIAS, 2003: 200, 203, 208, 213; CROSSKEY & HOWARD, 1997: 38; 2004: 30.
HERNÁNDEZ & SHELLEY, 2005: 6.
Trichodagmia laticalx ENDERLEIN, 1934b: 291.
Trichodagmia latidigitus ENDERLEIN, 1936: 129.
Because S. seriatum is very close to S. mexicanum, we will include only the differences.
The figures presented are in accordance with the description of S. mexicanum.
Female and Male: Similar to S. mexicanum; coloration slightly darker, sensory vesicle
of maxillary palp longer, with ratio of vesicle/palpomere length = 1.7-3.0, and cibarium
with small spicules at base of cornuae (Fig. 113 O). Claw as in figure 113 P; female
genitalia as in figures 113 Q-S and male genitalia as in 114 I-K.
Pupa: Frontoclypeus without or with very few tubercles (Fig. 115 T) and exposed
portion of thorax with tubercles arranged in 2+2 longitudinal stripes, leaving other areas
smooth (Figs. 116 I, J); cocoon thinner, allowing pupa to be seen, not reinforced on
aperture border, and with small dorsal projection (Fig. 115 I). Gill with filaments relatively
open and with length of 1.6-1.8 mm. (Fig. 116 I).
Larva: General aspect as in figure 117 C; maximum length surpassing S. mexicanum
length (8.7-9.6 mm); cephalic apotome as in figure 117 E; postgenal cleft deep with ratio
of length hypostomium/postgenal bridge = 1.3-1.5 (Fig. 117 R); cephalic fan with 55-60
rays; antennae with proximal article relatively short (Fig. 117 I), with ratio of articles 1:1.4-
1.7:0.7-1. Mandibular teeth as in figure 117 M. Lateral sclerite of thoracic proleg as in
figure 117 S. Anal ring with 200-320 rows of hooks and fewer diverticula; rectal papillae
(9-14 on each lobe) (Fig. 117 U).
550 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Distribution: Argentina: Jujuy; Bolivia: Beni; Colombia: Valle; Ecuador: Canar,


Chimborazo, (Esmeraldas), Pichincha; Peru: Cuzco, Lima; Trinidad; Venezuela: Bolivar,
Distrito Federal, Táchira.
Bionomics: Immature stages breed in fast, clear streams, attached to stones. Female
feeding behavior is unknown.
Discussion: As previously explained, S. seriatum and S. mexicanum are very similar
and the adults are practically inseparable. The pupae show several differences. The pupa
of S. mexicanum has a thicker cocoon, with a reinforced border and lower aperture without
an anterior projection, the gills are a little longer and arranged in a tighter bunch, and the
frontoclypeus and exposed portion of the thorax have thick tubercles homogeneously
disposed without bare areas. The larvae show small differences, such as more hooks in the
anal ring. Material studied from Argentina, Ecuador (states of Pichincha, Chimborazo and
Canar),Venezuela (Bolivar and Tachira) and Trinidad show the characteristic pupa of S.
seriatum and those of Guatemala and Mexico are as typically described for S. mexicanum.
We consider these differences justification for maintaining S. seriatum as a valid species.

Simulium (Hemicnetha) smarti VARGAS (Figs. 113 T, U; 114 M; 115 J, U; 116 K.)
Simulium (Eusimulium) smarti VARGAS, 1946: 332-335; DALMAT, 1955: 199-204.
Simulium (Hemicnetha) smarti: VARGAS & DÍAZ NÁJERA, 1957b: 174, 175, 177, 178, 180, 263, 326;
COSCARÓN, 1987: 36; IBÁÑEZ-BERNAL, 1992: 195-212; CROSSKEY & HOWARD, 1997: 38;
2004: 30.
Female: Wing length 4.0-4.4 mm. Scutum dark brown with grayish pollinosity,
without special ornamentation; hair dark brown. Scutellum brownish, slightly lighter than
scutum. Metanotum as for scutum, but with posterior edge light brownish gray. Legs
yellowish with apex of femora, tibiae and tarsomeres brownish. Abdomen brownish gray.
tergites II-V velvety with tergal plates darkened, VI-VIII shiny. Cibarium basally with
spiculae on cornuae edge and on median area. Sensory vesicle of maxillary palp half of
palpomere length. Sc with hair along entire length. Gonapophysis relatively short with
rounded apex and nearly as wide at base as length (Fig. 113 T), half of distal surface with
microtrichia; paraproct relatively short, nearly as wide at base as length, with rounded apex
and entire surface with microtrichia (Fig. 113 U).
Male: Wing length 3.9-4.2 mm. Scutum dark brown grayish with 1+1 thin
submedian and lateral gray pollinose stripes, covered by golden groups of recumbent
hair. Scutellum, metanotum and legs as in female. Abdomen dark brownish, tergites V-
VII with 1+1 lateral silvery pruinosity. Ratio of length/width of hind basitarsus = 3.3.
Gonostylus elongated, about 2.5-2.9 longer than wide, with slight outer lateral
protuberance (Fig. 114 M). Ratio of length of gonostylus/gonocoxite = 2.3. Ventral
plate short, narrow with elevated keel.
Pupa: Cocoon shoe shaped with basal projection (Fig. 115 J); length at base 3.8-4.9
mm, height of anterior portion 2.7-3.2 mm. Gills 2.6 mm, mostly covered by cocoon.
Frontoclypeus and exposed portion of thorax with moderate number of tubercles, small
and rounded (Fig. 115 U); trichomes short and single. Gills with 18 branches, branching
close to base, with primary and secondary branches very short (Fig. 116 K).
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 551

Larva: Maximum length 1.9-2.5 mm. Cephalic apotome with positive spots arranged
along midline, and with 1+1 submedian and basal spots. Ratio of proximal, medial and
distal antennal articles = 1:1.5:0.6. Cephalic fan with 45-55 rays. Postgenal cleft acuminate
apically, postgenal bridge slightly shorter than hypostomium; ratio of hypostomium/
postgenal bridge = 1.2. Lateral sclerite of thoracic proleg with about 74 teeth. Anal ring
with about 450-460 rows of hooks. Rectal papillae with 17-20 diverticula on each lobe.
Distribution: Guatemala: Chimaltenango, El Quiché, Huehuetenango, Sacatepequ-
ez, Solalá; Mexico: Chiapas, Oaxaca.
Bionomics: Simulium smarti breeds in small waterfalls and torrential streams of
clear water (0-23 °C), attached to rocks and aquatic vegetation. Females bite livestock
(DALMAT, 1955).
Discussion: The adults of S. mexicanum and S. smarti are very similar, not showing a
pronounced keel on the basal plate of the males and with no median group of spiculae on
the female cibarium. The major differences are in the pupa, with that of S. mexicanum
having more tubercles on the frontoclypeus and thorax, and the gill having 12 branches.

Simulium (Hemicnetha) hieroglyphicum PETERSON, VARGAS & RAMÍREZ PÉREZ (Fig. 116 L.)
Simulium (Hemicnetha) hieroglyphicum PETERSON et al., 1988: 76; CROSSKEY & HOWARD, 1997: 38; 2004: 30.
(Based on original description).
Female: Wing length 4.0-4.5 mm. Scutum brownish with silvery pollinose margins;
with posterior illumination, 3 slender dark stripes extending from anterior to posterior
edge; scutum covered by golden groups of recumbent hair. Scutellum yellowish brown
with golden yellow and black setae. Metanotum with silver pollinosity. Pleurae and
abdomen brownish black. Length of palpomere/sensory vesicle = 1.9. Cibarium with
spiculae on central area; Sc with hair covering most of vein. Paraproct and gonapophysis
as in S. mexicanum.
Male: Wing length 3.9-4.2 mm. Scutum, scutellum and metanotum as in female.
Abdomen brownish black dorsally, yellowish laterally; tergites II-VII with 1+1 silvery
pollinose lateral spots. Gonostylus similar to that of S. seriatum, but with rather thin
median lateral protuberances; ratio of length/width = 2.3; ratio of length of gonostylus/
gonocoxite = 2. Ventral plate with elevated median keel as in S. smarti.
Pupa: Cocoon shoe shaped but lower anteriorly than in other Simulium (Hemicnetha)
species, projected anteroventrally and with 2-4 festoons. Length basally 6.3-7.0 mm, gills
3.0 mm. Frontoclypeus and exposed portion of thorax without tubercles but with strong
rugosity as in S. bricenoi; trichomes single. Gill very unique with short trunk and 1 thick
basal branch projecting anteriorly and posteriorly over head and thorax, respectively, with
60-90 small thin branches emerging from basal branch (Fig. 116 L).
Larva: Maximum length 13.0-13.5 mm. Cephalic apotome with positive spots.
Antenna surpassing length of cephalic fan stem; ratio of proximal, medial and distal
articles = 1:2:0.6. Cephalic fan with 52-59 rays. Postgenal bridge slightly longer than
hypostomium length. Anal ring with 500-550 rows of 50-55 hooks each. Rectal papillae
with 20-41 diverticula on each lobe.
Distribution: Costa Rica: Alajuela.
552 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Bionomics: The species breeds in shaded torrential streams about 3 m wide with a
stone bottom. Larvae are found on rocks and aquatic vegetation. Females have not been
found biting humans.
Discussion: The adult is similar in coloration and genitalia to those of S. mexicanum,
S. seriatum and S. smarti but the pupa of S. hieroglyphicum has rugosity on the frontoclypeus
and thorax and the gill has 60-90 branches. The pupa of S. bricenoi has similar rugosity but
the gill has 8 branches, and the cocoon is anteriorly more elevated.

Simulium (Hemicnetha) freemani VARGAS & DÍAZ NÁJERA


Simulium (Dyarella) freemani VARGAS & DÍAZ NÁJERA, 1949: 289-292.
Simulium (Hemicnetha) freemani VARGAS & DÍAZ NÁJERA, 1957b: 153, 175, 177-179, 263, 314-315;
COSCARÓN, 1987: 35; IBÁÑEZ-BERNAL, 1992: 98-111; CROSSKEY & HOWARD, 1997: 38; 2004: 30;
ADLER et al., 2004: 370.
Female: Wing length 4.0 mm. Scutum dark brown with yellowish pile, with 1
median wide brown vitta separated medially by thin dark brown stripe, and 1+1 light
brown sublateral vittae edged by dark gray pollinosity that also covers lateral margins
and prescutellum area. Scutellum light yellowish. Legs yellowish with apex of median
and hind femora and tarsomeres dark brown. Abdomen blackish with tergal plates of
segments III-V brownish. Sensory vesicle of maxillary palp half of palpomere length.
Cibarium with small spiculae on central area. Gonapophysis subtrapezoidal and
rounded apically, with inner edge lacking microtrichia, as in S. seriatum; paraproct
relatively short, about as long as wide.
Male: Wing length 4.0-4.2 mm. Scutum dark brown, edged all around except
anteromedially with gray pollinosity; covered by yellow hair with green shiny metallic
color. Scutellum light yellowish. Ratio of length/width of hind basitarsus = 3.8.
Gonostylus elongated with median internal prominence well developed; ratio of
length/width = 2.6-2.8. Ventral plate with prominent median carina, appearing in
outline like primitive ax.
Pupa: Cocoon shoe shaped, without expansion on anterior edge, gill free. Length
basally 2.7-3.1, height in front 2.0-2.2 mm, gills 2.0-2.1mm. Frontoclypeus and exposed
portion of thorax with rounded tubercles moderately abundant on frontal area, and more
numerous on thorax; facial area with smaller tubercles. Cephalic area and thorax with
trichomes single. Gill with 8 branches arising from short basal trunk.
Larva: Undescribed from Neotropical area. ADLER et al. (2004) show dorsally
positive spots on cephalic apotome, and deep and acute postgenal cleft, with postgenal
bridge shorter than hypostomium length.
Distribution: Mexico: Guanajuato, Guerrero, Michoacan, Oaxaca; USA: Arizona,
New Mexico, Utah.
Bionomics: The immatures were collected in rocky streams at altitudes of 1600-
2000 m. The female feeding habits are unknown.
Discussion: The closest species is S. smarti, having males with a similar laterally
compressed ventral plate, but that species has a wider hind basitarsus and a pupal gill with
18 branches.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 553

Simulium (Hemicnetha) earlei VARGAS, MARTÍNEZ PALACIOS & DÍAZ NÁJERA (Fig. 115 K.)
Simulium (Dyarella) earlei VARGAS et al., 1946: 118-120; DALMAT, 1955: 176-181.
Simulium (Hemicnetha) earlei VARGAS & DÍAZ NÁJERA, 1957b: 175-177, 180, 256, 312; COSCARÓN, 1987:
35; IBÁÑEZ-BERNAL, 1992: 81-98; SHELLEY et al., 2002: 141-144; CROSSKEY & HOWARD, 1997:
37; 2004: 30.
?Simulium (Hemicnetha) dehnei FIELD, 1969a: 162.
Female: Wing length 2.6-3.8 mm. Scutum yellowish brown to dark brown with gray
pollinosity, with frontal light 1+1 submedian and lateral light grayish vittae running 2/3 of
scutum length; with posterior light scutum darker and submedian vittae get thinner and
blackish; scutum covered by grouped whitish recumbent hair. Scutellum yellowish to dark
brown and metanotum dark brown. Pleura brown with grayish pollinosity. Legs yellowish
with middle and hind femur apices, tibiae and tarsomeres dark brown. Abdomen blackish,
tergites I-II with grayish pruinosity; tergal plates of segments III-V light brown, VI-IX shiny.
Cibarium with spiculae on central area and over pharyngeal membrane. Ratio of length of
sensory vesicle of maxillary palp/palpomere = 2.5. Paraproct typical for subgenus;
gonapophysis subtriangular, elongated with acute apex, similar to that of S. brachycladum.
Male: Wing length 2.8-3.7 mm. Scutum dark brown with golden recumbent hair;
with posterior light, 1+1 elongated submedian light grayish pollinose stripes separated on
median longitudinal line by thin blackish stripe running to anterior 2/3 of scutum length;
scutellum, metanotum pleura and legs as in female. Abdomen blackish with 1+1 silver
pruinose spots on tergites II-IV, VI-VII. Ratio of length/width of hind basitarsus = 4.8.
Gonostylus with conspicuous prominences on lateral edges of median area, as in S.
seriatum, ratio of length/width = 2.1, ratio of length/width of gonostylus/gonocoxite =
3.2. Ventral plate wide, membranous with abundant hair, as in S. seriatum.
Pupa: Cocoon shoe shaped with strong anteromedian projection leaving most of gill free
(Fig. 115 K). Length basally 3.5-4.5 mm; dorsally 3.0-3.6 mm. Gills: 1.3-1.6 mm. Frontoclypeus
and exposed portion of thorax mostly smooth, with tubercles on facial area of frontoclypeus,
fewer on anterior and posterior lateral areas of thorax. Cephalic trichomes single, thoracic with
2-5 branches. Gill with 16 (15-17) branches emerging from short trunk with very short primary
branches; branches filamentous, acute apically, striated and covered by spiculae.
Larva: Maximum length 8.7-9.6 mm. Cephalic apotome with positive spots. Ratio of
proximal, medial and distal antennal articles = 1:1.6-1.8:0.8-0.9. Postgenal cleft subcircu-
lar; postgenal bridge as long as or longer than hypostomium, ratio of hypostomium/
postgenal bridge length = 0.8-1.0. Cephalic fan with 40-51 rays. Anal ring with 200-230
rows with 27-30 hooks each. Rectal papillae with 7-10 diverticula on each lobe.
Distribution: Belize; Costa Rica; Guatemala; Mexico: Colima, Distrito Federal,
Guerrero, Mexico, Michoacan, Morelos, Nueva Leon, Oaxaca, Veracruz.
Bionomics: Simulium earlei was collected in localities between 200 and 1900 m of
altitude in fast-flowing streams with water temperatures of 13-25 °C and pH of about 7.
The larval stages were found attached to rocks and aquatic vegetation. There are no
records of females biting humans; in Belize they are zoophilic (SHELLEY et al., 2002).
Discussion: The adult scutum ornamentation and genitalia have similarities with S.
brachycladum, S. guerrerense, S. keenani, S. solarii and S. yepocapense. Differences from S. keenani
554 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

are explained under that species. The other species are easy to separate by their lower
number of gill branches, but there are specimens of S. earlei with 15 branches that can be
confused with S. solarii; the latter species has a cocoon that is lower anteriorly,
gonapophyses that are wider medially, and a ventral plate that is shorter and with a long
median process.
Simulium dehnei is included here as a synonym with doubts, as presented by CROSSKEY &
HOWARD (2004). This species is known only from a female from Panama and the type
material is lost. From the description, S. dehnei is similar to S. earlei, S. pulverulentum and S.
yepocapense, but as is common in this subgenus, females closely resemble one another. Without
other stages, it is difficult to establish the species identity, and S. dehnei could be positioned
as species inquirendae.

Simulium (Hemicnetha) keenani FIELD


Simulium (Hemicnetha) keenani FIELD, 1969a: 157-162; COSCARÓN, 1987: 30; CROSSKEY & HOWARD,
1997: 38; 2004: 30.
(Diagnosis based on original description).
Female: Scutum brownish with silvery submedian vittae lyre shaped, and with
recumbent silver hair intermixed with some erect black hair; scutellum light brown,
metanotum brown blackish. Sensory vesicle of maxillary palp slightly shorter than palpomere;
ratio of sensory vesicle/palpomere length = 2.3. Cibarium without denticles. Sc with hair
surpassing basal 3/4 of vein. Genitalia as in S. rubrithorax with paraproct slightly shorter.
Male: Scutum brownish, with some illumination nacreous; scutellum brown, metan-
otum brownish black, as in female Genitalia as in S. earlei, with ventral plate subtrapezoidal
and gonostylus with median lateral prominence well developed.
Pupa: Gill with 16 filamentous branches emerging very close to short basal trunk.
Larva: Unknown.
Distribution: Panama: Canal Zone.
Bionomics: Unknown.
Discussion: Simulium earlei is very similar, especially in the number of pupal gill
branches and the male genitalia. The principal difference is in the gonapophysis, which is
more elongated in S. earlei. The absence of type material causes more doubts about the
species validity; the name might be a synonym of S. earlei. More topotypical material and
cytological and molecular studies are needed for better identification.

Simulium (Hemicnetha) hinmani VARGAS, MARTINEZ-PALACIOS & DÍAZ-NÁJERA


Simulium (Dyarella) hinmani VARGAS et al., 1946: 120-122, 178.
Simulium (Hemicnetha) hinmani: VARGAS & DÍAZ NÁJERA, 1957b: 153, 155-177, 179, 180, 263; COSCARÓN,
1987: 36; CROSSKEY & HOWARD, 1997: 38 ; 2004: 30; IBÁÑEZ-BERNAL, 1992: 127-144.
Female: Wing length 4.0-4.3 mm. Scutum brown-orange with golden hair; in frontal
illumination, with median and 1+1 sublateral lighter vittae, separated by 1+1 submedian
darker pollinose stripes, as in S. brachycladum; with posterior illumination, median and
submedian vittae become darker, scutellum yellowish; metanotum dark brown. Legs as in
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 555

S. guerrerense. Abdomen brownish gray with silver pollinose spot on tergite II, and 1+1
smaller spots on III-V. Cibarium with small spiculae on median area, as in S. brachycladum.
Sensory vesicle of maxillary palp slightly shorter than half of palpomere length. Sc with
hair surpassing level of branching of R2+3 and R4+5. Gonapophysis elongated with
acute apex, as in S. brachycladum. Paraprocts as in S. brachycladum, but with abundant
microtichia on half distal.
Male: Wing length 4.2-4.4 mm. Scutum coloration as in female, changing intensity
of coloration on submedian and sublateral stripes with change of light position. Legs as in
female. Abdomen dark brown grayish with 1+1 grayish pollinose spots on tergites II, IV-
VII. Gonostylus long, 1.5-2.2 times gonocoxite length, and with slight lateral protuberanc-
es, as in S. bricenoi; ventral plate wide, similar to that of S. seriatum.
Pupa: Cocoon shoe shaped, with strong anterior elevation, covering about half of
basal gill. Length basally 3.6-3.7 mm high on anterior portion 1.6-1.9 mm. Frontoclypeus
smooth, exposed portion of thorax with numerous acute, short spine-like tubercles on
median dorsal area. Cephalic trichomes single and short, thoracic trichomes single but
relatively long. Gill with 10 branches, emerging from short basal trunk; filaments striated
and acute apically.
Larva: Maximum length about 8.0 mm. Cephalic apotome with positive spots.
Cephalic fan with about 45 rays. Ratio of proximal, medial and distal antennal articles =
1:1.4:0.8. Postgenal cleft relatively short with arc shape apex; ratio of postgenal bridge/
hypostomium = 1.5. Anal ring with about 225 rows of hooks. Rectal papillae with about
15 diverticula on each lobe.
Distribution: Mexico: Chiapas, Distrito Federal, Mexico, Oaxaca, Veracruz.
Bionomics: Unknown.
Discussion: Simulium yepocapense is the closest species, with similar coloration and
female and male genitalia, anteriorly elevated cocoon, and number of gill branches.
However, in the pupa it lacks acute tubercles on the thoracic dorsum and multibranched
thoracic trichomes, and in the larva, it has a deeper acute postgenal cleft. Cytological and
molecular studies would help resolve specific differences.
Simulium guerrerense and S. pulverulentum also have 10 gill branches. The first species
also has the cocoon elevated anteriorly and no tubercles on the thorax, the male ventral
plate narrower distally, and the female gonapophysis shorter and wider medially. Simulium
pulverulentum has a lower anterior portion on the cocoon an absence of tubercles on the
pupal thorax, a ventral plate with a median process and a shorter gonapophysis.

Simulium (Hemicnetha) yepocapense DALMAT (Fig. 113 V.)


Simulium (Dyarella) yepocapense DALMAT, 1949: 548-553; 1955: 204-208; VARGAS & DÍAZ NÁJERA, 1954: 60.
Simulium (Hemicnetha) yepocapense: VARGAS & DÍAZ NÁJERA, 1957b: 154, 175-177, 179, 180, 332, 333;
COSCARÓN, 1987: 36; CROSSKEY, 1987: 453; PETERSON et al., 1988: 85; IBÁÑEZ-BERNAL, 1992:
244-260; CROSSKEY & HOWARD, 1997: 38; 2004: 31.
Simulium (Dyarella) ardeni DALMAT, 1953: 35-40; 1955: 171-176.
Female: Wing length 3.4 mm. Scutum brownish gray with hair light gray-golden and
blackish intermixed; coloration changing with illumination: with frontal light, 1+1 submedi-
556 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

an and lateral dark grayish pollinose stripes, as in S. brachycladum; with posterior light,
submedian vittae thinner and anterior area darker. Scutellum and metanotum brownish gray.
Legs yellowish brown with most of apices of segments brown-grayish. Abdomen brownish,
pale on segments I-II, VI-IX reddish brown. Cibarium as in S. brachycladum; sensory vesicle
of maxillary palp shorter than half of palpomere length; Sc with hair on basal half.
Gonapophysis elongate and apically acute (Fig. 113 V), paraprocts rounded.
Male: Wing length 3.4 mm. Scutum dark brownish gray with golden and blackish pile,
with 1+1 submedian gray pollinose bands separated medially by thin longitudinal stripe, and
lateral margin of scutum whitish pruinose. Abdomen brown with 1+1 gray pruinose spots
on tergites II, IV-VI. Ratio of length/width of hind basitarsus = 3.8-4.0. Gonostylus
elongate with inner and outer curvature, as in S. seriatum; ratio of length/width = 1.9; ratio
of length of gonostylus/gonocoxite = 2.1-2.2. Ventral plate wide, as in S. earlei.
Pupa: Cocoon shoe shaped, very elevated anteriorly, covering gills, length basally 3.5-
4.0 mm, maximum height 1.8 mm, gills 1.3-1.5 mm. Frontoclypeus and exposed portion of
thorax without tubercles, except small platelets on facial area of frontoclypeus and
posterolateral area of thorax. Head trichomes single, thoracic trichomes with 2-5 branches.
Gill with 10 branches emerging from short basal trunk; filaments striated and acute apically.
Larva: Maximum length 7.0-8.0 mm. Cephalic apotome with positive spots; antenna
surpassing apex of cephalic fan stem; ratio of proximal, medial and distal antennal articles
= 1:1.9-2:1.4-1.5. Cephalic fan with about 45 rays. Postgenal cleft acute apically, postgenal
bridge slightly shorter than hypostomium length. Anal ring with about 220 rows of hooks.
Rectal papillae with 11 diverticula each.
Distribution: Guatemala: Chimaltenango, El Quiche, Huehuetenango, Suchitepe-
quez; Mexico: Chiapas, Durango.
Bionomics: According to DALMAT (1955), this species breeds in small torrential
creeks attached to rocks and aquatic vegetation from sea level to 2400 m. Females have
zoophilic habits.
Discussion: Simulium yepocapense is a peculiar species with characters that permit
identification. The female is similar in coloration and genitalia to S. brachycladum and S.
cristalinum. The male is close to S. earlei in the shape of the ventral plate and to S. bricenoi in
the shape of the gonostylus, but these species have 16 and 8 pupal gill branches,
respectively. The pupal gill with 10 branches is similar to that of S. pulverulentum and S.
guerrerense but in the former species, the cocoon does not cover the gills, and in the latter
species, the aperture is cylindrical with the same height anteriorly and posteriorly.

OVIEDOI species group

Simulium oviedoi RAMÍREZ PÉREZ (Figs. 118 A-F; 119 A-D, H-O.)
Simulium oviedoi RAMÍREZ PÉREZ, 1971: 363; 1983: 66; COSCARÓN, 1987: 27; CROSSKEY & HOWARD,
1997: 82; 2004: 77.
Female: Wing length 3.0 mm. Scutum blackish with 1 median and 1+1 sublateral
longitudinal darker areas. Scutellum brownish; metanotum dark brown, pollinose. Legs
brownish gray with apical portions of femur, tibia and tarsomeres dark brown. Abdomen
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 557

blackish, with 1+1 silvery spots on tergite II. Eighth sternite and gonapophysis as in S.
rivasi; paraproct about as long as wide at base (Fig. 118 A ); genital fork as figure 118 D.
Male: Wing length 2.6 mm. Scutum coloration as in female. Abdominal tergites II, V-
VII with 1+1 silvery spots. Ventral plate with very short concavity basally (Fig. 118 E).
Gonostylus slightly shorter than gonocoxite (Fig. 118 F); ratio of length/width at base = 1.8.
Pupa: Cocoon aperture relatively lower basally (Fig. 119 A), length basally 3.2 mm,
dorsally 3.0-3.4 mm, maximum length 4.0-4.4 mm; gills 1.4-1.7 mm. Frontoclypeus and
exposed portion of thorax with tubercles acuminate, spine-like (Figs. 119 B, D); cephalic
trichomes with 1-3 branches, thoracic trichomes with 2-4 branches. Gills as in figure 119 C.
Larva: Maximum length 7.0-8.0 mm. General aspect as in figure 119 H. Antenna
with very thick articles (Fig. 119 I); ratio of proximal, medial and distal articles = 1:1.8-
2.4:2-2.4. Comb of cephalic fan rays with similar teeth length (Fig. 119 J). Mandible as in
figure 119 K. Postgenal cleft subtriangular (Fig. 119 L); hypostomium median tooth
slightly longer than lateral teeth (Fig. 119 M). Lateral sclerite of thoracic proleg as in figure
119 N, Anal sclerite with abundant hair (Fig. 119 O), anal ring with 100-160 rows.
Distribution: Venezuela: Merida.
Bionomics: Unknown.
Discussion: This species is very close to S. rivasi, which has rounded and granulose
pupal tubercles. Simulium hinmani also has acuminate tubercles.

Simulium rivasi RAMÍREZ PÉREZ (Figs. 118 B-D, G, H; 119 E-G.)


Simulium rivasi RAMÍREZ PÉREZ, 1971: 359-360; COSCARÓN, 1987: 27; CROSSKEY, 1987: 485; CROSSKEY
& HOWARD, 1997: 82; 2004: 77.
Simulium rivai and Simulium ribasi: incorrect spellings.
Female: Size and coloration as in S. oviedoi but with more evident light, thin median
and 1+1 submedian vittae. Paraproct slightly curved apically (Fig. 118 B). Eighth sternite
with about 30 setae on each side, gonapophysis slightly sclerotized on internal margin (Fig.
118 C); ratio of length/width at base = 1; genital fork as in figure 118 D.
Male (pharate): Genitalia as in S. oviedoi but gonostylus proportionately longer (Fig.
118 H) and ventral plate with deeper concavity at base (Fig. 118 G).
Pupa: Cocoon similar to that of S. oviedoi. Length basally 5.6-6.2 mm, gills 2.8-3.0
mm. Frontoclypeus and exposed portion of thorax with abundant granulose tubercles
(Figs. 119 E, G); trichomes with 2-5 branches. Gills as in figure 119 F.
Larva: Maximum length 9.0-9.5 mm. Mandible and cephalic fan as S. oviedoi. Ratio
of proximal, medial and distal antennal articles = 1:2.7:2.8. Anal ring with about 120 rows
with 20-23 hooks each.
Distribution: Venezuela: Merida.
Bionomics: Unknown.
Discussion: Simulium rivasi is very similar to S. oviedoi, as described for that species.
The small differences between these species, based on specimens from the same place,
suggest that it would be useful to obtain more material from different localities and to do
cytological and molecular studies to assure the identity of the species and their
supraspecific taxonomic position.
558 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

C
A

F
E

Figure 118. A-D: Female; A: S. (Hemicnetha) oviedoi, cercus and paraproct. B-D: S. (H. ) rivasi; B:
cercus and paraproct; C: eighth sternite and gonapophysis; D: genital fork. E-H: Male. E-F: S. (H.)
oviedoi; E: ventral plate; F: gonocoxite and gonostylus. G-H: S. (H.) rivasi; G: ventral plate; H:
gonocoxite and gonostylus.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 559

J C I

G
E
L

K
H

Figure 119. A-G: Pupa; A-D: S. (Hemicnetha) oviedoi; A: lateral view; B: frontoclypeus; C:
anterodorsal portion of thorax and gill; D: thoracic tubercles and trichomes enlarged. E-G: S. (H.)
rivasi; E: frontoclypeus; F: anterodorsal portion of thorax and gill; G: thoracic tubercles and
trichomes enlarged. H-O: Larva; S. (H.) oviedoi; H: lateral view; I: antenna; J: comb of cephalic fan
ray; K: mandibular teeth; L: postgenal cleft and hypostomium; M: hypostomium teeth of anterior
margin; N: lateral sclerite of thoracic proleg; O: anal sclerite.
560 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Species inquirendae

Simulium tarsale WILLISTON


Simulium tarsale WILLISTON, 1893: 268-269; STONE, 1969: 316-317; HERNÁNDEZ & SHELLEY, 2005: 5-6.
Simulium clavipes MALLOCH, 1914: 40-41.
STONE’s (1969) redescriptions of females and males of this species from Dominica,
which are in accordance with the type material, and the later study of types by
HERNÁNDEZ & SHELLLEY (2005), indicate similarities with other Simulium (Hemicnetha)
species. The absence of knowledge of the larval stage, and in particular the pupa, makes
it impossible to decide if S. tarsale is conspecific with some other well-known species, such
as S. earlei, S. bricenoi, S. mexicanum or S. virgatum, with which it is similar. An important
difference is the absence in S. tarsale of subbasal tooth on the female claw and the blunt
anterolateral apodemes on the lateral branches of the genital fork. We prefer to maintain
Simulium tarsale as species inquirenda, pending more information.
Distribution: Dominica; Guadeloupe.

Simulium falculatum ENDERLEIN


Simulium falculatum ENDERLEIN, 1929: 327 (= Simulium coffeae VARGAS, 1945a: 4).
Simulium (Hemicnetha) falculatum: HERNÁNDEZ & SHELLEY, 2005: 3-5.
This species is externally similar to S. paynei and without knowledge of the genitalia,
it was synonymized with doubt (COSCARÓN, 1987). Later, ADLER et al. (2004: 373) indicated
that S. falculatum was distinct from S. paynei and removed it from synonymy. HERNÁNDEZ &
SHELLEY (2005), through genitalia studies, found differences and agreed with the
revalidation of S. falculatum. Their digital illustrations (Figs. 50, 69, 78) show a short
gonapophysis and paraproct and, in particular, an elongated stem of the genital fork,
resembling the characters of S. pictipes HAGEN and some Simulium (Hearlea) species. The
absence of knowledge of the male and larva does not permit placement of S. falculatum in
a Simulium subgenus. Without more information we consider it as species inquirenda.

Simulium tarsatum MACQUART


Simulium tarsatum MACQUART, 1845 (1844): 20.
Simulium (Hemicnetha) tarsatum: HERNÁNDEZ & SHELLEY, 2005: 6.
According to HERNÁNDEZ & SHELLEY (2005), this species, known only from the
poorly preserved female holotype (not male, as originally published) from Colombia,
without a specific locality, is similar to S. mexicanum with which it was synonymized. As
mentioned under S. mexicanum, we consider it better to maintain the well-known species
name, S. mexicanum, until other studies with more material, including all life stages of S.
tarsatum, preferably from the type area, can be examined.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 561

Simulium (Hearlea) VARGAS, MARTÍNEZ PALACIOS & DÍAZ NÁJERA (Figs. 120-127; 152.)
Simulium (Hearlea) VARGAS et al., 1946: 104.
Type-species: Simulium canadense HEARLE, 1932: 14-15.
Female: Wing length 2.5-5.0 mm. General coloration blackish brown; scutum
blackish with 1+1 silvery vittae sometimes joining 1+1 anterior submedian subtriangular
or subquadrate spots, and bordered laterally and posteriorly by silver pruinose areas. Legs
yellowish to grayish brown, with apical portions dark brown to black. Frons convergent
below, fronto-ocular triangle deep, about as long as wide; maxillary palpus with sensory
vesicle elongate, nearly 1/2 length of basal palpomere. Cibarium broadly U-shaped, with
basal portion often thickened, medially cleft, and without strong teeth, although often
with wrinkles or small serrations on pair of sublateral subovoidal prominences. Basal
sector of R bare. Sc with 6-18 setae. Hind basitarsus with length/width ratio 5.7-6.0; claw
with small, subbasal tooth. Gonapophysis surpassing edge of sternite VIII, subtriangular,
with blunt apex and gradually concave internal margins; cerci generally subquadrate,
Paraprocts low, sparsely haired, truncate distally, with abundant short hair; genital fork
with stout rods, apically expanded arms bearing strong anteriorly directed apodemes;
spermatheca ovoid, with internal spicules.
Male: Wing length 2.6-5.0 mm. General coloration similar to that of female,
differing as follows: scutum generally velvety black with anterior 1/3 with whitish
pruinosity. Sc frequently bare; hind basitarsus with length/width ratio of 2.7-3.6.
Gonocoxite subquadrate, with small protuberance on external margin; gonostylus thin,
elongate, 1/3 longer than gonocoxite, with small haired subbasal carina posteriorly, and 1
subapical spine; ventral plate generally subquadrate, with narrow median carina; en-
doparameres with wide base and numerous strong hooks; median sclerite subrectangular.
Pupa: Cocoon slipper or shoe shaped, typically coarsely woven, no threads evident,
with or without reinforced anterior edge; length basally 3.5-4.5 mm (range = 2.4-7.0 mm).
Gill inflated, gradually flattened, pseudosegmented, and lacking secondary branches to
cylindrical, smooth, with secondary branches. Frontoclypeus gradually swollen basally.
Frontoclypeus and anterodorsum of thorax smooth or covered with abundant impressed
platelets or rounded or pointed elevated granules. Frontoclypeus with 2+2 frontal and
1+1 facial setae; thorax with 3(4)+3(4) dorsocentral and 2+2 dorsolateral stout trichomes.
Abdominal chaetotaxy as in figure 125. Setae of abdominal tergite I relatively short, hair-
like, and with small platelets along anterior edge; setae of abdominal tergites II-VI short
and slender. Tergites II-IV with 4+4 retrorse hooks, those of tergite II smaller; tergites V-
VIII with minute spine-combs. Tergite X with or without terminal hooks. Sternites IV-VII
with small, wrinkled areas.
Larva: Body length 6.5-12.0 mm. Distal portion of body typically flattened dorsally
and gradually swollen ventrally (Fig. 127 G); body without evident setae. Cervical sclerites
free or adjoining posterior edge of postgena. Antenna longer than stalk of cephalic fan,
with proximal article elongate and deeply incised, creating segmented appearance; medial
article longer than proximal and distal. Cephalic apotome faintly darkened basally,
typically lacking discernible head spots. Cephalic fan with 39-54 primary rays. Mandible
with strong lateral process, robust intermediate and internal teeth, and 2 thin marginal
562 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

teeth. Postgenal cleft bluntly to acutely tapered apically; hypostomial bridge generally
shorter than hypostomium height, ratio of hypostomium/hypostomial bridge = 0.7-
1.5:1.0; anterior margin of hypostomium curved or straight, with median tooth longer
than corner teeth. Anal sclerite sometimes encircling posterior circlet, and distal portion
of abdomen with accessory plates. Anal papillae of 3 lobes, with 15-78 secondary lobules.
Posterior circlet with 102-400 rows of 16-27 hooks.
Bionomics: The immature stages are found typically on stones or vegetation in
torrenticolous streams with temperatures of 10-21 °C and at altitudes of 1000-3000 m.
Feeding habits of females are unknown.
Phylogenetic analyses show two well-differentiated clades within Simulium (Hearlea),
named JUAREZI species group and CAROLINAE species group.
Hearlea closely resembles other Neotropical subgenera, particularly Hemicnetha ENDERLEIN,
Trichodagmia ENDERLEIN and Thyrsopelma ENDERLEIN (COSCARÓN, 1987). These subgenera
have recently been included in the same subgeneric group with Freemaniellum CROSSKEY,
Anasolen ENDERLEIN, Xenosimulium CROSSKEY, and Metomphalus ENDERLEIN (MIRANDA-
ESQUIVEL, 1999; MIRANDA-ESQUIVEL & COSCARÓN, 2001). These morphological similarities
incline us to believe that Hearlea might belong to a Gondwanian lineage that likely passed
into Central America or Mexico (from South America?) during the late Cretaceous or early
Paleocene, presumably during a brief period of contact.

KEYS TO SIMULIUM (HEARLEA) SPECIES

Females
Not included are Simulium nigrocornis and S. temascalense, which are inseparable from other
species, and S. chiriquiense and S. paracarolinae which are unknown.

1 Scutum with silvery submedian vittae joining 1+1 anterior silvery spot (Figs. 120
A, B) ...................................................................................................................................... 2
– Scutum with silvery submedian vittae not joining 1+1 anterior silvery spots (Figs.
120 K, L) .............................................................................................................................. 7
2(1) Wing length 2.5-2.7 mm ......................................................................... Simulium estevezi
– Wing length > 3.0 mm ...................................................................................................... 3
3(2) Cibarium without 1+1 spindle-shaped phalanges in space between arms ..............
...................................................................................................................... Simulium ayrozai
– Cibarium with 1+1 spindle-shaped phalanges in space between arms (Fig. 120 M)
................................................................................................................................................ 4
4(3) Scutum with anterior submedian 1+1 silvery spots subquadrate (Figs. 120 A, B)
................................................................................................................................................ 5
– Scutum with anterior submedian 1+1 silvery spots subtriangular (Figs. 120 K, L)
................................................................................................................................................ 6
5(4) Posterior edge of tergites II-V grayish ............................................ Simulium canadense
– Posterior edge of tergites II-V not grayish ......................................... Simulium juarezi
6(4) Cibarium with anterior rim thin (Fig. 120 C); wing length 3.1 mm ..........................
................................................................................................................... Simulium delatorrei
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 563

– Cibarium with anterior rim thick (Fig. 120 M); wing length 3.4-3.6 mm ................
................................................ Simulium dalmati, Simulium ethelae, Simulium larvispinosum
7(1) Cibarium with acute indentation in space between arms (Fig. 120 G) .................. 8
– Cibarium with shallow indentation in space between arms (Fig. 120 M) ............ 11
8(7) Scutum with anterior 1+1 submedian silver spots subtrapezoidal to subquadrate
................................................................................................................................................ 9
– Scutum with anterior 1+1 submedian silver spots subtriangular ..............................
........................................................................................................ Simulium microbranchium
9(8) Wing length 2.7-3.1 mm ................................................................................................. 10
– Wing length 3.4 mm ............................................................................ Simulium menchacai
10(9) Wing length 2.7-2.8 mm; scutum with anterior submedian 1+1 silver spots
subtrapezoidal ....................................................................................... Simulium capricorne
– Wing length 2.9-3.1 mm; scutum with anterior submedian 1+1 silver spots
subquadrate ................................................................................................. Simulium burchi
11(7) Wing length about 5.0 mm ............................................................... Simulium contrerense
– Wing length less than 4.0 mm ....................................................................................... 12
12(11) Wing length 3.7-3.9 mm; scutum with anterior submedian silvery spots
subtrapezoidal ......................................................................................... Simulium carolinae
– Wing length < 3.2 mm; scutum with anterior submedian silvery spots
subtriangular .......................................................... Simulium gorirossiae, Simulium johnsoni

Males
No male key is presented due to lack of characters.

Pupae
Unknown: Simulium paracarolinae.

1 Gill branches cylindrical, smooth, and with 4 or more small secondary branches
(Fig. 124). Cocoon generally slipper-shaped (Figs. 124 A, J) .................................... 2
– Gill branches flattened or subcylindrical, with noticeable wrinkling or annulations,
and lacking small secondary branches (Figs. 122, 123). Cocoon more typically
shoe-shaped (Figs. 122 A, J, L; 123 A, G) .................................................................... 7
2(1) Secondary gill branches smaller than primary anterodorsal branch. Ventral branch
with 2 secondary acuminate processes; anterior edge of cocoon complete, lacking
noticeable threads .............................................................................................................. 3
– Secondary gill branches as large as primary anterodorsal branch (Figs. 124 J, K).
Ventral branch with 6-7 secondary acuminate processes; anterior edge of cocoon
ragged, with noticeable threads ......................................................... Simulium menchacai
3(2) Gill with primary anterodorsal branch acuminate (Figs. 124 F, G) ........................ 4
– Gill with primary anterodorsal branch blunt (Figs. 124 A-E, H, I) ........................ 5
4(3) Frontoclypeus and thorax with sparse granules ............................Simulium gorirossiae
– Frontoclypeus and thorax without granules ............................... Simulium temascalense
5(3) Frontoclypeus and thorax with abundant granules. Cocoon with reinforced
anterior edge ........................................................................................... Simulium carolinae
564 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

– Frontoclypeus and thorax without elevated granules. Cocoon without, or with


only slightly, reinforced anterior edge ........................................................................... 6
6(5) Secondary gill branches strongly spined apically (Figs. 124 H, I). Frontoclypeus
without platelets ...................................................................................... Simulium johnsoni
– Secondary gill branches blunt or weakly spined apically (Figs. 124 A-D).
Frontoclypeus and thorax with impression of minute non-elevated granules
especially on edge of disc ............................................................ Simulium larvispinosum
7(1) Gill with 2 branches .......................................................................................................... 8
– Gill with 3 branches ........................................................................................................ 12
8(7) Cocoon shoe-shaped (Figs. 122 A, J, L) ........................................................................ 9
– Cocoon mostly slipper-shaped (Figs. 122 D, F) ........................................................ 11
9(8) Dorsal gill branch 1.8 times longer than ventral branch and strongly curved (Figs.
122 J, K) ....................................................................................... Simulium microbranchium
– Dorsal gill branch 0.6-1.0 times as long as ventral branch and only mildly curved
(Figs. 122 B, L-N) ............................................................................................................. 10
10(8) Gill protruding well beyond anterior margin of cocoon (Figs. 122 L-N); apex of
gill branches blunt .................................................................................Simulium nigricorne
– Gill largely concealed by cocoon (Figs. 122 A, B); apex of gill branches acuminate
...................................................................................................................... Simulium ayrozai
11(8) Dorsal gill branch acuminate, subequal in thickness to ventral branch, and only
slightly curved apically (Figs. 122 F-I) ............. Simulium ethelae, Simulium chiriquiense
– Dorsal gill branch blunt, twice as thick as ventral branch, and strongly curved
apically (Figs. 122 D, E) ...................................................................... Simulium capricorne
12(7) Frontal gill branch subglobose, inflated, tuberculated, not annulated (Figs. 123 N,
O) ......................................................................................................................................... 13
– Frontal gill branch subcylindrical, scarcely or not tuberculated, annulated ........ 14
13(12) Two frontal gill branches inflated, ventral branch with ventral area membranous;
another ventrolateral branch small and subannulated ................. Simulium contrerense
– Only 1 frontal gill branch inflated, plus 1 ventrolateral and another dorsolateral
branch thinner and subannulated (Figs. 123 N, O) ........................... Simulium juarezi
14(12) Dorsal gill branch attenuated distally (Figs. 122 C; 123 C-I) .................................. 15
– Dorsal gill branch thickened and capitate distally (Figs. 123 A, B, J-M) .............. 17
15(14) Dorsal gill branch with small ventral protuberance at about midlength (Figs. 123
C, G) ................................................................................................................................... 16
– Dorsal branch lacking ventral protuberance (Fig. 122 C) ........... Simulium canadense
16(15) Dorsal gill branch with blunt apex (Figs. 123 C-F). Frontoclypeus with platelets
laterally and medially with only punctuation (Fig. 121 H) ........... Simulium delatorrei
– Dorsal gill branch with acuminate apex (Figs. 123 G-I). Frontoclypeus with
platelets on entire surface (Fig. 121 I) ................................................. Simulium estevezi
17(14) Dorsal gill branch globose apically, with well-developed tubercles. Cocoon shoe-
shaped (Figs. 123 A) .................................................................................. Simulium burchi
– Dorsal gill branch with 3 apical prominences, lacking tubercles (Figs. 123 J-M).
Cocoon slipper-shaped (Figs. 123 J) ....................................................Simulium dalmati
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 565

Larvae
Simulium nigricorne and S. chiriquiense are unknown.

1 Hypostomium with anterior edge convex and with intermediate teeth above level
of corner teeth (Fig. 126 J). Distal portion of abdomen with accessory plates
(Figs. 127 B-E) .................................................................................................................... 2
– Hypostomium with anterior edge straight and with intermediate teeth below level
of corner teeth (Fig. 126 B). Distal portion of abdomen without accessory plates
................................................................................................................................................ 8
2(1) Distal portion of abdomen with only 1+1 accessory dorsolateral plates; ventral
arm short, not encircling base of posterior circlet (Figs. 127 B, C); anal ring with
about 400 rows of hooks ..............................................................Simulium paracarolinae
– Distal portion of abdomen with 2+2 accessory dorsolateral plates (Figs. 127 D,
E); anal sclerite ventral arm encircling base of posterior circlet (Figs. 127 D-I);
anal ring with < 300 rows of hooks .............................................................................. 3
3(2) Distal portion of abdomen without ventrolateral accessory plates ..........................
.................................................................................................................... Simulium carolinae
– Distal portion of abdomen with stout ventrolateral accessory plates (Figs. 127 D,
H, J) ....................................................................................................................................... 4
4(3) Lobes of accessory ventrolateral plate shallowly incised (Figs. 127 H, I [v1]).
Segment VIII without ventrolateral tubercles ............................................................. 5
– Lobes of accessory ventrolateral plate deeply incised (Fig. 127 D [v1]). Segment
VIII with well-sclerotized ventrolateral tubercles (Figs. 127 D-F [vp]) ................. 6
5(4) Cephalic fan with 43-48 primary rays; area before to anal sclerite with 1+1
dorsal subrectangular flattened accessory plates (Figs. 127 H, I [df]) ...................
............................................................................................................ Simulium larvispinosum
– Cephalic fan with 56 primary rays; area before to anal sclerite without dorsal
accessory plates ....................................................................................... Simulium johnsoni
6(4) Accessory ventrolateral plates of distal portion of abdomen with anteriorly directed
acuminate processes; anal ring with 310 rows of hooks ............... Simulium menchacai
– Accessory ventrolateral plates of distal portion of abdomen with anterolaterally
directed acuminate processes; anal ring with 242-282 rows of hooks ................... 7
7(6) Accessory ventrolateral plates of distal portion of abdomen strongly toothed
(Fig. 127 D [vl]); ventrolateral papillae of sternite VIII with acuminate tubercles
(Figs. 127 D-F [vp]) .............................................................................Simulium gorirossiae
– Accessory ventrolateral plates of distal portion of abdomen with serrated margin
(Fig. 127 J [vl]); ventrolateral papillae of sternite VIII smooth .................................
.............................................................................................................. Simulium temascalense
8(1) Rectal papillae with 60-78 lobules .................................................................................. 9
– Rectal papillae with 45 or fewer lobules ..................................................................... 12
9(8) Rectal papillae with 72-78 lobules; length of mature larva < 7.0-7.7 mm .......... 10
– Rectal papillae with 60-66 lobules; length of mature larva > 12.0 mm ...................
................................................................................................................. Simulium contrerense
566 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

10(9) Anal ring with about 114 rows of hooks. Cephalic fan with 39-40 primary rays
................................................................................................................... Simulium delatorrei
– Anal ring with about 172-204 rows of hooks. Cephalic fan with 46-57 primary
rays ...................................................................................................................................... 11
11(10) Anal ring with 172-178 rows of hooks; cephalic fan with 46-54 primary rays ......
........................................................................................................................ Simulium burchi
– Anal ring with 194-206 rows of hooks; cephalic fan with 56-57 primary rays ......
........................................................................................................ Simulium microbranchium
12(8) Body length 8.1-10.0 mm; hypostomium with 11-17 lateral setae ........................ 13
– Body length 7.0-8.0 mm; hypostomium with 10 or fewer lateral setae ................ 15
13(12) Cephalic fan with 42-44 primary rays; ratio of length of hypostomium/
hypostomial bridge = 0.8 .......................................................................Simulium dalmati
– Cephalic fan with 48-56 primary rays; ratio of length of hypostomium/
hypostomial bridge = 1.0 ............................................................................................... 14
14(13) Anal ring with 18-20 hooklets per row; rectal papillae with 15-24 lobules .............
.................................................................................................................. Simulium canadense
– Anal ring with 24-27 hooklets per row; rectal papillae with 36-45 lobules .............
...................................................................................................................... Simulium ayrozai
15(12) Body length 7.0 mm; cephalic fan with 36-40 primary rays ............ Simulium ethelae
– Body length 7.3-8.0 mm; cephalic fan with 42-50 primary rays ............................ 16
16(15) Body length 8.0 mm; anal ring with 98-104 rows of 24-30 hooks ......................
.....................................................................................................................Simulium estevezi
– Body length 7.5-7.7 mm; anal ring with 115-145 rows of 15-24 hooks .............. 17
17(16) Anal ring with 145 rows of about 17 hooks ....................................... Simulium juarezi
– Anal ring with 115-135 rows of 20-24 hooks ................................ Simulium capricorne

CLAVES DE ESPECIES DE SIMULIUM (HEARLEA)


Hembras
No incluidas Simulium nigricorne y S. temascalense por inseparables de otras especies, S.
chiriquiense y S. paracarolinae por desconocida.

1 Escudo con bandas plateadas submedianas uniéndose a 1+1 manchas plateadas


anteriores (Figs. 120 A, B) ................................................................................................ 2
– Escudo con bandas plateadas submedianas no uniéndose a 1+1 manchas
plateadas anteriores (Figs. 120 K, L) .............................................................................. 7
2(1) Largo ala 2,5-2,7 mm .............................................................................. Simulium estevezi
– Largo ala >3,0 mm ............................................................................................................ 3
3(2) Cibario con 1+1 elevación en forma de uso en el espacio entre las ramas (Fig.
120 M) .................................................................................................................................. 4
– Cibario sin 1+1 elevación en forma de uso en el espacio entre las ramas ..............
...................................................................................................................... Simulium ayrozai
4(3) Escudo con 1+1 manchas plateadas anteriores submedianas subcuadradas (Figs.
120 A, B) .............................................................................................................................. 5
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 567

– Escudo con 1+1 manchas plateadas anteriores submedianas subtriangulares (Figs.


120 K, L) .............................................................................................................................. 6
5(4) Margen posterior de los tergitos II-V grisáceo .............................. Simulium canadense
– Margen posterior de los tergitos II-V no grisáceo ............................. Simulium juarezi
6(4) Cibario con el borde anterior fino (Fig. 120 C); largo ala 3,1 mm ............................
................................................................................................................... Simulium delatorrei
– Cibario con el borde anterior grueso (Fig. 120 M); largo ala 3,4-3,6 mm ...............
................................................ Simulium dalmati, Simulium ethelae, Simulium larvispinosum
7(1) Cibario con espacios entrecortados suavemente aguzados entre las ramas (Fig. 120
G) ........................................................................................................................................... 8
– Cibario con espacios entrecortados bajos entre las ramas (Fig. 120 M) .............. 11
8(7) Escudo con 1+1 mancha plateada submediana subtrapezoidal a subcuadrada .......... 9
– Escudo con 1+1 mancha plateada submediana subtriangular ....... Simulium microbranchium
9(8) Largo ala 2,7-3,1 mm ...................................................................................................... 10
– Largo ala 3,4 mm ................................................................................. Simulium menchacai
10(9) Largo ala 2,7-2,8 mm; escudo con 1+1 manchas plateadas anteriores submedianas
subtrapezoidales ................................................................................... Simulium capricorne
– Largo ala 2,9-3,1 mm; escudo con 1+1 manchas plateadas anteriores submedianas
subcuadradas ............................................................................................... Simulium burchi
11(7) Largo ala alrededor de 5,0 mm ........................................................ Simulium contrerense
– Largo ala menos de 4,0 mm .......................................................................................... 12
12(11) Largo ala 3,9 mm; escudo con 1+1 manchas plateadas anteriores submedianas
subtrapezoidales ..................................................................................... Simulium carolinae
– Largo ala < 3,2 mm; escudo con 1+1 manchas plateadas anteriores submedianas
subtriangulares ...................................................... Simulium gorirossiae, Simulium johnsoni

Machos
No se presenta clave por escasez de caracteres.

Pupas
Desconcida: Simulium paracarolinae.

1 Ramas de la branquia subcilíndrica, lisa, con 4 o más ramas secundarias (Fig. 124);
capullo típicamente en forma de chinela (Figs. 124 A, J) ......................................... 2
– Ramas de la branquias achatadas o subcilíndricas con notoria rugosidad o
anillación y sin pequeñas ramas secundarias (Figs. 122, 123); capullo generalmente
en forma de zapato (Figs. 122 A, J, L; 123 A, G) ....................................................... 7
2(1) Rama secundaria ventral de las branquias más pequeñas que la rama primaria
anterodorsal; rama ventral con 2 prolongaciones secundarias aguzadas; margen
anterior del capullo con tejido compacto ..................................................................... 3
– Ramas secundarias ventrales de la branquia tan larga como la primaria anterodorsal
(Figs. 124 J, K); rama ventral con 6-7 prolongaciones secundarias aguzadas; margen
anterior del capullo deshilachado ....................................................... Simulium menchacai
3(2) Branquia con rama primaria anterodorsal aguzada (Figs. 124 F, G) ....................... 4
568 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

– Branquia con rama primaria anterodorsal roma (Figs. 124 A-E, H, I) .................. 5
4(3) Frontoclípeo y tórax con tubérculos esparcidos ...........................Simulium gorirossiae
– Frontoclípeo y tórax sin tubérculos ............................................. Simulium temascalense
5(3) Frontoclípeo y tórax con abundantes tubérculos; capullo con el borde anterior
reforzado .................................................................................................. Simulium carolinae
– Frontoclípeo y tórax al máximo con algunos tubérculos esparcidos; capullo sin o
con solo un ligero refuerzo en el borde anterior ........................................................ 6
6(5) Ramas secundarias de las branquias con fuertes espinas (Figs. 124 H, I);
frontoclípeo sin tubérculos ................................................................... Simulium johnsoni
– Ramas secundarias de las branquias romas o con débiles espinas (Figs. 124 A-D);
frontoclípeo y tórax con pequeños tubérculos no elevados hasta aguzados
especialmente en el dorso del tórax ........................................... Simulium larvispinosum
7(1) Branquia con 2 ramas principales: dorsal y ventral .................................................... 8
– Branquia con 3 ramas principales: dorsal, mediana y ventral ................................. 12
8(7) Capullo en forma de zapato (Figs. 122 A, J, L) ........................................................... 9
– Capullo en forma de chinela (Figs. 122 D, F) ............................................................ 11
9(8) Rama dorsal de la branquia 1,8 veces más larga que la ventral y fuertemente
curvada (Figs. 122 J, K) ............................................................ Simulium microbranchium
– Rama dorsal de la branquia 0,6-1,0 veces más larga que la ventral y débilmente
curvada (Figs. 122 B, L, N); ápices de las ramas de la branquia romas ............... 10
10(9) Branquia sobresale bien más allá del borde anterior del capullo (Figs.122 L-N);
ápice de las ramas romo ......................................................................Simulium nigricorne
– Branquia bien cubierta por el capullo (Figs. 122 A, B); ápice de las ramas de las
branquias aguzadas .................................................................................. Simulium ayrozai
11(8) Rama dorsal de la branquia aguzada, casi igual en grosor a la rama ventral y solo
ligeramente curvada apicalmente (Figs. 122 F-I) ..........................................................
.................................................................................. Simulium ethelae, Simulium chiriquiense
– Rama dorsal de la branquia roma, el doble de gruesa que la rama ventral y
fuertemente curvada apicalmente (Figs. 122 D, E) ...................... Simulium capricorne
12(7) Rama mediana de la branquia subglobosa, inflada, tuberculada, no anillada,
excepto basalmente (Figs. 123 N, O) ........................................................................... 13
– Rama mediana de la branquia subglobosa, subcilíndrica, escasamente o no tuberculada
o anillada ............................................................................................................................. 14
13(12) Ramas dorsal y media de la branquia infladas, la mediana membranosa
ventralmente; rama ventral pequeña y subanillada ...................... Simulium contrerense
– Rama mediana de la branquia inflada; ramas dorsal y ventral finas, subanilladas
(Figs. 123 N, O) .........................................................................................Simulium juarezi
14(12) Rama mediana de la branquia redondeada o desde débil a fuertemente aguzada
distalmente (Figs. 122 C; 123 C-I) ................................................................................ 15
– Rama mediana de la branquia engrosada y capitada distalmente (Figs. 123 A, B,
J-M) ..................................................................................................................................... 17
15(14) Rama dorsal de la branquia con pequeña protuberancia ventral cerca de la mitad
del largo (Figs. 123 C, G) ............................................................................................... 16
– Rama dorsal de la branquia sin protuberancia ventral (Fig. 122 C) ...... Simulium canadense
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 569

16(15) Rama dorsal de la branquia con el ápice romo (Figs. 123 C-F); frontoclípeo con
tubérculos lateralmente, medianamente solo con puntuaciones (Fig. 121 H) ........
................................................................................................................... Simulium delatorrei
– Rama dorsal de la branquia con ápice aguzado (Figs. 123 G-I); frontoclípeo con
tubérculos en toda la superficie (Fig. 121 I) ....................................... Simulium estevezi
17(14) Rama dorsal de la branquia globosa apicalmente, con tuberosidades bien
desarrolladas; capullo con forma de zapato (Fig. 123 A) .................. Simulium burchi
– Rama dorsal de la branquia con 3 prominencias apicales faltando tuberosidades
(Fig. 123 J-M; capullo con forma de chinela (Fig. 123 J) ................Simulium dalmati

Larvas
Simulium nigricorne y S. chiriquiense desconocidas.

1 Hipostomio con margen anterior convexo y con dientes intermedios arriba del
nivel de los dientes laterales (Fig. 126 J); porción posterior del abdomen con
placas accesorias (Figs. 127 B-E) .................................................................................... 2
– Hipostomio con margen anterior recto, dientes intermedios más cortos que los
dientes laterales (Fig. 126 B); porción posterior del abdomen sin placas accesorias
................................................................................................................................................ 8
2(1) Porción posterior del abdomen con solamente 1+1 placas accesorias
dorsolaterales; sostenes ventrales del esclerito anal cortos no rodeando la base
del anillo anal (Figs. 127 B, C); anillo anal con alrededor de 400 hileras de
ganchos .............................................................................................Simulium paracarolinae
– Porción posterior del abdomen con 2+2 placas accesorias dorsolaterales (Figs.
127 D, E); sostenes ventrales del esclerito anal rodeando la base del anillo anal
(Figs. 127 D, I); anillo anal con < 300 hileras de ganchos ........................................ 3
3(2) Porción posterior del abdomen sin placas accesorias ventrolaterales ......................
.................................................................................................................... Simulium carolinae
– Porción posterior del abdomen con robustas placas accesorias ventrolaterales
(Figs. 127 D, H, J) .............................................................................................................. 4
4(3) Lóbulos de la placa accesoria ventrolateral con incisiones no profundas (Figs. 127
H, I [vl] ); segmento VIII sin papilas ventrolaterales ................................................. 5
– Lóbulos de la placa accesoria ventrolateral con incisiones profundas (Figs. 127 D
[vl] ); segmento VIII con bien definidas papilas esclerotizadas ventrolaterales
(Figs. 127 D-F [vp] ) .......................................................................................................... 6
5(4) Abanico cefálico con 43-48 rayos; área anterior al esclerito anal con 1+1 placas
accesorias dorsales, subrectangulares, achatadas (Figs. 127 H, I [df]) ......................
............................................................................................................ Simulium larvispinosum
– Abanico cefálico con 56 rayos; área anterior al esclerito anal sin placas accesorias
dorsales ...................................................................................................... Simulium johnsoni
6(4) Placas accesorias de la porción posterior del abdomen con prolongaciones
aguzadas anteriormente dirigidas; anillo anal con 310 hileras de ganchos ..............
.................................................................................................................. Simulium menchacai
570 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

– Placas accesorias ventrolaterales de porción posterior del abdomen sin pro-


longaciones aguzadas anteriormente dirigidas; anillo anal con 242-282 hileras de
ganchos ................................................................................................................................. 7
7(6) Placas accesorias de la porción posterior del abdomen fuertemente dentadas (Fig.
127 D [vl] ); papilas ventrolaterales del VIII esternito con tubérculos aguzados
(Figs. 127 D-F [vp]) ........................................................................... Simulium gorirrossiae
– Placas accesorias de la porción posterior del abdomen con borde serrado (Fig. 127
J [vl] ); papilas ventrolaterales del VIII esternito lisas ............. Simulium temascalense
8(1) Papilas rectales con 60-78 divertículos .......................................................................... 9
– Papilas rectales con 45 o menos divertículos ............................................................. 12
9(8) Papilas rectales con 72-78 divertículos; largo de la larva madura < 7,0-7,7 mm
.............................................................................................................................................. 10
– Papilas rectales con 60-66 divertículos; largo de la larva madura > 12,0 mm .......
................................................................................................................. Simulium contrerense
10(9) Anillo anal con alrededor de 114 hileras de ganchos; abanico cefálico con 30-40
rayos ........................................................................................................ Simulium delatorrei
– Anillo anal con alrededor de 172-204 hileras de ganchos; abanico cefálico con 46-
57 rayos .............................................................................................................................. 11
11(10) Anillo anal con 172-178 hileras de ganchos; abanico cefálico con 46-54 rayos .....
........................................................................................................................ Simulium burchi
– Anillo anal con 194-206 hileras de ganchos; abanico cefálico con 56-57 rayos .....
........................................................................................................ Simulium microbranchium
12(8) Largo del cuerpo 8,0-10,0 mm; hipostomio con 11-17 cerdas laterales .............. 13
– Largo del cuerpo 7,0-8,0 mm; hipostomio con 10 o menos cerdas laterales ..... 15
13(12) Abanico cefálico con 42-44 rayos; relación largo hipostomio/puente postgenal
= 0,8 ............................................................................................................Simulium dalmati
– Abanico cefálico con 48-56 rayos; relación largo hipostomio/puente postgenal
= 1,0 .................................................................................................................................... 14
14(13) Anillo anal con 18-20 ganchos por hilera; papilas rectales con 15-24 divertículos
.................................................................................................................. Simulium canadense
– Anillo anal con 24-27 ganchos por hilera; papilas rectales con 36-45 divertículos
...................................................................................................................... Simulium ayrozai
15(12) Largo del cuerpo 7,0 mm; abanico cefálico con 36-40 rayos .......... Simulium ethelae
– Largo del cuerpo 7,3-8,0 mm; abanico cefálico con 42-50 rayos .......................... 16
16(15) Largo del cuerpo 8,0 mm; anillo anal con 98-104 hileras de 24-30 ganchos cada
una ............................................................................................................... Simulium estevezi
– Largo del cuerpo 7,5-7,7 mm; anillo anal con 115-145 hileras de 15-24 ganchos
cada una ............................................................................................................................. 17
17(16) Anillo anal con 145 hileras con 17 ganchos cada una ...................... Simulium juarezi
– Anillo anal con 115-135 hileras con 20-24 ganchos cada una ............................
.................................................................................................................. Simulium capricorne
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 571

JUAREZI species group

Simulium (Hearlea) ayrozai VARGAS (Figs. 122 A, B.)


Simulium ayrozai VARGAS, 1945a: 74.
Simulium (Hearlea) ayrozai: VARGAS et al., 1946: 106, 111, 164, 181, 184, 185; VARGAS & DÍAZ NÁJERA,
1957b: 166, 167, 170, 172, 173, 282; DÍAZ NÁJERA & VULCANO, 1962: 93, 108; COSCARÓN et al.,
2004: 10-11.
Female: Wing length 3.4 mm. General coloration blackish. Scutum velvety black,
with 1+1 silvery vittae joining anterior subtriangular spots with whitish pruinose
prescutellar area. Fronto-ocular triangle longer than wide. Cibarium broadly U-shaped,
with smooth edge, lacking ovoid sublateral prominences. Sc pilose, with 21 hairs. Hind
basitarsus with length/width ratio of 6:1; claw with subbasal tooth. Genitalia as in S.
capricorne: sternite VIII with median sclerotized area and 17-19 trichomes per side,
gonapophysis with internal edge smooth and sclerotized along inner 1/2; cerci with distal
margin straight; Paraprocts well sclerotized, with abundant hair distally.
Male: Wing length 3.4 mm. Scutum velvety black. Sc with few hairs. Hind basitarsus
with length/width ratio = 4.2. Genitalia as in S. capricorne, although ventral plate more
elevated distally.
Pupa: (Figs. 122 A, B). Cocoon shoe shaped, anterior margin reinforced, coarsely
woven; basal length 3.5-4.7 mm. Frontoclypeus and anterodorsal portion of thorax
smooth, with sparse tubercles; frontoclypeus with 2 small frontal and 1 large facial setae
per side. Gill with 2 wide, curved, pseudo annulated branches (particularly along inner
surface) whose apices meet along longitudinal midline, forming ring; gill typically not
extended beyond anterior margin of cocoon; dorsal branch 0.9 mm, with small acute
process distally; ventral branch 1.0 mm. Abdomen without terminal hooks.
Larva: Body length 8.0-10.0 mm. General coloration grey. Cephalic apotome
uniformly light brown, gradually darkened mediobasally and small anteromedian and
submedian spots. Cephalic fan with 52-54 rays. Hypostomium with straight anterior
margin, 14-17 lateral setae per side, and 4+4 discal setae. Mandible with 2 rows of 7-8
internal teeth, 2 thin marginal teeth, and 1 large latero-mandibular process. Ratio of
proximal, medial and distal antennal articles = 1:1.5:1.0. Ratio of length of hypostomi-
um/hypostomial bridge = 1:1. Postgenal cleft dome-shaped, with narrow anteromedian
incision. Cervical sclerites joining postgenae posteriorly. Lateral sclerite of thoracic proleg
with 46-52 teeth. Anal ring with about 150 rows of 24-27 hooks. Rectal papillae of 3
compound lobes, with 12-15 diverticula per lobe; dorsal lobe larger than ventral ones.
Distribution: Guatemala: Solalá, Totonicapan; México: México, Distrito Federal.
Bionomics: Unknown.
Discussion: Simulium microbranchium is very close structurally to S. ayrozai, especially
in features of the pupal gill, but the former species has the gill branches narrower and
slightly longer than the ventral branch and a thickened anterior margin on the cocoon.
Simulium microbranchium differs from S. ayrozai slightly in the larva, such as more greenish
coloration, a posterior circlet with 194-206 hooklet rows, and rectal papillae with 25
lobules.
572 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Simulium (Hearlea) burchi DALMAT (Figs. 123 A, B.)


Simulium (Hearlea) burchi DALMAT, 1951: 44 (1): 42-47; 1955: 72, 81, 76, 82, 239-244; COSCARÓN et al.,
2004: 11-12.
Female: Wing length 2.9-3.1 mm. General coloration brown-blackish. Frons and
clypeus light brown with silver pollinosity; scutum velvety brown-blackish, with 1+1
silvery submedian vitta not joining anteriorly with 1+1 subquadrangular silvery pruinose
spots and posteriorly with whitish pruinose prescutellar area; lateral and anterolateral
edges with silver bands; scutum pilosity dense, golden to silvery greenish. Pleurae light
grey-brown, with silver pollinosity. Legs yellowish brown, with apices of femora, tibiae,
and tarsomeres dark brown. Wing veins pale brown. Abdomen brownish, with tergite II
silvery; tergites VI-IX waxy. Cibarium concave medially, slightly indented. Sc pilose.
Genitalia structurally as in S. capricorne.
Male: Wing length 3.0-3.1 mm. Scutum velvety black edged by silvery band along
periphery with some light covering of anterior 1/4 area. Legs as in female, but apices of
femora, tibiae, and tarsomeres black. Sc without setae. Length/width ratio of hind
basitarsus = 3.9. Genitalia with gonocoxite having rounded internal protuberance;
gonostylus elongate, ratio of gonostylus length/gonocoxite length = 2.1; ventral plate
with small median carina similar to that of S. delatorrei; endoparameres with numerous,
strong hooks.
Pupa: (Figs. 123 A, B). Cocoon shoe-shaped, with weak texture and reinforced
anterior edge. Length basally 3.8 mm, dorsal length 2.8 mm. Frontoclypeus smooth,
lacking granules. Thorax with moderate number of tubercles. Gill strongly annulated,
especially dorsally, comprised of 3 primary branches; anterodorsal branch largest (length
= 1.4-1.5 mm), annulated, distally capitate, with small tubercles dorsally and apically;
ventrolateral branch medium-sized (length = 1.0 mm), flattened, with peculiar longitudi-
nal crest along internal margin; posterodorsal branch small, 1/5 or less length of other
branches. Tergite X without terminal hooks.
Larva: Length (mature) 7.3 mm. General coloration grey to tan. Cephalic apotome
with narrow dark median stripe basally. Cephalic fan with 46-54 primary rays. Antenna
surpassing stalk of cephalic fan; ratio of proximal, medial and distal antennal articles =
1.0:1.5:1.0. Hypostomium with median and corner teeth longer than others, 14-15 lateral
setae per side, and 3+3 discal setae. Postgenal cleft dome shaped, with narrow median
anterior incision; ratio of length of hypostomium/hypostomial bridge = 1.1:1.0. Lateral
plate of thoracic proleg with 47-50 setae. Anal ring with 172-178 rows of 20-24 hooks.
Rectal papillae with about 78 diverticula, 22 on dorsal (central) lobe and 28 diverticula on
each lateral lobe. Anal sclerite simple.
Distribution: Guatemala: El Quiché, Huehuetenango.
Bionomics: Simulium burchi occurs in medium-sized torrential creeks on stones or
leaves and twigs.
Discussion: This species resembles S. dalmati based on the pupal gill, but the gill of
S. dalmati has a more uniform, only slightly distally expanded, dorsal branch and a longer
posterior branch. Furthermore, the cocoon of S. burchi is boot-shaped and more coarsely
woven. In females of S. dalmati, the 1+1 silver anterior spots are subtriangular and join
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 573

vittae that extend further posteriorly. The original description of S. burchi states that the
vittae join the anterior 1+1 subquadrangular spots, but in the two pinned paratypes from
the USNM, the vittae do not fully join the anterior silvery spots. Larvae of S. dalmati differ
from those of S. burchi by their larger size, longer hypostomial bridge (index of 0.8), and
smaller number of diverticula on the rectal papillae.

Simulium (Hearlea) canadense HEARLE (Fig. 122 C.)


Simulium canadense HEARLE, 1932: 14 (as Simulium virgatum ssp.); 1932: 15.
Simulium (Hearlea) canadense: VARGAS & DÍAZ NÁJERA, 1957b: 262-284; DÍAZ NÁJERA & VULCANO,
1962: 110; PETERSON, 1981: 381; PETERSON & KONDRATIEFF, 1995: 33-34; COSCARÓN et al.,
2004: 12-14.
Simulium (Hemicnetha) canadense: ADLER et al., 2004: 369-370.
Female: Wing length 3.4-3.8 mm. General coloration dark grayish. Scutum blackish
grey, clothed in closely adpressed silvery to greenish pilosity, with longitudinal submedian
silvery vittae joining with 1+1 subtrapezoidal silvery anterior spots (changing to black
with opposite light), lateral and posterior areas silvery grey pollinose; scutellum and
metanotum brownish black; pleurae grayish pollinose. Abdomen grayish black, with lateral
margin of tergite II and posterior 1/5 of tergites III-V silvery grey pollinose; tergites III-
V blackish, subquadrate. Frons convergent below; fronto-ocular triangle about as wide as
high. Sensory vesicle of palpus about 1/2 of basal palpomere length, with abundant
tubercles. Mandible with 12-27 teeth. Lacinia with 12-14 retrorse teeth. Basal portion of
cibarium thick, smooth, medially concave, with 1+1 ovoid lateral prominences. Sc with 13
setae. Length/width ratio of hind basitarsus 5.7-6.4. Sternite VIII darkened medially, with
about 14-20 hairs per side. Gonapophysis subtriangular, concave internally and blunt
distally; cercus curved distally, paraprocts subrectangular, with relatively abundant hair
internally and transverse sulcus subbasally; genital fork with median stem capitate basally
and with lateral arms wide, bearing stout anteriorly directed apodemes; spermatheca
ovoid, homogeneously sclerotized.
Male: Wing length 3.1-3.3 mm. Scutum black, with anterior 1/3 light grayish
pollinose, separated by black area and surrounding 1+1 subtriangular silvery spots.
Length/width ratio of hind basitarsus = 3.8. Gonostylus about 1.8 times gonocoxite
length, with small internally directed basal process; gonocoxite elevated laterally; ventral
plate with median carina; endoparamere with strong hooks.
Pupa: Cocoon slipper shaped, coarsely woven, reinforced along anterior edge. Gill
membranous with 3 subsegmented branches; anterodorsal branch largest, blunt apically,
ventral branch directed anteriorly, roughly 1/2 width of dorsal branch and digitiform
distally (Fig. 122 C); posterior branch 1/4 length of other branches, curved posterome-
dially. Frontoclypeus and thorax with abundant small tubercles. Cephalic and thoracic
trichomes single, stout. Abdomen with spines reduced.
Larva: Length (mature) 8.0-9.0 mm. Cephalic apotome without ornamentation, head
ventrally darkened basomedially. Cephalic fan with 48-56 primary rays. Ratio of length of
hypostomium/hypostomial bridge = 1:1. Hypostomium with anterior margin straight,
median tooth longer than corner teeth, 11-15 lateral setae per side, and 2-4 discal setae.
574 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Postgenal cleft dome shaped, with anterior incision. Mandible with internal teeth arranged
in 2-3 rows of 5-8 teeth, 2 thickened marginal teeth, and 1 large latero-mandibular
process. Ratio of proximal, medial and distal antennal articles = 1:1.3-1.4:1.0-1.2; medial
article with 3-4 subdivisions. Lateral sclerite of thoracic proleg with 24 teeth. Anal sclerite
with hair among struts. Anal ring with 102-158 rows of 18-20 hooks. Rectal papillae with
15-24 diverticula (5-8 per lobe), dorsal diverticulum of each lobe greatly enlarged.
Distribution: Canada: British Columbia; USA: Arizona, California, Colorado,
Florida, Idaho, Montana, Nevada, New Mexico, Oregon, South Dakota, Utah, Washing-
ton, Wyoming; Mexico: Distrito Federal, Durango, Mexico, Oaxaca, Puebla, Veracruz.
Bionomics: Larvae are found on vegetation in cool, clear streams.
Discussion: This species is structurally similar to S. burchi and S. dalmati but the
former species has a shoe-shaped cocoon and a more strongly capitate, tuberculed,
anterior gill branch; the latter species has both major gill branches multilobed apically.

Simulium (Hearlea) capricorne DE LEÓN (Figs. 120 D-J; 121 B; 122 D, E; 126 A-C; 127 A.)
Simulium capricornis DE LEÓN, 1945: 71-72.
Simulium (Hearlea) capricornis: DALMAT, 1950: 72, 76, 82, 86; l955, 244-249; COSCARÓN, 1987: 34;
COSCARÓN et al., 2004: l4-l6.
Simulium (Simulium) deleoni VARGAS, 1945b: 72.
Simulium (Hearlea) deleoni: VARGAS & DÍAZ NÁJERA, 1957b: 294; VARGAS et al., 1946: 182.
Female: Wing length 2.7-2.8 mm. General coloration brownish dark. Head blackish,
eyes dark brown, frons and clypeus brown, with silver pruinosity; scape and pedicel light
brown, flagellum dark grayish brown, palpus and proboscis dark brown to blackish;
palpus with long blackish hair. Scutum dark brown to black, with faint grayish pollinosity,
uniformly distributed recumbent hair, and 1+1 anterior submedial, subtrapezoidal black
spots not joined to 1+1 silvery vittae; prescutellar area and lateral edges of scutellum with
abundant, long golden hair, pleurae brown-grey pollinose; metanotum brown, silvery
tomentose. Wing veins light brown. Legs light brown, with darkened apices on femur,
tibiae and tarsomeres, tarsomeres of fore leg black. Abdomen dark brown; tergites II-V
with dark brown median spots, tergite II with 1+1 silvery spots. Frons convergent below
(Fig. 120 D); fronto-ocular triangle wider than high (Fig. 120 E). Basal portion of
cibarium slightly wrinkled along median edge, with 1+1 submedian, subovoid prominenc-
es (Fig. 120 G). Mandible with 11+27 teeth. Lacinia with 13+17 retrorse teeth. Sensory
vesicle of maxillary palpus slightly shorter than length of basal palpomere. Basal sector of
R without hair; Sc with about 15 hairs. Length/width ratio of hind basitarsus = 5.2:1.0
(Fig. 120 F). Sternite VIII well sclerotized, with about 14 hairs per side; gonapophysis with
distal edge externally curved and medially directed apices (Fig. 120 H). Cercus flattened
distally. Paraproct subrectangular, with internal edge curved and with small longitudinally
positioned crest (Fig. 120 I); genital fork (Fig. 120 J) with median stem long, distally
capitate and lateral arms only moderately expanded apically.
Male: Wing length 2.6-2.8 mm. General coloration same as for female but slightly
darker. Scutum velvety black, with anterior 1+1 wide silver pruinose spot continuous with
thin lateral silvery stripe that joins silver prescutellar area. Legs darkened, especially
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 575

A B K L

D J F

E
C

G I

N
M
O

H
P

Figure 120. Female. A-B: S. (Hearlea) juarezi; A: scutum with front illumination; B: scutum with posterior
illumination. C: S. (H.) delatorrei, cibarium. D-J: S. (H.) capricorne; D: frons; E: fronto-ocular triangle; F:
hind basitarsus; G: cibarium; H: eighth sternite and gonapophysis; I: cercus and paraproct; J: genital fork.
K-L: S. (H.) microbranchium; K: scutum with frontal illumination; L: scutum with posterior illumination.
M-P: S. (H.) larvispinosum; M: cibarium; N: claw; O: genital fork; P: spermatheca.
576 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

0.5 mm

0.05 mm
A

C
E
0.1 mm

0.05 mm

0.05 mm
G

F B 0.1 mm

0.1 mm

Figure 121. A-G: Male; A: S. (Hearlea) juarezi, scutum; B: S. (H.) capricorne, hind basitarsus; C-E: S.
(H.) delatorrei; C: gonocoxite and gonostylus; D: ventral plate; E: endoparameres and median
sclerite; F-G: S. (H.) larvispinosum; F: gonocoxite and gonostylus; G: ventral plate. H-I: Pupa,
frontoclypeus of S. (H.) capricorne and S. (H.) delatorrei, respectively.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 577

A 1 mm

1 mm

0.4 mm

H
C

E
F

0.3 mm

I
J

L
N

Figure 122. Pupa and gill. A-B: S. (Hearlea) ayrozai; A: lateral view; B dorsal view. C: S. (H.)
canadense. D-E: S. (H.) capricorne; D: lateral view; E: ventral view. F-I: S. (H.) ethelae; F: lateral view; G:
lateral view; H: dorsal view; I: lateral internal view. J-K: S. (H.) microbranchium; J: lateral view; K:
dorsal view. L-N: S. (H.) nigricorne; L: lateral view; M: latero internal view; N: antero dorsal view.
578 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

prothoracic and mesothoracic coxae and most of metathoracic leg. Abdomen dark brown,
with 1+1 silvery spots on tergite II. Sc bare. Hind basitarsus wide, length/width ratio = 3.0-
3.2 (Fig. 121 B). Gonocoxite subquadrate, with posterior prominence. Gonostylus elongate,
about twice as long as gonocoxite, with internal row of setae basally, and stout subapical
spur; ventral plate with sinuous posterior margin and short longitudinal median carina.
Pupa: (Figs. 122 D, E). Cocoon slipper shaped, coarsely woven, reinforced along
anterior edge; basal length 3.5-4.3 mm; dorsal length 3.3-4.1 mm. Frontoclypeus with only
lateral and ventral edges having few smooth apically rounded tubercles (Fig. 120 H) and
2+2 frontal and 1+1 facial simple trichomes. Anterodorsum of thorax with scarce to
abundant acuminate granules and 5 stout, simple trichomes per side. Gill with 2 main
branches, dorsal and ventral; branches thickened, subannulated, dorsoventrally flattened,
strongly curved apically; dorsal branch largest, with strongly downturned apex; ventral
branch curved along contour of cocoon. Maximum length of branch = 2.2 mm.
Abdomen with spine-combs on tergites VII and VIII. Terminal spine absent.
Larva: Length 7.3-7.7 mm. Coloration yellowish (in alcohol). Head light brown;
cephalic apotome brownish and slightly darkened basally, without special ornamentation.
Cephalic fan with 42-47 rays. Hypostomium with straight anterior margin, median tooth
longer than corner teeth (Fig. 126 B), 8-10 lateral setae per side, and 1+1 to absent discal
setae. Hypostomial bridge longer than hypostomium, ratio of length of hypostomium/
hypostomial bridge = 0.8-1. Postgenal cleft mildly curved, without apical incision (Fig.
126 A). Antenna longer than stalk of cephalic fan; medial article annulated; ratio of
proximal, medial and distal articles = 1:1.1-1.5:1.0-1.4. Mandible with 2 rows of 4-5
internal teeth, 2 marginal teeth, and 1 curved lateral mandibular process (Fig. 126 C).
Lateral sclerite of thoracic proleg with about 38 teeth arranged in 12-14 groups. Anal
sclerite with sparse hair, ventral branches relatively elongate (Fig. 127 A). Anal ring with
115-135 rows of 20-24 hooks. Rectal papillae trilobate, and 4-6 diverticula per lobe.
Distribution: Guatemala: Alta Verapaz, Chimaltenango, El Quiché, Guatemala,
Huehuetenango, Sacatepequez, Sololá, Totonicapán; Mexico: Chiapas, Distrito Federal,
Durango, Mexico, Morelos, Oaxaca, Veracruz.
Bionomics: Unknown.
Discussion: The closest species to S. capricorne is S. estevezi, which can be
differentiated by the absence of an anterior collar on the cocoon, presence of a process
on the dorsal gill branch, and a cibarium with a thickened anterior edge; and S. burchi,
which has a wide apically blunt dorsal branch, as does S. canadense. Pupal gill morphology,
as used in the accompanying key, permits separation of S. capricorne from structurally
similar species such as S. ethelae, S. dalmati and S. microbranchium.

Simulium (Hearlea) chiriquiense FIELD


Simulium (Hearlea) chiriquiense FIELD, 1967b: 194-199; VARGAS et al., 1977: 137; COSCARÓN et al.,
2004: 16-17.
It was not possible to examine topotypical material of this species. Based on morphology
of the pupal gill in the original description, it could be considered conspecific with S. ethelae
DALMAT. Material from Costa Rica identified as S. chiriquiense by VARGAS et al. is similar to that
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 579

of S. capricorne, based on illustrations of the pupa (Figs. 3, 4). Genitalia of adults cannot be
reliably differentiated from those of several other Simulium (Hearlea) species.
Distribution: Panama: Cerro Punta (Boquete), Chiriqui; Costa Rica: Vicinity of
San José.
Bionomics: Unknown.

Simulium (Hearlea) contrerense DÍAZ NÁJERA & VULCANO


Simulium (Hearlea) contrerense DÍAZ NÁJERA & VULCANO, 1962: 94-99; COSCARÓN et al., 2004: 17.
(Description based on DÍAZ NÁJERA & VULCANO, 1962).
Female: General coloration blackish. Scutum velvety blackish, with 1+1 silvery
vittae not joining anteriorly with 1+1 submedian silvery triangular spots; lateral and
posterior edges silvery pruinose. Abdomen brownish, with posterior edge of tergites
yellowish. Basal portion of cibarium concave medially, without wrinkled margin and thick
rim. Genitalia similar to those of S. capricorne.
Male: Wing length 5.0 mm. General coloration blackish. Scutum velvety black,
bordered with whitish pruinosity and clothed in golden pile. Scutellum brown, with light
brown posterior edge. Abdominal tergites black, with yellowish hair. Sc haired. Gonocox-
ite, gonostylus and endoparameres typical for subgenus (as in S. capricorne); ventral plate
with prominent median carina.
Pupa: Length (basal) 7.0 mm; dorsal 6.0 mm; gill 3.5 mm. Cocoon shoe shaped,
gradually elevated, closed basally on anterior 1/4, compactly woven, not reinforced along
anterior margin. Thorax with 5 simple trichomes per side. Gill inflated, with 2 major
branches frontally and 1 small ventrolateral branch; dorsal branch stout, subglobulose,
tapered distally, sclerotized and wrinkled dorsally, ventral branch membranous and
translucent ventrally, with subconical lateral prominence, and subannulations; ventrolater-
al branch small, thin, and curved medially.
Larva: Length 12.0 mm. Head brownish, body yellowish grey, without ventral
tubercles. Cephalic fan with 42 primary rays. Hypostoma with 15-17 lateral setae per side
and 4+4 discal setae. Mandible with 6-7 internal and 2 marginal teeth. Ratio of proximal,
medial and distal antennal articles = 1:2.2:1.9. Postgenal cleft subtriangular. Anal ring with
185 rows of about 16 hooks. Rectal papillae with 60-66 diverticula, the dorsal lobe largest.
Distribution: Mexico: Distrito Federal., Mexico (3000 m).
Bionomics: Unknown.
Discussion: The unique gill, with its inflated dorsal and ventral branches, membra-
nous and translucent ventrally, distinguishes this species from all others. Simulium juarezi
has a similar gill, but only its dorsal branch is markedly inflated.

Simulium (Hearlea) dalmati VARGAS & DÍAZ NÁJERA (Figs. 123 J-M.)
Simulium (Hearlea) dalmati VARGAS & DÍAZ NÁJERA, 1948b: 337-341; 1957b: 166, 169, 170, 171, 173,
290; DÍAZ NÁJERA & VULCANO, 1962: 93, 122; COSCARÓN et al., 2004: 18.
Female: (Diagnosis of female based on VARGAS & DÍAZ NÁJERA, 1948b). Wing
length 3.5 mm. General coloration blackish, with golden pilosity. Scutum black with
580 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

F E
0.5 mm

I G
H

L K

B
A
1 mm

Figure 123. Pupa and gill. A-B: S. (Hearlea.) burchi; A: lateral view; B: dorsal view. C-F: S. (H.)
delatorrei; C: lateral view; D: dorsolateral view; E: dorsal view; F: lateral internal view. G-I: S. (H.)
estevezi; G: lateral view; H: dorsal view; I: ventral view. J-M: S. (H.) dalmati; J: lateral view; K: dorsal
view; L: frontal view; M: ventral view. N-O: S. (H.) juarezi; N: lateral view; O: dorsal view.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 581

1+1 silvery vittae, joining anteriorly with 1+1 silvery subtriangular spots. Legs
yellowish, with apices of femora, tibiae, and tarsomeres black. Basal portion of
cibarium smooth, with thickened anterior margin gradually elevated sublaterally. Sc
haired. Genitalia as in S. capricorne.
Male: Wing length 3.0 mm. Coloration similar to that of female. Scutum black, with
grayish pruinosity on anterior 1/3 interrupted medially. Sc with scarce hair. Length/width
ratio of hind basitarsus = 3.6. Ventral plate with short, haired, median carina.
Pupa: (Fig. 123 J). Cocoon slipper shaped coarsely woven, anterior edge reinforced.
Length at base 5.0 mm. Gill (Figs. 123 K-M) inflated, pseudosegmented, comprised of 3
branches; dorsal and ventral branches subequal in size; posterior branch 1/4 or less length
of other branches; dorsal branch 1.4 mm, subcylindrical, capitate, directed anterolaterally,
with 3-4 small apical protuberances (Fig. 123 K); ventral branch 1.3 mm, laterally
flattened, blunt apically, curved anteromedially, with 1 or more small apical protuberances
(Fig. 123 L); posterior branch 0.4 mm, acuminate, directed posteromedially (Fig. 123 K).
Frontoclypeus and thorax with sparse and dissimilarly sized tubercles.
Larva: Length 9.0 mm. Coloration grayish green; cephalic apotome without head
spots, slightly darkened mediobasally. Cephalic fan with 42-44 primary rays. Antenna as
long as stalk of cephalic fan, median article with 2 whitish subannulations. Mandible with
strong curvature on superior side, with 6 internal and 2 marginal teeth, the last well
developed; ratio of proximal, medial and distal antennal articles = 1:1.3-1.4:1.0-1.1.
Postgenal cleft gradually curved, with narrow apical incision. Hypostoma with straight
anterior margin, 11-13 lateral setae per side, and 3+3 discal setae; median tooth longer
than others; ratio of hypostomium/hypostomial bridge = 0.8. Anal ring with 144 rows of
21-23 hooks each. Rectal papillae with 18-24 diverticula, consisting of large dorsal and 5-
7 smaller ventral diverticula per lobe.
Distribution: Mexico: Veracruz.
Bionomics: Unknown.
Discussion: The female is very similar to those of S. delatorrei and S. larvispinosum.
The pupa is structurally close to S. canadense and S. burchi, but S. canadense has a slipper-
shaped cocoon and S. burchi has a more anteriorly erect cocoon with a finer textured
weave. The larva is similar to that of S. ethelae, but has 36-39 lobules on the rectal papillae.
The larva of S. delatorrei is also similar, but it has 114 rows of hooks on the posterior
circlet and 7-9 lateral hypostomial setae per side.

Simulium (Hearlea) delatorrei DALMAT (Figs. 120 C; 121 C-E, I; 123 C-F.)
Simulium (Dyarella) delatorrei DALMAT, 1950: 137-147.
Simulium (Hearlea) delatorrei: DALMAT, 1951: 53; 1955: 253-258; VARGAS & DÍAZ NÁJERA, 1957b: 170,
172, 174; DÍAZ NÁJERA & VULCANO, 1962: 93; COSCARÓN et al., 2004: 19-20.
Female: Wing length 3.1 mm. General coloration dark brown, clothed with yellow to
greenish pile, very similar to S. capricorne; scutum with 1+1 submedian silvery vittae extending
from subtriangular anterior silvery spots to silvery pollinose prescutellar area, changing from
whitish silver to black depending on lighting. Frons convergent below, fronto-ocular triangle
about as high as wide. Basal portion of cibarium smooth, with thin anterior edge and small
582 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

median concavity (Fig. 120 C). Sensory vesicle of maxillary palpus with abundant tubercles,
about 1/2 of basal article length. Genitalia similar to those of S. capricorne.
Male: Wing length 3.0 mm. General coloration similar to that of female. Eyes dark
brown; scutum dark brown, bordered by silver pruinosity with anterior and posterior areas
wider than lateral areas, scutum with abundant closely appressed silvery pilosity; (in
specimens without pilosity, light position enables vittae to be seen on grey tomentous
pruinosity); pleurae brown with silver pollinosity. Legs brownish, with long yellowish
brown hair. Abdomen brown blackish, with 1+1 silver spots laterally on tergites VI-VIII.
Length/width ratio of hind basitarsus = 3.3. Genitalia as in figures 121 C-E.
Pupa: Cocoon slipper shaped, closed anteriorly, compactly woven, threads not evident,
reinforced on anterior edge (Figs. 123 C-F). Length (basal) 3.2-3.3 mm; (dorsal) 2.5-2.8 mm.
Frontoclypeus with abundant, small tubercles laterally, continued internally by microscopic
punctation, with 2+2 thin frontal and 1+1 stout and spine-like simple facial trichomes (Fig.
121 I). Thorax dorsally with abundant and minute tubercles more developed and conical
along midline and with 5 spine-like trichomes per side. Gill 1.8 mm, comprised of 3
branches; filaments stout, gradually tapered, pseudosegmented; dorsal branch largest,
subcylindrical, apically blunt, directed anteriorly, with noticeable ventrolateral protuberance
submedially; ventral branch 1/2 width of dorsal branch, curved medially; posterior branch
smallest, directed posteromedially. Abdomen lacking terminal hooks.
Larva (description based on DALMAT, 1955): Body length 7.0 mm. General color
grey, cephalic apotome darkened mediobasally. Cephalic fan with 39-40 primary rays.
Postgenal cleft dome shaped, with deep anteromedian incision. Hypostomium with
anterior margin straight and 7-9 lateral setae per side. Ratio of hypostomium/hypostomial
bridge = 1.1. Mandible with 6 internal teeth disposed in 3 rows, 2 marginal teeth, second
slightly longer than 1/2 length of first. Lateral sclerite of thoracic proleg with 40-42 comb
teeth. Anal ring with 114 rows of 19-22 hooks each. Rectal papillae with 72-75 diverticula,
24-25 per lobe.
Distribution: Guatemala: Chimaltenango, El Quiché, Guatemala, Huehuetenango,
Quezaltenango, San Marcos, Sololá, Totonicapán; Mexico: Chiapas.
Bionomics: Unknown.
Discussion: The closest species is S. estevezi but differences in the pupa and larva
enable their separation. The Díaz & Vulcano (1962) illustration of larvae from Mexico
shows that the anal papillae have only 5 lobules per lobe and 8 lateral hypostomial setae,
differing from Guatemalan populations.

Simulium (Hearlea) estevezi VARGAS (Figs. 123 G-I.)


Simulium (Hearlea) estevezi VARGAS, 1945b: 71; VARGAS & DÍAZ NÁJERA, 1948b: 341; DÍAZ NÁJERA &
VULCANO, 1962: 92; COSCARÓN et al., 2004: l9-20.
(Diagnosis of female and male based on VARGAS, 1945b and VARGAS & DÍAZ
NÁJERA, 1957b).
Female: Wing length 2.5 mm. Scutum blackish, with 1+1 silvery vittae joined with
anterior subtriangular silvery spots. Cibarium without ovoid sublateral prominences.
Genitalia structure similar to that of S. capricorne.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 583

Male: Wing length 2.7 mm. Genitalia structurally similar to that of S. capricorne.
Ventral plate with haired median carina.
Pupa: Length (basal) 3.6 mm; (dorsal) 2.8 mm. Cocoon shoe shaped, compactly
woven, threads not evident, slightly reinforced anterior edge. Frontoclypeus covered by
platelets; thorax with abundant tubercles. Gill with 3 acuminate, pseudosegmented
branches; dorsal branch largest, subcylindrical, directed anteroventrally, with median,
internally directed prominence, and sharply upturned acuminate apex; ventral branch
laterally flattened, curved ventromedially; posterior branch 1/3 or less length of other
branches, directed posteromedially (Figs. 123 G-I).
Larva: Length 8.0 mm. General coloration grayish. Cephalic apotome uniformly
yellowish colored, or with slightly darkened spots forming cross. Antenna extended
anteriorly beyond apex of cephalic fan stalk; ratio of proximal, medial and distal articles =
1:1.3-1.4:1.1-1.2; medial article appearing subsegmented into 3 pieces. Mandible with 2
rows of 7 internal teeth, and 2 internal teeth, the anterior one larger. Cephalic fan with 44-
50 primary rays. Postgenal cleft gradually narrowed apically, with small apical incision.
Hypostoma with anterior margin straight. Ratio of hypostomium/hypostomial bridge =
0.9. Hypostomium with 8 lateral setae per side and 2+2 discal setae. Lateral plate of
thoracic proleg sclerite with about 40 setae. Anal ring with 98-104 rows of 18-19 hooks
each. Rectal papillae with 24-30 diverticula, 8-10 per lobe.
Distribution: Mexico: Chiapas, Distrito Federal, Guerrero, Oaxaca, Veracruz.
Bionomics: Unknown.
Discussion: Externally, the female is very close to those of S. ayrozai, S. dalmati and
S. ethelae. The pupa of these species, however, is quite distinct. The pupa of S. estevezi is
closest to that of S. delatorrei, but it has the dorsal gill branch thicker and blunt apically, the
medioventral process smaller, and a more basally flattened cocoon. The larva of S. estevezi
is closest to that of S. delatorrei, but the anal ring has 114 rows of hooks, the rectal papillae
have 24-25 diverticula per lobe, and the postgenal cleft is significantly shorter.

Simulium (Hearlea) ethelae DALMAT (Figs. 122 F-I.)


Simulium (Hearlea) ethelae DALMAT, 1950: 143-148; 1951: 52-53; 1955: 258-263; VARGAS & DÍAZ
NÁJERA, 1957b: 167, 171, 174, 298 (as Simulium capricorne); DÍAZ NÁJERA & VULCANO, 1962: 93;
COSCARÓN et al., 2004: 21-22.
Female: Wing length 3.6 mm. Scutum velvety black with 1+1 anterior silvery vittae
joining 1+1 anterior silver pruinose subtriangular spots and light grayish prescutellar area.
Basal portion of cibarium smooth, concave medially, with thickened margin bearing 1+1
lateral subovoidal prominences. Length/width ratio of hind basitarsus = 4.7:1.0. Genitalia
as in S. capricorne.
Male: Wing length 3.4 mm. Coloration of scutum velvety blackish with silvery
pruinose spot on anterior 1/4; abdominal ornamentation and genitalia morphology
similar to those of other S. (Hearlea) species, with ventral plate subovoidal with median
carina. Length/width ratio of hind basitarsus = 3.3.
Pupa: Length (basal) 2.4 mm. Cocoon slipper-shaped, closed anteriorly, compactly
woven, threads not evident, with anterior reinforcement. Gill with 2, subequally sized,
584 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

wrinkled, curved, gradually flattened, pseudosegmented, acuminate branches lacking


granules or spicules (Figs. 122 F-I); dorsal branch slightly longer than ventral branch, with
apex curled ventrally. Thorax with sparse acuminate tubercles; frontoclypeus mostly
smooth, with rounded tubercles only along margins.
Larva: Length 7.0 mm. General coloration grayish. Cephalic apotome with only
small spots on midline and submedian area. Cephalic fan with 36-40 primary rays.
Mandible with 5 or 6 internal teeth and 2 thin marginal teeth. Hypostoma with 12-16
lateral setae per side. Postgenal cleft narrow distally, deeply incised; ratio of hypostomi-
um/hypostomial bridge = 0.9. Lateral sclerite of thoracic proleg with 36-42 teeth. Rectal
papillae with 36-39 diverticula, 12-13 diverticula per lobe. Anal ring with 150-170 rows of
18-20 hooks each.
Distribution: Guatemala: Chimaltenango, Guatemala, Huehuetenango, Quezalt-
enango, Solalá; Mexico: Chiapas.
Bionomics: Unknown.
Discussion: The closest species morphologically is S. capricorne, but the dorsal gill
branch is more flattened and apically blunt and has a median crest and a small
protuberance dorsobasally. The female differs in that the scutal vittae do not join the
anterior subtrapezoidal spots. Simulium chiriquiense has an acuminate dorsal gill branch,
frontoclypeus with numerous tubercles, and wing length (female 3.8 mm and male 3.0
mm) more similar to that of S. ethelae.

Simulium (Hearlea) juarezi VARGAS & DÍAZ NÁJERA (Figs. 120 A, B; 121 A; 123 N, O.)
Simulium (Hearlea) juarezi VARGAS & DÍAZ NÁJERA, 1957b: 205, 262, 304; DÍAZ NÁJERA & VULCANO,
1962: 100; COSCARÓN et al., 2004: 22-23.
Female: Wing length 3.3-3.8 mm. General coloration blackish brown. Head brown,
with frons and clypeus grey pollinose. Scape and pedicel light brown; flagellum dark
brown. Palpus and proboscis dark brown. Scutum brown, with grayish pruinosity, and
yellowish-green to brown decumbent hair varying in color with light position. Scutum
with 1+1 submedian vittae that are silver whitish with posterior lighting and blackish with
anterior lighting, extending from 1+1 subquadrate anterior spots to prescutellar area;
median blackish stripe about 3 times wider than silvery submedian stripes (Figs. 120 A, B);
humeral spots light grey pollinose as are narrow lateral edges and prescutellar area;
abdomen dark brown. Legs yellowish brown, with apices of femora, tibiae, and
tarsomeres blackish. Basal portion of cibarium smooth, with thickened rim, and 1+1
subovoidal sublateral prominences. Genitalia similar to those of S. capricorne.
Male: Wing length 3.2-3.6 mm. General coloration as in female but more darkened.
Scutum velvety blackish, with anterior 1/3 grayish pollinose and separated medially by
blackish area (Fig. 121 A). Legs mostly dark brownish, with closely adpressed yellowish
brown short hair and blackish long hair. Length/width ratio of hind basitarsus = 4.0.
Genitalia structurally similar to that of S. capricorne.
Pupa: Cocoon slipper-shaped, closed anteroventrally, with reinforced anterior edge,
compactly woven, frequently covered by sand particles. Length (basal) 3.4-4.0 mm;
(dorsal) 3.2-3.5 mm. Frontoclypeus and anterodorsum of thorax with abundant, rounded,
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 585

variably sized tubercles. Gill with 3 semi-membranous branches; dorsal branch largest,
strongly clavate, with numerous tubercles dorsally and annulations basally; ventral branch
laterally flattened, annulated, strongly curved medially; posterodorsal branch acuminate,
subannulated, only slightly shorter than ventral branch, and directed dorsomedially. (Figs.
123 N, O).
Larva (diagnosis based on DÍAZ NÁJERA & VULCANO, 1962): Length 7.5 mm. General
coloration greenish dorsally and yellowish ventroapically. Cephalic apotome darkened
basally and medially. Cephalic fan with 50 primary rays. Hypostoma with 8-10 lateral setae
per side. Postgenal cleft dome shaped, narrowed distally. Anal ring with about 145 rows of
17 hooks. Rectal papillae with 5-7 diverticula per lobe, and median anterior diverticulum
larger than others.
Distribution: México: Chiapas, México, Michoacán, Oaxaca, Veracruz.
Bionomics: Unknown.
Discussion: The peculiar respiratory organ with its large, subglobose, superficially
granulose dorsal branch distinguishes this species. The closest species with inflated gill
branches is S. contrerense.

Simulium (Hearlea) microbranchium DALMAT (Figs. 120 K, L; 122 J, K.)


Simulium (Simulium) microbranchium DALMAT, 1949: 538-544.
Simulium (Hearlea) microbranchium DALMAT, 1951: 50-52; 1955: 72, 76, 82, 87, 268-272; COSCARÓN et
al., 2004: 23-24.
Female: Wing length 3.4 mm. Frons and clypeus brownish-grey silvery pollinose.
Thorax brownish black. Scutum brownish dark, with silver submedian vittae not joining
anterior 1+1 silvery subtrapezoidal spots, but joining silvery pollinose prescutellar area;
area between these spots black, although changing light position changes this silver color
to black (Figs. 120 K, L). Scutum with golden, closely adpressed pilosity. Shape and
proportion of palpal sensory vesicle as in S. capricorne. Cibarium with median portion
concave, hyaline, with serrated margins. Sc haired. Length/width ratio of hind basitarsus
= 5.2. Genitalia structurally similar to those of S. capricorne.
Male: Wing length 3.4 mm. Scutum velvety black with whitish pruinosity anteriorly
and bordered with narrow silvery band on periphery. Scutum pilosity golden. Sc haired.
Genitalia typical for subgenus; ventral plate with small median crest. Gonostylus about
twice length of gonocoxite height at area of insertion; gonocoxite externally 1/3 longer
than internal height.
Pupa: Cocoon weakly shoe-shaped, closed and slightly enlarged anterobasally, with
reinforced edge (Fig. 122 J). Length (basal) 3.5 mm. Frontoclypeus and anterodorsum of
thorax with sparse tubercles laterally around base of frontoclypeus, and sparse on anterior
area of thorax. Gill comprised of 2 laterally flattened, dorsally pseudosegmented,
acuminate branches (Fig. 122 K); dorsal branch (length = 1.6 mm) directed anteriorly,
forming about 90° angle with gill base; ventral branch (length = 0.9 mm) only slightly
medially directed.
Larva (diagnosis based on DALMAT, 1951): Body length 7.7 mm. General coloration
grey. Cephalic apotome concolorous, with small black spots along midline and submedian
586 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

anterior area. Cephalic fan with 56 or 57 primary rays. Hypostoma with median tooth and
corner teeth longer than other teeth. Postgenal cleft curved with narrow anteromedian
incision. Lateral sclerite of thoracic proleg with 37-39 teeth. Anal ring with 194-206 rows
of hooks. Rectal papillae with about 75 diverticula, about 25 per lobe. Anal sclerite simple,
lacking scales.
Distribution: Guatemala: El Quiché, Huehuetenango, Quetzaltenango, Sololá, To-
tonicapán.
Bionomics: Unknown.
Discussion: This species is very similar to S. ayrozai, but its dorsal gill branch is
projected anteriorly well beyond the anterior margin of the cocoon.

Simulium (Hearlea) nigricorne DALMAT (Figs. 122 L-N.)


Simulium (Hearlea) nigricornis DALMAT, 1950: 148-151; 1955: 76, 82, 273-275; COSCARÓN et al., 2004:
24-25.
(Diagnosis of female and male based on DALMAT, 1955).
Female: General coloration unknown. Cibarium with definite indentation on
internal edge. Cerci about 2 times wider than high with curved edge; paraproct with
abundant hair distally; genital fork with median rod thin, elongate.
Male: General coloration unknown. Gonocoxite with strong prominence basally
along internal edge; ventral plate with small median carina.
Pupa: (Figs. 122 L-N). Cocoon shoe-shaped, with anterior edge reinforced; length
(basal) 2.9 mm; maximum 3.6 mm. Frontoclypeus with sparse tubercles. Thorax with
tubercles. Gill extended well beyond anterior margin of cocoon, comprised of 2
subannulated, flattened branches; dorsal branch 0.9 mm, darkened on distal 3/4,
distended apically, with blunt apex; ventral branch 1.5 mm, curved, elongate, thinning
distally, with small projection distally. Abdominal terminal hooks absent.
Larva: Unknown.
Distribution: Guatemala: Chimaltenango.
Bionomics: Unknown.
Discussion: The presence of two annulated branches places S. ayrozai, S. ethelae and
S. microbranchium close to S. nigricorne, but the short, wide, distally darkened dorsal branch
permits separation.

CAROLINAE species group

Simulium (Hearlea) carolinae DE LEÓN


Simulium carolinae DE LEÓN, 1945: 73-75.
Simulium (Hearlea) carolinae: VARGAS et al., 1946: 122-124, 182; DALMAT, 1955: 72, 76, 82, 86, 249-253;
DÍAZ NÁJERA & VULCANO, 1962: 93, 112; COSCARÓN et al., 2004: 25-26.
Female: Wing length 3.7-3.9 mm. Coloration blackish brown grayish with vittae not
joining 1+1 anterior subquadrate silvery spots, pilosity adpressed, abundant, tin colored;
pleurae light grayish, silvery pollinose; legs yellowish grayish with concolorous hair, with
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 587

apices of femur, tibia, hind basitarsus, and tarsite blackish. Cibarium smooth, concave
medially, with 1+1 sublateral ovoidal prominences. Sc haired. Length/width ratio of hind
basitarsus = 4.8. Genitalia structurally similar to those of S. larvispinosum.
Male: Wing length 3.2-3.6 mm. Scutum velvety black, homogeneously pigmented, or
with light area on anterior 1/5, abundant pilosity tin to golden. Legs generally darkened.
Length/width ratio of hind basitarsus = 2.8. Genitalia similar to those of S. larvispinosum.
Gonocoxite slightly elevated laterally; ventral plate with large median carina.
Pupa: Cocoon slipper shaped, reinforced along anterior edge. Length (basally) 3.0
mm. Thorax covered with numerous tubercles. Gill rigid, similar to that of S. larvispinosum,
except main dorsal branch thicker and blunter apically, secondary branches relatively
short, and ventral branch strongly curved.
Larva: Length 7.8 mm. General coloration light grayish brown. Cephalic apotome
darkened slightly along midline basally. Cephalic fan with 42-44 primary rays. Mandible
with preapical teeth well developed, 5-6 internal teeth arrayed in 2 rows; marginal teeth
with 2 denticles well developed and 1 accessory at half of internal teeth position. Ratio of
proximal, medial and distal antennal articles = 1:1.6-1.8:1.1-1.2. Hypostomium with
prominent median tooth and 11-15 lateral setae per side. Postgenal cleft dome shaped,
only slightly acuminate distally, without deep anteromedian incision. Lateral sclerite of
thoracic proleg with 24 teeth in 7-8 groups. Anal sclerite with ventral branches encircling
posterior circlet. Dorsolaterally on posterior portion of abdomen there are 2+2 plates
flattened and conical, very sclerotized and covered by acuminate tubercles and some
spines with wide base. Anal ring with 190-200 rows of 25-27 hooks each. Rectal papillae
with 24-33 diverticula, 8-11 diverticula per lobe.
Distribution: Mexico: Chiapas, Oaxaca, Veracruz; Guatemala: Alta Verapaz,
Chimaltenango, Guatemala, Quezaltenango, Sololá, Suchitepequez.
Bionomics: Unknown.
Discussion: The form of the pupal gill and absence of 1+1 ventrolateral papillae on
the eighth segment of the larva is shared with S. larvispinosum, but the latter has 1+1
accessory ventrolateral accessory plates on the anal sclerite.

Simulium (Hearlea) gorirossiae VARGAS & DÍAZ NÁJERA (Figs. 124 F, G; 126 D-G; 127 D-F.)
Simulium (Hearlea) gorirossii VARGAS & DÍAZ NÁJERA, 1957b: 199-202; DÍAZ NÁJERA & VULCANO,
1962: 92; COSCARÓN et al., 2004: 26-28 (incorrect original spelling).
Female: Wing length 3.2 mm. Coloration blackish. Head dark brown, with frons and
clypeus silvery pollinose. Scutum velvety black with 1+1 silvery vittae joined with submedian
anterior subtriangular silvery spots; laterally bordered by narrow whitish grey pruinose band
joining whitish prescutellar area. Abdomen blackish. Fronto-ocular triangle about as long as
wide; cibarium with basal portion as in S. larvispinosum, with hyaline edge and 1+1 small
rounded submedian processes. Sc bare. Genitalia typical for subgenus, with cercus
rectangular, flattened distally; paraprocts with a transverse tegumentary fold and very haired
apically; gonapophysis subtriangular, with blunt apex similar to that of S. larvispinosum.
Male: Wing length 3.0 mm. Scutum velvety black with anterior 1/3 pruinose,
continued by narrow band edging laterally and meeting with whitish grey pruinose
588 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

1 mm

C D

E B

0.5 mm

H I
F

G K

Figure 124. Pupa and gill. A-E: S. (Hearlea) larvispinosum; A: lateral view; B: dorsal view; C: lateral
external view; D: latero internal view; E: dorsolateral view. F-G: S. (H.) gorirossiae; F: dorsal view; G:
ventrolateral view. H-I: S. (H.) johnsoni; H: dorsal view; I: lateral external view. J-K: S. (H.) menchacai;
J: lateral view; K: dorsal view.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 589

prescutellar area. Abdomen blackish. Sc bare. Genitalia similar to those of S. larvispinosum,


except ventral plate more elevated medially.
Pupa: Length: basally 2.4-2.7 mm; dorsally 1.5-1.7 mm. Cocoon slipper shaped, not
closed anteriorly and thickly woven, compact with very narrow anterior rim, showing
open weave. Clypeus and thoracic dorsum with moderately to abundant, small and
acuminate tubercles; clypeus and thorax with typical strong single trichomes. Gill very
similar to that of S. larvispinosum, but with large frontolateral branches narrower,
acuminate apically, with spine at tip similar to other branches, first secondary branch more
elongated and insertion area more sclerotized; in dorsal view, frontolateral branches more
open (Figs. 124 F, G). Abdominal terminal hooks absent.
Larva: Length 6.5-7.0 mm. General coloration light yellowish grey. Cephalic
apotome uniformly colored, slightly darkened at base (Fig. 126 D); cephalic fan with 50-
52 primary rays. Antenna longer than stalk of cephalic fan; ratio of proximal, medial and
distal antennal articles = 1:1.4:0.7; medial article with pseudosegmentation (Fig. 126 E).
Mandible with strong curvature on superior side (Fig. 126 F); internal teeth arrayed in 2
rows with 4 teeth, with first and second teeth very curved, 2 marginal teeth, and 1 robust
and long latero- mandibular process, 1/3 of which surpasses inferior edge of mandible
(Fig. 126 G). Hypostoma with anterior margin as in S. larvispinosum, with prominent
median tooth and intermediate teeth longer than corner teeth; 14-21 lateral setae per side.

III
III

V
V

VII
VII

IX

Figure 125. S. (Hearlea) sp., abdominal chaetotaxy of pupa.


590 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Postgenal cleft deep, without anterior acuminate incision; ratio of hypostomium/


hypostomial bridge = 0.9-1.1. Comb of thoracic proleg lateral sclerite with 47-55 teeth.
Abdominal segment VIII with 1+1 lateroventral, apically sclerotized papillae (Figs. 127 D,
F [vp]). Anal sclerite with ventral branch projected distally forming ring around anal ring,
laterally bordered by very sclerotized and spinous accessory plates of 2+2 conical
dorsolateral projections [dd] and 1+1 flattened more expanded, ventrolateral, acuminate
projections (Fig. 127 D [vl]), abundant acuminate tubercles and sparse minute trichomes;
and 1+1 dorsolateral, irregularly bordered, elongate plates positioned diagonally to lateral
lobes of rectal papillae (Fig. 127 E [df]) and 1+1 short and sclerotized ventrolateral papilla
slightly anterior (Fig. 127 F [vp]). Anal ring with 272-282 rows of 30-35 hooks each. Rectal
papillae with 18-30 diverticula, 6-10 per lobe.
Distribution: México: Chiapas, Veracruz; Guatemala: Sololá.
Bionomics: Unknown.
Discussion: Simulium (Hearlea) larvispinosum, S. menchacai, S. carolinae, S. johnsoni and S.
temascalense are very closely related to S. gorirossiae, but they can be distinguished by subtle
differences in the pupal gills and accessory plates of the larval anal sclerite. Simulium
larvispinosum has a pupal gill with the dorsofrontal branch more inflated and blunt apically,
with the first secondary branch a little shorter (Figs. 124 A-D). The anal sclerite and
accessory plates of the abdominal distal area of the larva are similar, but the ventrolateral
plates lack hard spinous projections, and anterodorsal to the rectal papillae are 1+1 porous
rectangular plates (Fig. 127 I) that in S. gorirossiae are more trapezoidal and obliquely
positioned (Fig. 127 E). Simulium menchacai has a very similar larval anal sclerite, accessory
plates and ventrolateral sclerotized papillae, but the pupal gill has stout secondary
branches as long as the dorsal and ventral branches. Simulium temascalense larvae also have
ventrolateral papillae on the eighth segment but they are less sclerotized, the postgenal
cleft lacks an acuminate incision and the projections of the ventrolateral accessory sclerite
anal plates are shorter. Also, the pupa has very reduced secondary gill branches. Simulium
johnsoni, like S. temascalense, has very reduced secondary gill branches, and in the larva, the
projections of the ventrolateral accessory plates are shorter, but S. johnsoni lacks
ventrolateral papillae on segment VIII of the larva. Simulium carolinae has a very similar
pupal gill, but the larva lacks the ventrolateral accessory plates on the distal portion of the
abdomen, and the ventrolateral papillae of the eighth segment. Simulium paracarolinae has
larvae with only 1+1 accessory dorsolateral plates on the distal portion of the abdomen,
and the ventral branches of the anal sclerite are elongated, of subequal diameter, and
enclose the anal ring.

Simulium (Hearlea) johnsoni VARGAS & DÍAZ NÁJERA (Figs. 124 H, I.)
Simulium (Hearlea) johnsoni VARGAS & DÍAZ NÁJERA, 1957b: 203-205; DÍAZ NÁJERA & VULCANO, 1962:
99-100; COSCARÓN et al., 2004: 28-29.
(Diagnosis of female and male based on descriptions from VARGAS & DÍAZ NÁJERA,
1957b and larva from DÍAZ NÁJERA & VULCANO, 1962).
Female: Wing length 3.1 mm. General coloration blackish. Scutum black with 2
submedian silvery vittae not joining 1+1 anterior subtriangular silvery spots. Anterior
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 591

portion of cibarium with thickened anterior edge, medially concave. Sc with 4-6 hair.
Genitalia similar to those of S. larvispinosum.
Male: Wing length 3.1 mm. Coloration blackish, scutum velvety black whitish on
anterior edge. Abdominal tergites blackish. Genitalia as in S. larvispinosum.
Pupa: Length basally 3.1 mm; dorsally 2.9 mm. Cocoon slipper shaped. Gill
membranous with general shape similar to that of S. larvispinosum but secondary branches
shorter and ventral branch more strongly curved; small branches acuminate, with well
sclerotized spine-like apices (Figs. 124 H, I). Frontoclypeus smooth. Thorax dorsally with
abundant small tubercles on median area, and sparse laterally. Posterodorsum of thorax
with small, sparse, acuminate tubercles.
Larva: Length 8.5 mm. Coloration grey. Cephalic apotome uniformly pigmented,
with small thin darkened area mediobasally. Cephalic fan with 54-56 primary rays. Ratio
of proximal, medial and distal antennal articles = 1:1.4:1.0. Mandible with preapical teeth
robust, internal teeth in 2 rows of 5; 2 large marginal teeth and 3; lateral mandibular
processes long and curved. Hypostoma with straight anterior margin and 10-12 lateral
setae per side; ratio of hypostomium/hypostomial bridge = 0.7. Postgenal cleft dome
shaped; abdominal segment VIII without ventrolateral tubercles. Anal ring with 242 rows
of 26-30 hooks each. Anal sclerite with ventral branches encircling anal ring; 2+2
dorsolateral accessory plates with subconical processes, and 1+1 wide lateroventral plates
bearing 4-5 short lateral processes. No ventrolateral papillae present. Rectal papillae with
21 diverticula, 7 per lobe, dorsalmost ones largest.
Distribution: México: Oaxaca, Veracruz.
Bionomics: Unknown.
Discussion: Simulium gorirossiae is very similar to S. johnsoni but the pupal gill has the
anterodorsal branch thinner and more acuminate, with a terminal spine, and the
secondary and ventral branches longer. The larva has longer projections on the accessory
ventrolateral plate, and 1+1 ventrolateral papillae on the eighth segment. Simulium
temascalense has a very similar pupal gill but the dorsal anterior branch is spined apically,
and the larva has ventrolateral papillae on segment VIII and the lateral projections of the
accessory ventrolateral plate are shorter. Simulium larvispinosum has larger secondary
branches and a more curved ventral branch on the pupal gill, and in the larva, 1+1
dorsolateral accessory rectangular plates before the anal papillae.

Simulium (Hearlea) larvispinosum DE LEÓN


(Figs. 120 M, P; 121 F, G; 124 A-E; 126 H; 127 G-I.)
Simulium (Hearlea) larvispinosum DE LEÓN, 1948: 5-23; DALMAT, 1955: 72, 76, 81, 86, 263-268; DÍAZ
NÁJERA & VULCANO, 1962: 93; COSCARÓN et al., 2004: 29-31.
Female: Wing length 3.4 mm. General coloration dark brown. Head blackish, frons
brownish silvery pollinose; scutum velvety brown blackish bordered by narrow silver
pruinose band, with 1+1 narrow (about 1/4 width of median black stripe) silvery vittae,
not reaching to 1+1 anterior subtriangular silvery spots and joining posterior silvery
prescutellar area. Frons with sides convergent below; fronto-ocular triangle about as long
as wide. Cibarium with thickened, medially concave anterior margin bearing 1+1 short,
592 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

smooth submedian processes (Fig. 120 M). Sc with 8-13 hair. Mandible with 10+25
serrations. Lacinia with 13-14 retrorse teeth. Claw with subbasal tooth (Fig. 120 N);
length/width ratio of hind basitarsus = 4.6-4.7. Sternite VIII with 14-16 hairs per side;
genital fork and spermatheca as in figures 120 O, P respectively; cercus subrectangular;
paraproct with longitudinal fold, and thickly haired distally.
Male: Wing length 3.4 mm. Scutum velvety black, bordered by silvery pruinosity, as
are humeral angles. Scutellum black. Abdomen blackish. Sc bare. Length/width ratio of
hind basitarsus = 3.5-3.6. Gonocoxite subquadrate, 1/2 of gonostylus length; gonostylus
with abundant pilosity mediobasally, and subterminal accuminate spur (Fig. 121 F).
Endoparameres with strong hooks; median sclerite wide basally; ventral plate subovoidal,
without evident median carina (Fig. 121 G).
Pupa: Length: basally 3.4 mm. Cocoon slipper shaped, compactly woven, threads
not evident; anterior edge slightly reinforced. Frontoclypeus protruded basally without
platelets but with impression of minute, very abundant tubercles; 1+1 facial and 2+2
frontal single, stout trichomes; thorax with small, acuminate, sparse tubercles on posterior
area and impression of tubercles on remainder of integument, with 5 stout, single, spine-
like trichomes per side. Gill rigid, granulose, cylindrical, with 2 principal branches: dorsal
branch largest, directed anteriorly; ventral branch smaller, curved, and also directed
anteriorly. Main dorsal branch subdivided basally into short secondary branches, the first
of which is short and directed dorsally, the second of which is bifurcate, posteriorly
directed, with 2 or 3 apical branches; the second laterointernal secondary branch also
bifurcate distally; the last frontally directed branch short and apically subdivided (Figs. 124
A-E); except large frontal branch, other branches are sclerotized, and spined apically.
Larva: Length 8.0 mm. Color yellowish to light brown (in alcohol). General aspect as
in figure 127 G. Cephalic apotome homogeneous without special designs, only darkened
on narrow basal stripe; cervical sclerites elongated, reaching edges; ratio of hypostomi-
um/hypostomial bridge = 1.2-1.5. Cephalic fan with 43-48 primary rays. Hypostoma with
median tooth stout, surpassing height of other teeth; corner teeth below intermediate
teeth; 12-17 lateral setae per side, without discal setae, and few lateral serrations. Postgenal
cleft dome shaped with deep projected anteriorly incision (Fig. 126 H). Antenna
surpassing length of cephalic fan stem. Ratio of proximal, medial and distal antennal
articles = 1:1.4-1.5:0.9. Mandible with 4-5 internal teeth; 2 marginal teeth, the second 1/2
length of anterior tooth. Lateral sclerite of thoracic proleg with 45-50 teeth. Anal sclerite
with ventral branch projected basally, forming ring around anal ring. Accessory plates
2+2, subconical, well sclerotized, dorsolaterally and apically positioned (Figs. 127 H, I,
[dd]), 1+1 ventrolateral subconical, subannulate, dentate on external edge (Fig. 127 I [vl])
and 1+1 dorsolateral subrectangular, flattened, anterior to rectal gill, (Figs. 127 H, I [df]).
No ventrolateral papillae present. Anal ring with about 152 rows of 23-26 hooks each.
Rectal papillae with 18-24 diverticula, 6-8 per lobe.
Distribution: Guatemala: Chimaltenango, Sololá, Suchitepequez; Mexico: Chiapas.
Bionomics: Unknown.
Discussion: Simulium johnsoni is very similar to S. larvispinosum, but the pupal gills of
these two species exhibit slight differences, as explained for the former species. Adults of S.
carolinae do not differ externally. The closest species based on the pupa gill is S. gorirossiae, but
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 593

0.2 mm

E
A
0.1 mm

0.0
H
5m
m
0.03 mm

C
0.01 mm

G 0.3 mm
J

L
0.05 mm

D
K 0.1 mm

Figure 126. Larva. A-C: S. (Hearlea) capricorne; A: postgenal cleft and hypostomium; B: hypostomial
dental border; C: mandibular dental area. D-G: S. (H.) gorirossiae; D: cephalic apotome and neck
sclerite; E: antenna; F: mandible, general aspect; G: mandibular dental area. H: S. (H.) larvispinosum,
cephalic capsule, ventral view. I-L: S. (H.) paracarolinae; I: cephalic capsule, ventral view; J:
hypostomial dental border; K: antenna; L: mandibular dental area.
594 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

these species exhibit differences in the larval accessory plates, with the ventrolateral plate
more subquadrate and having long, acuminate projections and the dorsolateral, flattened
plates being positioned diagonally at the sides of the anal gill in S. gorirossiae. Simulium
gorirossiae also differs in that it has sclerotized ventrolateral papillae on the eighth segment.

Simulium (Hearlea) menchacai VARGAS & DÍAZ NÁJERA (Figs. 124 J, K.)
Simulium (Hearlea) menchacai VARGAS & DÍAZ NÁJERA, 1957b: 208-210; DÍAZ NÁJERA & VULCANO,
1962: 101-102; COSCARÓN et al., 2004: 31-32.
Female: Wing length 3.5 mm. General coloration blackish. Thorax blackish brown;
scutum with abundant adpressed silvery pilosity that somewhat obscure submedian silvery
stripes; vittae not joining 1+1 anterior subtrapezoidal silvery spots; scutellum brownish,
metanotum and pleurae blackish; legs yellowish brown. Abdomen brownish with tergites
II-IV pale. Fronto-ocular triangle about as long as high. Metathoracic basitarsus length/
width ratio = 0.5-1.1. Cibarium wrinkled, with anterior edge thickened, translucent,
concave medially and without lateral subovoidal prominences; Sc with 15-17 trichomes.
Genitalia as in S. larvispinosum.
Male: Wing length 3.0 mm. Coloration blackish; scutum velvety black with anterior
area whitish. Sc with 2-3 trichomes. Metathoracic basitarsus length/width ratio = 3.0-3.1.
Genitalia similar to those of S. larvispinosum.
Pupa: Length: basally 3.5 mm; dorsally 3.0 mm. Cocoon slipper shaped, closed basally,
with reinforced edge slightly projected laterodorsally; weave loose and coarse anteriorly. Gill
well sclerotized, cylindrical, comprised of 9-10 terminal branches; 3 primary gill branches:
dorsal one short, oriented posteriorly, and curved with rounded apex; median branch short,
with 3 secondary branches of approximately equal length; dorsal secondary branch bifurcated
basally, median secondary branch subdivided apically into 3 branches, third secondary branch
also with 3 subdivided branches, although on different plane; primary ventral branch curved
and longer than other three main branches (Figs. 124 J, K). All branches apically acuminated.
Frontoclypeus smooth, and with robust spiniform setae, frontal setae separated and superior
very long; thorax with abundant, small, rounded tubercles.
Larva: (Diagnosis based on VARGAS & DÍAZ NÁJERA, 1957 and also DÍAZ NÁJERA &
VULCANO, 1962). Length (mature) 7.5-8.0 mm. Coloration dark greenish. Cephalic
apotome homogeneously pigmented, without special ornamentation. Cephalic fan with
50-55 primary rays. Hypostomium with anterior margin curved and 15-17 lateral setae per
side. Postgenal cleft dome shaped, with apical incision. Anal sclerite, accessory plates and
ventrolateral papillae similar to those of S. gorirossiae, but with ventrolateral plates having
shorter accuminate projections and directed only anteriorly. Anal ring with about 310
rows of hooks. Rectal papillae with 18-36 diverticula.
Distribution: Mexico: Oaxaca.
Bionomics: Unknown.
Discussion: The larva is very close to those of S. gorirossiae and S. temascalense, based
on a similar anal sclerite and accessory plate and the presence of ventrolateral papillae on
the larva. However, the peculiar form and arrangement of the pupal gill branches permits
their separation.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 595

Simulium (Hearlea) paracarolinae COSCARÓN (Figs. 126 I-L; 127 B, C.)


Simulium (Hearlea) paracarolinae COSCARÓN et al., 2004: 32-33.
Larva: Length (mature) 7.0-8.5 mm. Coloration light brown yellowish (in alcohol).
Cephalic apotome light brown, with darkened base, and darkened medial longitudinal stripe on
basal 1/3 and 1+1 small submedian spots near midline of sclerite. Cephalic fan with 44-48
primary rays. Medial antennal article with 4 annulations (Fig. 126 K); ratio of proximal, medial
and distal antennal articles = 1:1.6:1.0. Postgenal cleft subtriangular, without anterior incision
(Fig. 126 I); ratio of hypostomium/hypostomial bridge = 1.2. Hypostomium (Fig. 126 J) with
median tooth longer than others; lateral setae 11-12 and 2 discal setae. Mandible with internal
teeth arrayed in 3-4 rows with 6-8 teeth each; 2 marginal teeth, the second very small (Fig. 126
L); mandibular lateral process unique, elongated, stout and curved. Lateral sclerite of thoracic
proleg with 42-46 teeth. Anal sclerite with ventral branches heavily sclerotized, relatively short,
not encircling posterior circlet, although enclosing approximately 1/3 of it; accessory plates
constituted of 1+1 well-sclerotized, dorsolateral conical plates covered with small setae (Figs.
127 B, C [ds]). No ventrolateral papillae. Anal ring with 398-400 rows of 42-48 hooks each.
Anal papillae with 18-21 diverticula, 6-7 per lobe, median lobe largest.
Distribution: Guatemala.
Bionomics: Unknown.
Discussion: Larval morphology permits differentiation of S. paracarolinae from all
other species. The presence of accessory plates on the anal sclerite and a curved anterior
hypostomial margin place this species into the CAROLINAE species group, for which the
larva is known for all species. The larva cannot be distinguished from the unknown larva of
other S. (Hearlea) species, as it could potentially be that of S. nigricorne, because pupal
morphology suggests it corresponds to another group with different characters in the larva,
such as absence of accessory plates on the distal portion of the abdomen, straight anterior
edge of the hypostomium and distinct mandibular morphology and postgenal cleft. The
closest species is S. carolinae, but it has the anal sclerite ventral branch forming a ring
encircling the rows of hooks of the anal ring, and also has 2+2 dorsolateral accessory plates.
The absence of a mature gill histoblast does not permit comparisons with other species.

Simulium (Hearlea) temascalense DÍAZ NÁJERA & VULCANO (Fig. 127 J.)
Simulium (Hearlea) temascalense DÍAZ NÁJERA & VULCANO, 1962: 102-105, 140, 142; COSCARÓN et al.,
2004: 33.
(Diagnosis of female and pupa based on DÍAZ NÁJERA & VULCANO, 1962).
Female: General coloration blackish. Scutum brown with 1+1 thin silvery vittae not
joining anterior 1+1 subtriangular silvery spots. Pleurae black pollinose. Femora and tibiae
yellowish, with black apices. Meso- and metathoracic coxae, trochanter, tarsomeres I-II,
and apex of tarsomere III black. Cibarium with smooth, thickened, medially concave
anterior margin. Genitalia similar to those of S. larvispinosum.
Male: Unknown.
Pupa: Cocoon (at base) 3.0 mm. Frontoclypeus and anterodorsum of thorax without
tubercles. Gill with general facies of S. carolinae, with anterior branch elongate, relatively
596 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

d 0.2 mm

ds

A v
0.05 mm
B

dd

d
v

ds
df
C
0.5 mm

H
df
J dd
vl

vp
I
dd
0.2 mm

vl

D
F df
dd 0.05 mm

E
gd
G 1 mm

Figure 127. A-J: Larva. A-D, H-J: Terminal portion of abdomen showing anal sclerite and
accessory plates (d = dorsal branch of anal sclerite, dd = dorsolateral conical accessory plates
double, d f = dorsolateral flattened accessory plate, d s = dorsolateral conical accessory plate single,
gd = diverticles of rectal papillae, v = ventral branch of anal sclerite, vl = ventrolateral accessory
plate, vp = ventrolateral papillae). A: S. (Hearlea) capricorne. A: anal sclerite. B-C: S. (H.) paracarolinae;
B: anal sclerite area in transparent slide view; C: abdomen posterodorsal view. D-F: S. (H.)
gorirossiae; D: abdomen posteroventral view; E: abdomen posterodorsal view, with rectal papillae
diverticles (gd); F: eighth segment ventrolateral papilla (vp) sclerotized. G-I: S. (H.) larvispinosum,
general aspect; G: lateral view; H: abdomen posterodorsal view; I: same in transparent slide view. J:
S. (H.) temascalense, abdomen posterolateral view.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 597

thin, with acuminate, spined apex, with 7 small subbasal branches and another curved and
directed posteroventrally; apex of branches spined.
Larva: Length (mature) 8.0 mm. Coloration dark greenish; head yellowish, cephalic
apotome without noticeable ornamentation. Cephalic fan with 50 primary rays. Mandible
with strong preapical tooth, 2 rows of internal teeth, and 2 large marginal teeth;
lateromandibular process long and curved. Ratio of proximal, medial and distal antennal
articles = 1:1.6:1.2. Comb of thoracic proleg sclerite with 38-39 teeth arranged in 10
groups. Hypostoma with anterior edge curved, median tooth longer than other teeth, and
15 lateral setae per side; ratio of hypostomium/hypostomial bridge = 0.8; postgenal cleft
mildly curved apically, without acuminate anterior incision. Segment VIII with 1+1
sublateral, sclerotized papillae. Anal ring with 242 rows of 26-30 hooks. Distal portion of
abdomen with 2+2 dorsolateral acuminate posteriorly directed processes, 1+1 wide
ventrolateral positioned processes having small lateral projections (Fig. 127 J [dd and vl]);
1+1 dorsolateral flattened plates and 1+1 ventrolateral papillae; ventral arms of anal
sclerite surrounding anal ring. Rectal papillae with 21-27 diverticula, 7-9 per lobe.
Distribution: Mexico: Michoacan, Oaxaca.
Bionomics: Unknown.
Discussion: Simulium johnsoni and S. larvispinosum have very similar pupal gills, but the
larvae lack ventrolateral papillae on segment VIII and the females have a blackish scutum
in these species.

Simulium (Trichodagmia) ENDERLEIN (Figs. 128-131; 153.)


Trichodagmia ENDERLEIN, 1934b: 288 (as genus).
Type-species: Simulium townsendi MALLOCH, 1912 = Trichodagmia latitarse ENDERLEIN, 1934b: 288.
Grenieriella VARGAS & DÍAZ NÁJERA, 1951: 141. Type-species: Eusimulium lahillei PATERSON &
SHANNON, 1927: 740.
Female: Big species (wing length 2.8-4.4 mm). Scutum brown-orange, dark brown to
blackish, generally ornamented with 1+1 submedian longitudinal grayish stripes joined
posteriorly, forming lyre-like pattern (Fig. 128 A); pilosity generally homogeneous but can be
grouped simulating scales. Abdomen dark brown with blackish, generally with 1+1 grayish
spots on tergite II. Frons relatively wide (Fig. 128 F) and fronto-ocular triangle about as long
as wide (Fig. 128 G). Cibarium well sclerotized with small teeth generally like spiculae on
median depressed area (Figs. 128 D, H). Claw with well developed subbasal tooth. Eighth
sternite well sclerotized with median portion depressed; gonapophysis subovoidal to
subtriangular with blunt apex and small anterior depression (Fig. 128 I). Genital fork with
wide anterior branches and strong, well-sclerotized apodeme (Fig. 128 K).
Male: Color brown-yellowish to black. Gonostylus nearly 1.5 times longer than
gonocoxite, subcylindrical, acuminate distally and about 2 times longer than wide at
base, generally with 1 apical spur. Ventral plate body wide, about 2.0 times wider than
long (Figs. 129 G, E, K, M), with short median process. Endoparamere branches
without hooks (Fig. 129 C).
Pupa: Cocoon shoe shaped, generally with aperture anteriorly elevated protecting gill;
weave thick. Frontoclypeus reinforced basally. Frontoclypeus and exposed portion of thorax
598 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

generally with abundant tubercles; trichomes small, multibranched. Gill with 12-20 branches
generally moderate thick and relatively short; abdomen with terminal spur very reduced.
Larva: Body shape dorsally frequently straight on subterminal portion as in S.
(Hemicnetha). Cephalic apotome dark brown, mostly homogeneous, darkened on basal
margin and scarcely ornamented. Antenna relatively thick, with medial article much longer
than proximal and distal; mandible with several internal teeth rows and thin marginal
teeth. Postgenal cleft deep, with postgenal bridge shorter than hypostomium length;
hypostomium with straight anterior margin. Anal sclerite with abundant hair, but no
scales. Anal ring with 150-360 rows with 23-40 hooks each. Rectal papillae with elongated
diverticula of 10-45 on each lobe.
Distribution: The subgenus has 7 species in the South American temperate west
sub-Andean and eastern mountains (Fig. 153).

KEYS TO SIMULIUM (TRICHODAGMIA) SPECIES

Females
1 Scutum grayish brown, hair arranged in groups; paraproct relatively short (Fig.
128 J), ratio of length/width basally = 1.3; cibarium teeth well developed (Fig. 128
H) .......................................................................................................... Simulium nigrimanum
– Scutum yellowish brown or reddish to blackish; paraproct relatively long (Figs.
128 C, E), ratio of length/width = 1.9-2.0; cibarium teeth reduced, like spicules
(Fig. 128 D) ......................................................................................................................... 2
2(1) Scutum blackish; paraproct well sclerotized; cercus with distal border straight (Fig.
128 E) ..................................................................................................... Simulium townsendi
3 Scutum yellowish brown to grayish brown or blackish; paraproct medially
sclerotized; cercus with distal border curved (Fig. 128 C) ........................................ 3
4(2) Scutum blackish golden to dark brown; wing length 2.8-3.8mm ..............................
................................................................ Simulium wgodzinskyorum, Simulium sumapazense
– Scutum blackish to reddish brown or grayish brown; wing length 3.9-4.4 mm .....
............................................... Simulium huairayucu, Simulium lahillei, Simulium muiscorum

Males
Not included: Simulium townsendi.

1 Gonostylus with about 4 apical spurs (Fig. 129 L); apical process of ventral plate
relatively long (Fig. 129 M) ............................................................. Simulium nigrimanum
– Gonostylus with 1 (or 2) apical spurs (Figs. 129 B, D, F, J); apical process of
ventral plate short (Figs. 129 B, E, G) (except Simulium wygodzinskyorum) ............. 2
2(1) Scutum reddish brown to brownish orange; ratio of length/width of hind
basitarsus = 4.5-5.0 ............................................................................................................ 3
– Scutum black; ratio of length/width of hind basitarsus = 3.5 ................................ 4
3(2) Scutum reddish brown, showing with some illumination 1+1 subtriangular darker
spots reaching lateral margin by thin anterior stripe (Fig. 129 A); ventral plate with
median process short (Fig. 129 B) ......................................................... Simulium lahillei
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 599

– Scutum brownish orange, showing with some illumination 1+1 oblique


subtriangular nacreous spots, reaching from antero-external angle to submedian
darker stripes (Fig. 129 H); ventral plate with median process long (Fig. 129 K)
...................................................................................................... Simulium wygodzinskyorum
4(2) Ratio of length/width of hind basitarsus = 3.5 ...........................................................
......................................................................... Simulium huairayacu, Simulium sumapazense
– Ratio of length/width of hind basitarsus = 2.5-2.7 ................... Simulium muiscorum

Pupae
Not included: Simulium townsendi.

1 Gill with 18-20 branches (Fig. 130 L); frontoclypeus and thorax without tubercles
............................................................................................................. Simulium nigrimanum
– Gill with 12 branches; frontoclypeus and thorax generally with numerous
tubercles (Fig. 130 F) ......................................................................................................... 2
2(1) Cocoon with anterior aperture border lower than posterior edge (Figs. 130 A, D)
................................................................................................................................................ 3
– Cocoon with anterior aperture border higher than posterior edge (Figs. 130 B, C)
................................................................................................................................................ 4
3(2) Gill branches very thick, emerging from base without trunk, branches apically acute
and with microscopic annulations only distally (Figs. 130 A, H) ......... Simulium lahillei
– Gill branches relatively thin, emerging from base with basal trunk, branches
apically blunt and with microscopic annulations from base (Fig. 130 J) .................
...................................................................................................... Simulium wygodzinskyorum
4(2) Frontoclypeus and exposed portion of thorax without tubercles (Fig. 130 G); gill
expanded (Fig. 130 K) .................................................................... Simulium sumapazense
– Frontoclypeus and exposed portion of thorax with numerous tubercles (Fig. 130
F); gill not expanded .......................................................................................................... 5
5(4) Ventral primary branches positioned far from other branches (Fig. 130 I) ............
................................................................................................................ Simulium huairayucu
– Ventral primary branches positioned near other branches ........ Simulium muiscorum

Larvae
No key to larvae is presented because of scarcity of material and characters.

CLAVES DE ESPECIES DE SIMULIUM (TRICHODAGMIA)


Hembras
1 Escudo castaño grisáceo, pelos dispuestos en grupos; paraprocto relativamente
corto (Fig. 128 J); relación largo/ancho en la base = 1,3; dientes del cibario bien
desarrollados (Fig. 128 H) ............................................................... Simulium nigrimanum
– Escudo castaño amarillento o rojizo a negruzco; paraprocto relativamente largo
(Figs. 128 C, E); relación largo/ancho = 1,9-2,0; dientes del cibario reducidos
como espículas (Fig. 128 D) ............................................................................................ 2
600 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

2(1) Escudo negruzco, paraprocto bien esclerotizado; borde distal del cerco recto (Fig.
128 E) ..................................................................................................... Simulium townsendi
– Escudo castaño amarillento a castaño grisáceo o negruzco, paraprocto
medianamente esclerotizado; borde distal del cerco curvado (Fig. 128 C) ........... 3
3(2) Escudo negruzco dorado a castaño oscuro; largo ala 2,8-3,8 mm ............................
............................................................... Simulium wygodzinskyorum, Simulium sumapazense
– Escudo negruzco a castaño rojizo o castaño grisáceo; largo ala 3,9-4,4 mm .........
............................................... Simulium huairayacu, Simulium lahillei, Simulium muiscorum

Machos
No incluida: Simulium townsendi.

1 Gonostilo con alrededor de 4 espolones apicales (Fig. 129 L); prolongación apical
de la placa ventral relativamente larga (Fig. 129 M) .................. Simulium nigrimanum
– Gonostilo con 1 (ó 2) espolones apicales (Figs. 129 B, D, F, J); prolongación apical de
la placa ventral corta (Figs. 129 B, E, G) (excepto Simulium wygodzinskyorum) ............. 2
2(1) Escudo castaño rojizo a castaño naranja; relación largo/ancho del basitarso
posterior = 4,5-5,0 ............................................................................................................. 3
– Escudo negro; relación largo/ancho del basitarso posterior = 3,5 ........................ 4
3(2) Escudo castaño rojizo mostrando con cierta iluminación 1+1 manchas oscuras
subtriangulares, llegando al borde lateral por una fina banda anterior (Fig. 129 A);
prolongación mediana de la placa ventral corta (Fig. 129 B) .......... Simulium lahillei
– Escudo castaño naranja mostrando con cierta iluminación 1+1 manchas
nacaradas subtriangulares oblicuas, llegando desde el ángulo antero externo hasta
las bandas submedianas más oscuras (Fig. 129 H); prolongación mediana de la
placa ventral larga (Fig. 129 K) ............................................. Simulium wygodzinskyorum
4(2) Relación largo/ancho del basitarso posterior = 3,5 .....................................................
......................................................................... Simulium huairayacu, Simulium sumapazense
– Relación largo/ancho del basitarso posterior = 2,5-2,7 ............. Simulium muiscorum

Pupas
No incluida: Simulium townsendi.

1 Branquia con 18-20 ramas (Fig. 130 L); frontoclípeo y tórax sin tubérculos .........
............................................................................................................... Simulium nigrimanum
– Branquia con 12 ramas; frontoclípeo y tórax generalmente con numerosos
tubérculos (Fig. 130 F) ...................................................................................................... 2
2(1) Capullo con borde frontal de la abertura más bajo que borde posterior (Figs. 130
A, D) ..................................................................................................................................... 3
Capullo con borde frontal de la abertura más alto que el borde posterior (Figs. 130
B, C) ...................................................................................................................................... 4
3(2) Ramas de las branquias muy gruesas, emergiendo desde la base sin tronco, ápice
de las ramas aguzado y con anillaciones microscópicas solo distalmente (Figs. 130
A, H) ............................................................................................................ Simulium lahillei
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 601

– Ramas de las branquias relativamente finas, emergiendo desde la base con un


tronco basal; ápice de las ramas romo y con anillaciones microscópicas desde la
base (Figs. 130 J) ...................................................................... Simulium wygodzinskyorum
4(2) Frontoclípeo y porción expuesta del tórax sin tubérculos (Fig. 130 G); branquia
expandida (Fig. 130 K) ................................................................... Simulium sumapazense
– Frontoclípeo y porción expuesta del tórax con numerosos tubérculos (Fig. 130 F);
branquia no expandida ...................................................................................................... 5
5(2) Rama primaria ventral de la branquia alejada de las otras ramas (Fig. 130 I) .........
................................................................................................................ Simulium huairayacu
– Rama primaria ventral dispuesta junta con las otras ramas ....... Simulium muiscorum

Larvas
No se presenta clave por escaso material y caracteres.

Simulium (Trichodagmia) townsendi MALLOCH (Fig. 128 E.)


Simulium townsendi MALLOCH, 1912: 651-652.
Simulium (Trichodagmia) townsendi: HERNÁNDEZ & SHELLEY, 2005: 12-15.
Simulium chalcocoma KNAB, 1914b: 85.
Simulium (Trichodagmia) chalcocoma: VARGAS & DÍAZ NÁJERA, 1953b: 139; MIRANDA ESQUIVEL &
COSCARÓN, 2001: 429; COSCARÓN ARIAS, 2003: 191.
Simulium (Grenieriella) chalcocoma: VARGAS & DÍAZ NÁJERA, 1951: 142; COSCARÓN, 1987: 38.
Simulium (Hemicnetha) townsendi: CROSSKEY & HOWARD, 1997: 38; 2004: 74
Trichodagmia ecuadoriense ENDERLEIN, 1934a: 193.
Trichodagmia latitarse ENDERLEIN, 1934b: 289.
Trtichodagmia strigata ENDERLEIN, 1934b: 290.
Female: Wing length 3.5-3.7 mm. Scutum brown and grayish with 1 thin median and
1+1 wider submedian and lateral stripes; submedian stripes joined posteriorly, forming
lyre pattern as in S. rubrithorax; with frontal illumination, vittae becoming blackish, and
with posterior light, median and sublateral longitudinal areas grayish and black; pilosity
recumbent, uniformly golden to silvery. Scutellum and metanotum brown and grayish.
Legs blackish with distal portion of hind tibia and basitarsus with silver pruinosity.
Abdomen blackish with 1+1 light grayish large lateral and posterior spots on tergite II,
thinner on tergites III-V; tergal plates II-V dark brown. Cibarium as in S. huairayacu;
fronto-ocular triangle slightly wider than high. Sensory vesicle of maxillary palp near 1/3
of palpomere length. Sc with hair all along vein. Claw with well-developed subbasal tooth.
Eighth sternite and gonapophysis as in S. nigrimanum, but the first more sclerotized and
with about 17 setae on each side; cerci and paraprocts well sclerotized; cerci with straight
border (Fig. 128 E); paraprocts about 2 times longer than wide; genital fork with anterior
apodemes relatively long and well sclerotized.
Male, pupa and larva: Unknown.
Discussion: The species is very close to S. huairayacu, S. muiscorum, S. sumapazense and
S. wygodzinskyorum; small differences permit them to be separated, such as the curved distal
border of the cercus, and the paraproct slightly sclerotized medially.
602 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

J
D

K
I

Figure 128. Female. A-C: S. (Trichodagmia) lahillei; A: scutum, B: genitalia, ventral view showing
eighth sternite and gonapophysis, cercus and paraproct, genital fork and spermatheca; C: cercus
and paraproct, slide view. D: S. (T.) huairayacu, cibarium median portion. E: S. (T.) townsendi, cercus
and paraproct. F-K: S. (T.) nigrimanum; F: frons; G: fronto-ocular triangle; H: cibarium; I: eighth
sternite and gonapophysis; J: cercus and paraproct; K: genital fork.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 603

We accept the synonyms proposed by HERNÁNDEZ & SHELLEY, 2005, except S.


muiscorum which shows differences such as the large size and short paraproct. The
similarity among females is the principal element used to synonymize this species, but
without knowledge of the other stages, it is doubtful if they are conspecific. Some species,
such as S. muiscorum, S. huairayacu and S. sumapazense, have very similar females, but it is
possible to separate them in the other stages, especially the immatures. Knowledge of
more topotypical material, including all life stages, and cytological and molecular methods,
will help to decide if S. muiscorum is a synonym of S. townsendi.

Simulium (Trichodagmia) lahillei (PATERSON & SHANNON)


(Figs. 128 A-C; 129 A-C; 130 A, H, N; 131 A-F.)
Eusimulium lahillei PATERSON & SHANNON, 1927: 740; PINTO, 1931: 678.
Simulium lahillei: VARGAS, 1945a: 153; WYGODZINSKY, 1949 : 306; 1953a: 317; STONE, 1963: 8.
Simulium (Trichodagmia) lahillei: CROSSKEY & HOWARD, 1997: 79; 2004: 74; MIRANDA ESQUIVEL &
COSCARÓN, 2001: 429, 435.
Simulium (Grenieriella) lahillei: VARGAS & DÍAZ NÁJERA, 1951: 141; COSCARÓN, 1987: 38; 1991: 276-278.
Female: Wing length 4.0-4.3 mm. Scutum brownish red with 1+1 submedian thin
light gray pollinose stripe converging to posterior yellowish area, forming lyre pattern,
with changing light position, thin stripes become blackish (Fig. 128 A). Scutum pile
uniformly yellowish to silvery. Scutellum and metanotum dark brown, the latter with
grayish pruinosity. Legs yellowish with silvery pilosity. Abdomen yellowish, tergite I darker
with yellow posterior margin; II-V matte with 1 median and 1+1 submedian darker spots,
and tergite II with 1+1 silvery anterior spots; VI-IX shinny. Gonapophysis subtriangular,
about as long as wide at base (Fig. 125 B). Paraproct longer than wide at base, curved
distally with slight anterior concavity (Fig. 128 C).
Male: Wing length 3.6-4.0 mm. Scutum brownish red with 1+1 small light grayish
anterior submedian subtriangular spots (Fig. 129 A); with changing light position, stripes
become darker and reach lateral border by thin anterior stripe; pilosity uniformly grayish.
Scutellum yellowish red; metanotum reddish to blackish; abdomen as in female. Ratio of
length/width of hind basitarsus = 4.5-5.0. Gonostylus nearly half as long as gonocoxite,
with curved apex and 1-2 apical spurs (Fig. 129 B); ventral plate with short median, relatively
acute process (Fig. 129 B); endoparameres and aedeagal membrane as in figure 129 C.
Pupa: Cocoon with aperture frontally projected, protecting gill branches (Fig. 130
A). Length basally 3.0-3.3 mm; dorsally 3.0-3.8 mm; maximum length 5.5-6.0 mm; gills
1.7-1.8 mm. Gills with 12 branches very thick, with acute apices; 4 primary branches
emerging from base (Fig. 130 H). Frontoclypeus and exposed portion of thorax with
small and abundant tubercles; trichomes with 5-8 branches and spiraled basal trunk as in
S. huairayacu. Abdominal chaetotaxy as in figure 130 N with very reduced terminal spur.
Larva: Maximum length 13.0 mm. Body general as in figure 131 A. Cephalic
apotome dark brown, mostly uniform, darker on basal border and with negative spots
(Figs. 131 B, C). Cephalic fan with 50-60 rays. Antenna thick (Fig. 131 D), ratio of
proximal, medial and distal articles = 1:1.8:1.1. Mandible with much reduced second
marginal tooth (Fig. 131 E). Postgenal cleft subtriangular (Fig. 131 F). Anal sclerite with
604 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

B L

A H

D
E

M C

Figure 129. Male. A-C: S. (Trichodagmia) lahillei; A: scutum, B: genitalia, ventral view, gonocoxite and
gonostylus, ventral plate; C: endoparameres and aedeagus. D-E: S. (T.) huairayacu; D: gonocoxite
and gonostylus; E: ventral plate. F-G: S. (T.) muiscorum; F: gonocoxite and gonostylus, dorsal view;
G: ventral plate. H-K: S. (T.) wygodzinskyorum; H: scutum; I: hind basitarsus; J: gonostylus; K: ventral
plate. L-M: S. (T.) nigrimanum; L: gonocoxite and gonostylus, dorsal view; M: ventral plate.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 605

hair and scales, the last in 2-4 rows. Anal ring with 300-360 rows with 30-40 hooks each.
Rectal papillae with about 10 diverticula on each lobe.
Distribution: Argentina: Catamarca, Córdoba, Jujuy, Salta, Tucumán. Bolivia;
Peru: Cuzco.
Bionomics: This species breeds in creeks of median discharge, fast current and
clear water, attached in groups on stones near the surface, frequently in winter and at the
beginning of spring. Females were collected on horses and biting humans.
Discussion: The closest species is S. huairayacu; differences are explained under that
species. Probably S. lahillei is the most important vector of Mansonella ozzardi in northern
Argentina (SHELLEY & COSCARÓN, 2001).

Simulium (Trichodagmia) wygodzinskyorum COSCARÓN & PY-DANIEL


(Figs. 129 H-K; 130 F, J.)
Simulium (Grenieriella) wygodzinskyorum COSCARÓN & PY-DANIEL, 1989: 319.
Simulium (Trichodagmia) wygodzinskyorum CROSSKEY & HOWARD, 1997: 79; 2004: 74; MIRANDA
ESQUIVEL & COSCARÓN, 2001: 429-435; COSCARÓN ARIAS, 2003: 191, 206.
Female: Wing length 3.5 mm. General coloration brownish. Scutum dark reddish
brown with 1+1 submedian stripes, with lyre pattern in some illumination; pilosity
uniformly silvery to yellowish, in accordance with light incidence. Scutellum and
metanotum dark brown, the latter with gray pruinosity. Legs yellowish with silvery pilosity.
Abdomen brownish yellow with darker tergal plates. Cibarium smooth and sclerotized
subbasally. Claw with short subbasal tooth. Genitalia as in S. lahillei; eighth sternite with
about 10 setae on each side; gonapophysis apically curved.
Male: Wing length 3.0-3.2 mm. Scutum yellowish brown with 1 median longitudinal
and 1+1 thin submedian darker stripes converging posteriorly; with other illumination,
1+1 anterior oblique nacreous spots can be seen from outer anterior angle to submedian
darker stripes (Fig. 129 H). Scutellum, metanotum and abdomen brown, changing to dark
brown on the latter and with 1+1 lateral silvery spots on tergites II and IV. Legs as in
female but darker; ratio of length/width of hind basitarsus = 4.8 (Fig. 129 I). Gonostylus
about 2 times longer than gonocoxite, relatively acuminate apically (Fig. 129 J). Ventral
plate with well-developed median process (Fig. 129 K).
Pupa: Cocoon relatively low anteriorly, similar to that of S. sumapazense but lower
than anterior portion of aperture; length basally 2.5-3.0 mm; dorsally 2.7-3.3 mm;
maximum length 3.4-4.8 mm, gills 1.0-1.2mm. Frontoclypeus and thorax with numerous
small and granulose tubercles (Fig. 130 F); basal portion of frontoclypeus reinforced.
Cephalic and thoracic trichomes multibranched, frontal trichomes reduced. Gill with 12
medially thick branches, with microscopic annulations on surface and acute apically; 4
short primary branches emerging from short basal trunk (Fig. 130 J).
Larva: Cephalic apotome dark brownish without much ornamentation and with
negative spots. Cephalic fan with 30-32 rays. Ratio of proximal, medial and distal antennal
articles = 1:2.5-2.8:1.2-1.5. Anal ring with 188-212 rows with 26-28 hooks each. Rectal
papillae with 17-19 diverticula on each lobe.
Distribution: Peru; Ecuador: Pichincha; Junín.
606 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

A
C B

E D

I
F J

K
L

Figure 130. Pupa. A-E: lateral view; A: S. (Trichodagmia) lahillei; B: S. (T.) huairayacu; C: S. (T.)
muiscorum; D: S. (T.) sumapazense; E: S. (T.) nigrimanum. F-G: frontoclypeus; F: S. (T.) wygodzinskyorum;
G: S. (T.) sumapazense. H-L: gills; H: S. (T.) lahillei; I: S. (T.) huarayacu; J: S. (T.) wygodzinskyorum; K: S.
(T.) sumapazense; L: S. (T.) nigrimanum. M: S. (T.) huairayacu, thoracic trichome enlarged. N: S. (T.)
lahillei, abdominal chaetotaxy.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 607

Bionomics: Unknown.
Discussion: Simulium lahillei and S. huairayacu are very close, but they are bigger, have
different male scutal ornamentation, darker color, especially in S. huairayacu, a ventral plate
with a short median process, and a pupal gill without a basal trunk and with thicker
branches in S. lahillei and a different arrangement in S. huairayacu. Simulium townsendi also is
similar but can be differentiated by the wider, more sclerotized paraproct, distally
flattened cerci, and general blackish coloration.

Simulium (Trichodagmia) huairayacu WYGODZINSKY (Figs. 128 D; 129 D, E; 130 B, I, M.)


Simulium huairayacu WYGODZINSKY, 1953a: 310; 1961: 460.
Simulium (Grenieriella) huairayacu: COSCARÓN, 1987: 38; 1991: 278.
Simulium (Trichodagmia) huairayacu CROSSKEY & HOWARD, 1997: 79; 2004: 74; MIRANDA ESQUIVEL &
COSCARÓN, 2001: 429, 435.
Female: Wing length 3.9-4.4 mm. Scutum brown-violaceous or reddish brown to
brownish orange, with 1+1 submedian grayish stripes anteriorly starting as a subtriangle
and converging posteriorly, forming lyre pattern. Scutellum yellowish brown and metano-
tum dark brown with gray pruinosity. Legs yellowish gray. Abdomen light yellowish gray
with blackish velvety spots on tergal plates of segments II-V, and slightly lighter on VI-VII
with dark grayish center; segments II-VI with 1+1 sublateral silvery spots, more
developed on II. Cibarium concave medially and with abundant spiniform teeth (Fig. 128
D). Fronto-ocular triangle shorter than wide; claw with subbasal tooth moderately
developed. Genital as in S. lahillei.
Male: Wing length 3.5-4.0 mm. Scutum velvety black, showing with some illumina-
tion silvery lateral and posterior edges. Scutellum dark brown; metanotum blackish; some
specimens lighter with reddish brown scutum. Legs brown to blackish. Abdomen velvety
black; tergites II, V, VII with 1+1 submedian silvery spots, those of V smaller and in some
specimens imperceptible. Ratio of length/width of hind basitarsus = 3.5. Genitalia very
similar to those of S. lahillei, but gonostylus shorter, about 1.5 times longer than
gonocoxite (Fig. 129 D). Ventral plate with short median process (Fig. 129 E).
Pupa: Cocoon with aperture anteriorly elevated (Fig. 130 B), protecting gills; length
basally 4.0-5.0 mm, on dorsum 2.6-3.0mm; maximum length 4.5-5.1mm; gills 1.4-1.6 mm.
Frontoclypeus and exposed portion of thorax with abundant tubercles; trichomes
frequently curled with thick basal trunk and 7-12 thin and elongated branches (Fig. 130
M). Gill with 12 thick branches with blunt apices, 4 relatively short primary branches
emerging from common place, with 2 ventral branches directed down and forward and
other directed dorsally, forming a bunch (Fig. 130 I).
Larva: Maximum length 10-11mm. Very similar to S. lahillei. Ratio of proximal,
medial and distal antennal articles = 1:2.3-2.7:1.3. Anal ring with about 250 rows with 30
hooks each. Rectal papillae with about 45 diverticula on each lobe.
Distribution: Argentina: Jujuy, Salta, Tucumán; Bolivia: Cochabamba, Potosí.
Bionomics: This species breeds in small to medium-sized torrential creeks in clear,
cold water and is found frequently with S. lahillei.
608 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Discussion: This species is very close to S. lahillei but is easy to separate by the pupal
gill shape. In the adults, it is possible to separate the male because S. lahillei is lighter
reddish brown.

Simulium (Trichodagmia) muiscorum BUENO, MONCADA & MUÑOZ DE HOYOS


(Figs. 129 F, G; 130 C.)
Simulium (Hemicnetha) muiscorum BUENO et al., 1979: 581-594.
Simulium (Trichodagmia) muiscorum: CROSSKEY & HOWARD, 1997: 79; 2004: 74; MUÑOZ DE HOYOS &
COSCARÓN, 1999: 183; MIRANDA ESQUIVEL & COSCARÓN, 2001: 429, 435.
Simulium (Grenieriella) muiscorum: COSCARÓN, 1987: 37.
Female: Wing length 3.9-4.1 mm. Coloration brown dark blackish. Scutum blackish
with small golden to grayish hair and some darkened; with frontal illumination, 1+1
subtriangular light gray spots continued posteriorly by thin dark stripes. Scutellum and
metanotum blackish, the latter shiny. Legs brown with apex of femora, tibiae and
tarsomeres blackish. Cibarium and genitalia as in S. huairayacu, claw with well-developed
subbasal tooth. Abdomen dark brown with 1+1 grayish spots laterally on tergites III-IV.
Gonapophysis, genital fork and cercus as in S. lahillei, but the former more rounded
apically; paraproctos as in S. townsendi but shorter and less sclerotized
Male: Wing length 3.5-3.8 mm. Scutum blackish with golden to gray pilosity;
scutellum, metanotum and legs as in female. Hind legs wide; ratio of length/width of
hind basitarsus = 2.5-2.7. Abdomen dark brown with tergal plates darkened and 1+1
big silvery spots on tergite II. Gonostylus and ventral plate as in figures 129 F, G,
respectively.
Pupa: Cocoon with aperture very elevated frontally, covering mostly of gill base (Fig.
130 C). Length basally 3.5-4.0 mm, maximum length 4.8-5.0 mm; gills 1.2 mm.
Frontoclypeus and exposed portion of thorax with numerous smooth tubercles. Facial
trichomes with 7-12 branches, frontal with 2-5, thoracic with 3-5 branches. Gill with 12
branches emerging from short basal trunk with 4 primary branches; gill filaments with
microscopic pseudoannulations and acute apex. Abdomen terminal spur absent.
Larva: Maximum length 8.5-9.0 mm. Cephalic apotome brownish, homogeneously
pigmented, only darkened on narrow basal margin and without definite spots. Cephalic
fan with 36-39 rays. Antenna surpassing apex of cephalic fan stem; ratio of proximal,
medial and distal antennal articles = 1:1.9-2:1.1-1.4. Mandible with thin marginal teeth, the
second very small. Postgenal cleft dome shaped, postgenal bridge nearly half of
hypostomium length. Thoracic proleg sclerite with about 47 teeth arranged in 15-17
groups. Anal sclerite without scales. Anal ring with about 196 rows of 30-34 hooks each.
Rectal papillae with 24-35 diverticula on each lobe.
Distribution: Colombia: Cundinamarca.
Bionomics: This species breeds in torrential creeks with rocky bottoms and clear,
cold water, attached to rocks or aquatic vegetation. Females bite cattle.
Discussion: Adult morphology is similar to that of the other species in the
subgenus. The closest species is S. huairayacu, with similar male blackish coloration. The
adults of S. lahillei and S. wygodzinskyorum are lighter reddish brown. The anterior elevated
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 609

aperture of the cocoon is very similar to that of S. huairayacu, but the gill branching, with
more open basal primary branches, plus the larger larval size, greater number of anal ring
rows and rectal papillae diverticula allow separation. It would be useful to do cytological
and molecular studies to reaffirm the validity of these species.

Simulium (Trichodagmia) sumapazense COSCARÓN & PY-DANIEL (Figs. 130 D, G, K.)


Simulium (Grenieriella) sumapazense COSCARÓN & PY-DANIEL, 1989: 321.
Simulium (Trichodagmia) sumapazense: CROSSKEY & HOWARD, 1997: 79; 2004: 74; MUÑOZ DE HOYOS &
COSCARÓN, 1999: 183.
Female: Wing length 3.2-3.7 mm. Scutum blackish golden with 1+1 grayish
submedian stripes forming lyre pattern, pilosity homogeneous. Paraprocts about as long
as wide at base.
Male: Wing length 2.8-3.8 mm. Scutum blackish. Ratio length/width of hind basitarsus = 3.5.
Pupa: Cocoon anterior aperture border higher than posterior border (Fig. 130 D).
Length basally 3.7 mm, on dorsum 3.3 mm, maximum length 5.5 mm; gills 0.9-1.2 mm.
Frontoclypeus and exposed portion of thorax without tubercles except facial area with
small tubercles (Fig. 130 G) and posterior area of thorax with numerous small acute
tubercles; cephalic trichomes with 3-7 and thoracic with 8-9 branches. Frontoclypeus
basal border reinforced. Gill with 12 branches arranged in 1 plane, emerging from short
basal trunk, with acute apex (Fig. 130 K).
Larva: Length over 7.0 mm. Antenna longer than cephalic fan stem. Anal ring with
about 150 rows of hooks. Rectal papillae with 20-26 diverticula on each lobe.
Distribution: Colombia: Cundinamarca.
Bionomics: Unknown.
Discussion: This species is near S. muiscorum but it can be separated by the
numerous tubercles on the frontoclypeus and exposed portion of the pupal thorax, higher
frontal aperture on the cocoon, and gill branches arranged very closely in a bunch.

Simulium (Trichodagmia) nigrimanum MACQUART


(Figs. 128 F-K; 129 L, M; 130 E, L; 131 H-M.)
Simulium nigrimanum MACQUART, 1838: 88; SHELLEY et al., 1984: 146-147.
Simulium (Trichodagmia) nigrimanum: CROSSKEY & HOWARD, 1997: 79; 2004: 74, MIRANDA ESQUIVEL &
COSCARÓN, 2001: 429, 435; HERNÁNDEZ & SHELLEY, 2005: 11-12.
Simulium (Grenieriella) nigrimanum: COSCARÓN, 1987: 37.
Simulium pruinosum LUTZ, 1910: 250-251, 263, 265-266; PINTO, 1931: 704; PY-DANIEL, 1989a: 256.
Simulium (Grenieriella) pruinosum: COSCARÓN, 1991: 280-281.
Grenieriell pruinosa: PY-DANIEL & MOREIRA SAMPAIO, 1995: 118.
Eusimulium pruinosum: LANE & PORTO, 1939:169, 174-175; LANE & VULCANO, 1943: 436.
Simulium spadicidorsum ENDERLEIN, 1934a: 194.
Female: Wing length 2.9-3.6 mm. General coloration grayish brown. Scutum and
scutellum light brown with soft gray pollinosity; scutum pilosity tin to silvery, arranged in
small groups. Metanotum velvety grayish brown; pleurae brown with gray pollinosity. Legs
610 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

A
K

E D H

Figure 131. Larva. A-G: S. (Trichodagmia) lahillei; A: larva, lateral view; B: cephalic apotome; C: basal
portion of head dorsally enlarged, D: antenna; E: mandibular teeth; F: postgenal cleft and
hypostomium; G: hypostomial teeth of anterior margin. H-M: S. (T.) nigrimanum; H: larva, in lateral
view, I: cephalic apotome; J: antenna; K: mandibular teeth, L: postgenal cleft and hypostomium, M:
rectal papillae diverticula.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 611

yellowish brown with distal portion of femora, distal 3/4 of tibia and tarsomeres blackish
brown. Abdomen dark brown; tergites VI-IX cereus, shiny with grayish tones. Frons
convergent (Fig. 128 F). Fronto-ocular triangle wider than high (Fig. 128 G). Basal portion
of cibarium with well-sclerotized border and well-developed teeth on median depressed
area, continued by slight serrations laterally (Fig. 128 H). Sensory vesicle of maxillary palp
about 1/4 of palpomere length. Sc with 12-15 hairs; claw with large subbasal tooth.
Eighth sternite with about 10 setae on each side. Gonapophysis subovoidal with rounded
apex (Fig. 128 I). Paraproct longer than wide, rounded apically and with small anterior
depression (Fig. 128 J). Genital fork with large posterior branches (Fig. 128 K).
Male: Wing length 2.5-2.7 mm. Scutum brown-yellow to dark brown, showing
with some light 1+1 submedian blackish stripes; pile golden. Abdomen dark brown with
1+1 silvery spots on tergite II and large spots on V-VIII. Gonocoxite subquadrate and
nearly as long as gonostylus; thick gonostylus about 1.9 times longer than wide, with
about 4 apical spurs (Fig. 129 L). Ventral plate with well-developed median apical
process (Fig. 129 M).
Pupa: Cocoon with aperture frontally elevated (Fig. 130 E); length at base = 3.0-3.8
mm, at dorsum 2.0-2.5 mm, maximum length 3.5-4.0 mm; gill 1.4-1.5 mm. Cocoon weave
thick but translucent. Frontoclypeus and exposed portion of thorax without tubercles,
small tubercles on facial area. Frontal trichomes single, facial bifid, thoracic short with 1-
4 branches. Gill relatively short with 18-20 filamentous branches, emerging from short
basal trunk and arranged in bunch (Fig. 130 L). Abdomen without terminal spur.
Larva: Maximum length 8.0 mm. Shape as in figure 131 H. Cephalic apotome
slightly darkened on posterior half and especially on basal border; median longitudi-
nal and 1+1 submedian spots; the 1+1 basal spots negative (Fig. 131 I). Antenna
relatively thin (Fig. 131 J), not surpassing apex of cephalic fan stem; ratio of
proximal, medial and distal articles = 1:1.5-1.6:1:1.1. Mandible with several rows of
internal teeth (Fig. 131 K), marginal teeth very thin. Hypostomium with median tooth
slightly beyond level of lateral teeth; postgenal bridge shorter than hypostomium
length (Fig. 131 L). Body integument with scarce and small filiform trichomes. Anal
ring with 146 rows of 23-25 hooks each. Lateral sclerite of proleg with about 36 teeth
arranged in 17-19 groups. Anal sclerite with abundant hair. Rectal papillae with 17-23
diverticula on each lobe (Fig. 131 M).
Distribution: Brazil: Espírito Santo, Goiás, Mato Grosso, Mato Grosso do Sul, Minas
Gerais, Pará, Paraná, São Paulo, Tocantins; Colombia: Vaupes; Paraguay; Venezuela.
Bionomics: This species breeds in small creeks with fast-flowing, clear water 0.20-
0.40 m deep. Females are very noxious to humans.
Discussion: Simulium nigrimanum is easy to separate from similar species of the
subgenus because the male has a gonostylus with one to four apical spurs and a longer
median process on the ventral plate.
SHELLEY et al. (1984: 146), based on a study of two females in the MNHN labeled as
S. nigrimanum (that were designated as syntypes), decided that pruinosum LUTZ is a synonym.
PY-DANIEL (1989: 256) did not agree with this designation because a male is mentioned in
the original description of S. nigrimanum. PY-DANIEL (1989) prefers to maintain S.
nigrimanum as species inquirenda.
612 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Simulium (Thyrsopelma) ENDERLEIN (Figs. 132-135; 154.)


Thyrsopelma ENDERLEIN, 1934b: 284 (as genus).
Type-species: Thyrsopelma brasiliense ENDERLEIN, 1934b: 284 = Simulium orbitale LUTZ, 1910: 231.
Female: Relatively big, wing length 2.0-3.6 mm. General coloration blackish. Scutum
reddish brown with gray pollinosity and hair yellow to silvery, grouped, simulating scales.
Abdomen black. Frons relatively wide. Fronto-ocular triangle about as high as wide to
higher than wide. Basal portion of cibarium smooth and well sclerotized, as for cornuae
and sometimes median area. Sensory vesicle of maxillary palp smaller than half of
palpomere. Basal sector of R haired; Sc with or without hair. Claw without subbasal tooth.
Eighth sternite with abundant hair; gonapophysis subovoidal, generally wider than long.
Male: Coloration similar to that of female, showing in some cases black silvery
lateral margin and 1+1 anterior spots as wide longitudinal stripes; abdomen blackish with
1+1 lateral silvery spots on segments II and smaller on V-VII. Gonostylus 1.5-2 times
longer than gonocoxite, subtriangular, curved on apical third and becoming thinner
distally with 1 apical spur. Ventral plate body wider than high with big median process.
Median sclerite subrectangular with deep median incision. Endoparameres without teeth.
Pupa: Cocoon shoe shaped projected ventrally, with reinforced border of aperture,
protecting base of gill branches. Frontoclypeus and exposed portion of thorax with
rounded or acuminate or spine-like tubercles on base of frontoclypeus and ventrolateral
area of thorax. Cephalic trichomes (2+2 frontal, 1+1 facial) and thoracic (5-6) single and
small. Gill branches 12-50, relatively thick, smooth, with apex acute and well sclerotized.
Larva: Body cuticle with single hairs, generally with racket shape. Cephalic apotome
weakly darkened basally and without special ornamentation. Antenna surpassing apex of
cephalic fan stem, with medial article about 2 times longer proximal or distal articles.
Cephalic fan rays with teeth of similar length and divergent at sides distally, appearing as
2 rows. Mandible with large preapical teeth nearly as long as apical tooth; marginal teeth
large. Postgenal cleft subovoidal, postgenal bridge slightly shorter than hypostomium
length. Hypostomium with median tooth lower than lateral teeth, and intermediate lateral
teeth very wide. Anal sclerite without scales, anal ring with 120-300 rows with 17-35 hooks
each. Rectal papillae with 5-30 diverticula on each lobe.
Simulium (Thyrsopelma) is a Neotropical taxon distributed in tropical South America. It
is composed of 5 species. It shares several characters with Simulium (Trichodagmia)
(MIRANDA ESQUIVEL & COSCARÓN, 2001), and some authors, such as CROSSKEY & HOWARD
(1997, 2004) and SHELLEY et al. (2000), include these species in Trichodagmia. Simulium
(Hemicnetha) also shows similarities with Thyrsopelma and with Trichodagmia, including the
morphology of the gonocoxite, gonostylus and paraproct; pupal cocoon and gill
branches; and proportion of antennal articles and large number of hook rows and hooks
in the anal ring of the larva.
Simulium (Thyrsoplema) also has similarities with Ethiopian Anasolen, Xenosimulium and
Freemaniellum (CROSSKEY, 1969; COSCARÓN, 1991; MIRANDA ESQUIVEL & COSCARÓN, 2001). The
similarities are the smooth cibarium, haired basal sector of R, absence of a subbasal tooth
on the claw, scutum not ornamented; cocoon with anterior projection, absence of spines on
the abdominal terminal tergites of the pupa; and absence of ventral tubercles on the larval
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 613

abdomen, elevated number of hook rows in the anal ring, presence of setae on the larval
cuticle, and short and blunt teeth on the hypostomium, all showing a strong relationship
with the Gondwanian fauna (COSCARÓN, 1991; MIRANDA ESQUIVEL & COSCARÓN, 2004).
In the absence of a world revision, we maintain the subgeneric separation, based
on morphological differences, as also indicated for the other Neotropical taxa that we
have treated.

KEYS TO SIMULIUM (THYRSOPELMA) SPECIES

Females
1 Scutum dark reddish brown; gonapophysis wider than long (Fig. 132 M);
paraproct subtriangular without anterior depression (Fig. 132 N) ............................
..............................................................................................................Simulium scutistriatum
– Scutum blackish; paraproct with anterior deep depression, distally with tail-like
process (Fig. 132 G); gonapophysis about as long as wide (Figs. 132 D, E, H) .......... 2
2(1) Paraproct continued distally, with relatively wide and curved membranous process
(Fig. 132 G) ............................................................................................ Simulium hirtipupa
– Paraproct truncate at short distance from base, with short, thin and straight
membranous process (Fig. 132 D) ................................................................................. 3
3(2) Claw with small subbasal tooth ........................................................ Simulium perplexum
– Claw without subbasal tooth (Fig. 132 C) .................................................................... 4
4(3) Wing length 1.8-2.5 mm; basal sector of R and Sc without hair ..............................
................................................................................................................... Simulium guianense
– Wing length 2.8-3.1 mm; basal sector of R with 20-40 hairs, Sc with 11-15 hairs
..................................................................................... Simulium orbitale, Simulium itaunense

Males
1 Scutum dark brown to blackish, without silvery humeral spots; ventral plate body
without lateral processes and with or without short median process (Fig. 133 K)
................................................................................................................................................ 2
– Scutum blackish, generally with silvery humeral spots that reach posterior silvery
area (Figs. 133 A, G); ventral plate body with lateral processes and long median
process (Figs. 133 D, I) ..................................................................................................... 3
2(1) Ventral plate without median process; gonostylus with apical spur thin and
pointed ................................................................................................... Simulium perplexum
– Ventral plate with short median process (Fig. 133 K); gonostylus with apical spur
thick and blunt apex (Fig. 133 J) ...................................................Simulium scutistriatum
3(1) Ventral plate with median process subpentagonal and wider at base than at apex
(Fig. 130 I); gonostylus relatively thin basally (Fig. 133 H) .......... Simulium hirtipupa
– Ventral plate with median process pear shaped, wider at apex than at base (Fig.
133 D); gonostylus relatively wide basally (Figs. 133 B, C) ....................................... 4
4(3) Scutum with 1+1 silvery spots extended from humeral area to posterior silvery
area as wide submedian stripes (Fig. 133 G); basal sector of R without hair ........
................................................................................................................... Simulium guianense
614 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

– Scutum with 1+1 silvery spots reduced mostly to humeral area, not reaching
posterior area (Fig. 133 A); basal sector of R with hair ............................................ 5
5(4) Wing length 2.5-2.7 mm; scutum with anterior 1+1 silvery spots elongated (Fig.
133 A) …. .................................................................................................. Simulium orbitale
– Wing length 2.8-3.0 mm; scutum with anterior 1+1 silvery spots rounded ...........
................................................................................................................... Simulium itaunense

Pupae
1 Gill with 20-50 branches .................................................................................................. 2
– Gill with 12 branches ........................................................................................................ 3
2(1) Gill with 21-24 branches (Fig. 134 B); cocoon aperture relatively well elevated
anteriorly (Fig. 134 A) ............................................................................. Simulium orbitale
– Gill with 46-52 branches; cocoon aperture weakly elevated anteriorly (similar to
Simulium hitipupa, Fig. 134 F) .............................................................. Simulium itaunense
3(1) Frontoclypeus and exposed portion of thorax covered by spines (Figs. 134 F, I);
gill branches covered by strong spiculae (Fig. 134 H) ................ Simulium hirtipupa
– Frontoclypeus and exposed portion of thorax without spines; gill branches
without spiculae or with very weak spiculae ................................................................ 4
4(3) Frontoclypeus and exposed portion of thorax with numerous apically acute tubercles
(Figs. 134 L, M); cocoon length basally 4.0-4.5 mm .....................Simulium scutistriatum
– Frontoclypeus and exposed portion of thorax smooth or only with small rounded
tubercles on frontoclypeus base (Fig. 134 C) or posterolaterally ..............................
............................................................................... Simulium guianense, Simulium perplexum

Larvae
Not included: Simulium perplexum.

1 Maximum length 10.0-10.5 mm; anal ring with 280-300 rows with 28-35 hooks
each; body cuticule with sparse trichomes filiform and racket shaped (Fig. 135 S)
..............................................................................................................Simulium scutistriatum
– Maximum length 6.8-8.8 mm; anal ring with 120-220 rows with 17-31 hooks each;
body cuticle with abundant trichomes racket shaped (Figs. 135 G, L) .................. 2
2(1) Anal ring with about 220 rows with 28-31 hooks each ................ Simulium hirtipupa
– Anal ring with 133-170 rows with 17-30 hooks each ...................................................
.................................................. Simulium guianense, Simulium itaunense, Simulium orbitale.

CLAVES DE ESPECIES DE SIMULIUM (THYRSOPELMA)


Hembras
1 Escudo castaño rojizo oscuro, gonapófisis más anchas que largas (Fig. 132 M);
paraprocto subtriangular sin depresión anterior (Fig. 132 N) ........Simulium scutistriatum
– Escudo negruzco; gonapófisis aproximadamente tan largas como anchas (Figs.
132 D, E, H); paraprocto con una depresión anterior profunda, generalmente
mostrando como teniendo una cola (Fig. 132 G) ....................................................... 2
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 615

2(1) Paraprocto continuado distalmente por una prolongación membranosa ancha y


curva (Fig. 132 G) ................................................................................. Simulium hirtipupa
– Paraprocto truncado a corta distancia de la base con una prolongación corta, fina,
membranosa (Fig. 132 D) ................................................................................................. 3
3(2) Uña con pequeño diente subbasal ................................................... Simulium perplexum
– Uña sin diente subbasal (Fig. 132 C) ............................................................................. 4
4(3) Largo ala 1,8-2,5 mm; sector basal de R y Sc sin pelos ................ Simulium guianense
– Largo ala 2,8-3,1 mm; sector basal de R con 20-40 pelos; Sc con 11-15 pelos .....
..................................................................................... Simulium orbitale, Simulium itaunense

Machos
1 Escudo castaño oscuro a negruzco; sin manchas humerales plateadas; placa
ventral sin prolongaciones laterales y con una corta prolongación mediana (Fig.
133 K) o ausente ................................................................................................................ 2
– Escudo negruzco generalmente con manchas humerales plateadas (Figs. 133 A,
G); placa ventral con prolongaciones laterales y prolongación mediana larga (Figs.
133 D, I) ............................................................................................................................... 3
2(1) Placa ventral sin prolongación mediana; gonostilo con espolón apical fino y
aguzado .................................................................................................. Simulium perplexum
– Placa ventral con una corta prolongación mediana (Fig. 133 K); gonostilo con
espolón apical grueso y con ápice romo (Fig. 133 J) ...............Simulium scutistriatum
3(1) Prolongación mediana de la placa ventral subpentagonal y más ancha en la base
que en el ápice (Fig. 130 I); gonostilo relativamente fino basalmente (Fig. 133 H)
................................................................................................................... Simulium hirtipupa
– Prolongación mediana de la placa ventral en forma de pera, con el ápice más
ancho que la base (Fig. 133 D); gonostilo relativamente ancho basalmente (Figs.
133 B, C) .............................................................................................................................. 4
4(3) Escudo con 1+1 mancha plateada extendida desde el área humeral hasta el área
plateada posterior, como una ancha banda submediana (Fig. 133 G); sector basal
de R sin pelos ......................................................................................... Simulium guianense
– Escudo con 1+1 mancha plateada reducida mayormente al área humeral, no
llegando al área posterior (Fig. 133 A); sección basal de R con pelos ................... 5
5(4) Largo ala 2,5-2,7 mm; escudo con 1+1 manchas plateadas anteriores, alargadas
(Fig. 133 A) ................................................................................................ Simulium orbitale
– Largo ala 2,8-3,0 mm; escudo con 1+1 manchas plateadas redondeadas ...............
................................................................................................................... Simulium itaunense

Pupas
1 Branquias con 20-50 ramas .............................................................................................. 2
– Branquias con 12 ramas .................................................................................................... 3
2(1) Branquias con 21-24 ramas (Fig. 134 B); abertura del capullo elevado
frontalmente (Fig. 134 A) ....................................................................... Simulium orbitale
– Branquias con 46-52 ramas; abertura del capullo débilmente elevado frontalmente
(similar a Simulium hirtipupa, Fig. 134 F) ........................................... Simulium itaunense
616 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

3(1) Frontoclípeo y porción expuesta del tórax cubierta por espinas (Figs. 134 F, I);
ramas de las branquias cubiertas por fuertes espículas (Fig. 134 H) ........................
................................................................................................................... Simulium hirtipupa
– Frontoclípeo y porción expuesta del tórax sin espinas; ramas de las branquias sin
espículas o muy débiles ..................................................................................................... 4
4(3) Frontoclípeo y porción expuesta del tórax con numerosos tubérculos con ápice
aguzado (Figs. 134 L, M); largo capullo en la base 4,0-4,5 mm .................................
..............................................................................................................Simulium scutistriatum
– Frontoclípeo y porción expuesta del tórax liso o solamente con pequeños
tubérculos redondeados sobre la base del frontoclípeo (Fig. 134 C) o latero
posteriormente ................................................... Simulium guianense, Simulium perplexum

Larvas
No incluida: Simulium perplexum.

1 Largo máximo 10,0-10,5 mm; anillo anal con 280-300 hileras con 28-35 ganchos
cada una; cutícula del cuerpo con escasos tricomas filiformes y en forma de
raqueta (Fig. 135 S) ..........................................................................Simulium scutistriatum
– Largo máximo 6,8-8,8 mm; anillo anal con 120-220 hileras con 17-31 ganchos
cada uno; cutícula del cuerpo con abundantes tricomas en forma de raqueta (Figs.
135 G, L) .............................................................................................................................. 2
2(1) Anillo anal con alrededor de 220 hileras con 28-31 ganchos cada una ...................
................................................................................................................... Simulium hirtipupa
– Anillo anal con 133-170 hileras con 17-30 ganchos cada una ...................................
................................................... Simulium guianense, Simulium itaunense, Simulium orbitale

Simulium (Thyrsopelma) orbitale LUTZ (Figs. 132 A-F; 133 A-F; 134 A-D; 135 A-G.)
Simulium orbitale LUTZ, 1910: 231-234, 251; LUTZ & MACHADO, 1915: 14-46; NEIVA & PENNA, 1916:
93; LUTZ et al., 1918: 107, 109, 165; LUTZ & NUNES TOVAR, 1928: 47; PINTO, 1931: 706, 708,
741, 756; WYGODZINSKY, 1953a: 303.
Simulium (Thyrsopelma) orbitale: COSCARÓN, 1987: 39; 1991: 287-288.
Simulium nigrimanum (nec MACQUART, 1838: 38): LUTZ, 1909: 137-138, 145-146; LUTZ, 1910: 215;
PINTO, 1931: 705-706, 726, 760.
Simulium albimanum LUTZ, 1909: 127, 132.
Simulium (Eusimulium) orbitale: D’ANDRETTA & D’ANDRETTA, 1946: 85-101.
Thyrsopelma brasiliense ENDERLEIN, 1934b: 284; STONE, 1963: 19; VULCANO, 1967: 17.
Simulium paraguayense (nec SCHROTTKY, 1909: 63): LANE & VULCANO, 1943: 437.
Thyrsopelma orbitale: PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 2004: 131-132.
Simullium (Trichodagmia) orbitale: CROSSKEY & HOWARD, 2004: 74.
Female: Wing length 2.8-3.0 mm. Scutum blackish with whitish pruinosity and hair
grouped like scales, silvery to yellowish, without special arrangement. Scutellum, pleurae
and metanotum blackish, the latter one shiny nacreous. Legs brown with darker apical
portion blackish. Abdomen velvety black on segments I-IV, black, cereus, shiny on V-VII;
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 617

II with silvery pruinosity. Cibarium smooth with sclerotized border (Fig. 132 B). Sc with
about 15-20 hairs. Fronto-ocular triangle slightly wider at base than high (Fig. 132 A).
Claw without subbasal tooth (Fig. 132 C). Eighth sternite with 24-41 setae on each side,
gonapophysis subovoidal, longer than wide and with dense hair (Fig. 132 E), paraproct
relatively narrow and rounded apically, ventrally membranous with short tail-like projec-
tion (Fig. 132 D), genital fork with distal portion expanded (Fig. 132 F).
Male: Wing length 2.5-2.7 mm. Basal portion of R with 20-40 hairs. Scutum blackish
with 1+1 diagonal anterolateral silvery spots reaching anterior 1/3, and joined anteriority
with silvery lateral and posterior margins (Fig. 133 A). Sc without hair. Ratio of length/
width of hind basitarsus = 6. Gonostylus narrow on distal half (Figs. 133 B, C). Ventral
plate with peculiar elongated, subglobose and hairy distal process (Figs. 133 D, E).
Endoparameres without teeth (Fig. 133 F).
Pupa: Cocoon aperture relatively low anteriorly (Fig. 134 A); length basally: 3.1-3.7
mm, on dorsum 2.2-2.7 mm, gills 0.6-0.7 mm. Frontoclypeus with tubercles on basal 1/3
and laterally (Fig. 134 C); exposed portion of thorax smooth. Cephalic trichomes small
and single. Gills with 21-24 branches, emerging without perceptible basal trunk; primary
branches very short; branches medially thick, pseudoannulated and bordered by spiculae;
apically acute with well-sclerotized terminal spine (Fig. 134 B). Abdominal chaetotaxy as
in figure 134 D.
Larva: Maximum length 6.7 mm. Shape as in figure 135 A. Cephalic apotome
scarcely darkened basally, continued anteriorly by median line (Fig. 135 B). Antennae
surpassing apex of cephalic fan stem; ratio of proximal, medial and distal articles =
1:2.2:1-1.1, shape as in figure 135 E. Cephalic fan with about 56 rays. Mandible with
second marginal tooth about half of first (Fig. 135 F). Hypostomium with anterior margin
straight (Figs. 135 C-D); postgenal bridge slightly longer than half of hypostomium
length. Body with hair-like scales racket shaped; length 0.019 mm (Fig. 135 G), arranged in
rows on anterior 5 segments and not ordered on following segments. Anal ring with 160-
166 rows with 25-30 hooks on each. Rectal papillae with 10-15 diverticula on each lobe.
Distribution: Argentina: Buenos Aires, Corrientes, Entre Rios, Misiones; Brazil:
Mato Grosso do Sul, Minas Gerais, Paraná, Rio Grande do Sul, Santa Catarina, São Paulo.
Bionomics: This species breeds in rivers with large discharge and fast flow attached
to submerged tree branches at a depth of about 0.5 m. Female feeding habits are
unknown. In Buenos Aires province, the highest prevalence is in autumn and winter
(COSCARÓN et al., 2000).
Discussion: The closest species is S. guianense, but it can be differentiated by the pupal
gill with 12 branches and male with elongated silvery spots the length of the scutum.

Simulium (Thyrsoplema) guianense WISE (Figs. 133 G; 134 E; 135 G 2.)


Simulium guianense WISE, 1911: 248; 1912: 43; SMART, 1940: 4.
Simulium (Trichodagmia) guianense: SHELLEY et al., 1997: 40-43; 2000: 183-184; CROSSKEY & HOWARD,
1997: 97; 2004: 74.
Simulium (Thyrsopelma) guianense: COSCARÓN, 1987: 39.
Thyrsopelma guianense PY-DANIEL & MOREIRA SAMPAIO, 1995: 119.
618 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

B L A

I
O

K
G
N F

E H

Figure 132. Female. A-F: S. (Thyrsopelma) orbitale; A: fronto-ocular triangle; B: cibarium; C: claw; D:
genitalia, ventral view showing eighth sternite and gonapophysis, cercus and paraproct (at left
mounted on slide) and genital fork; E: eighth sternite and gonapophysis; F: genital fork. G-H: S.
(T.) hirtipupa; G: cercus and paraproct; H: eighth sternite and gonapophysis. I-O: S. (T.) scutistriatum;
I: hair of scutum; J: palpomere with sensory vesicle, K: claw; L: cibarium; M: eighth sternite and
gonapophysis; N: cercus and paraproct; O: genital fork.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 619

G A

E D

C
H

I
K
J

Figure 133. Male. A-F: S. (Thyrsopelma) orbitale; A: scutum; B: genitalia in ventral view showing
gonocoxite and gonostylus; C: gonocoxite and gonostylus; D-E: ventral plate, ventral view (D) and
lateral view (E); F: endoparameres and aedeagal membrane. G: S. (T.) guianense, scutum. H-I: S. (T.)
hirtipupa; H: gonocoxite and gonostylus; I: ventral plate. J-K: S. (T.) scutistriatum; J: gonocoxite and
gonostylus; K: ventral plate.
620 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Simulium pintoi D’ANDRETTA & D’ANDRETTA, 1946: 101-103; SHELLEY et al., 1979: 41-43; LACEY &
CHARLWOOD, 1980: 496; LACEY, 1981: 330.
Simulium (Thyrsoplema) pintoi COSCARÓN, 1987, 39; 1991: 288-290.
Simulium ortizi RAMÍREZ PÉREZ, 1971: 336-339.
“A”, “B”, “C”,”D” CHARALAMBOUS et al.,1996: 111-120 (Simulium guianense cytoform).
Female: Wing length 1.8-2.5 mm. Coloration of scutum, scutellum, metanotum pleurae
and abdomen as in S. orbitale. Sc without hair. Basal sector of R with single row of hairs,
Cibarium, paraproct and cercus as in S. orbitale; eighth sternite with about 20 setae on each side.
Male: Wing length 1.9-2.3 mm. Scutum velvety black with 1+1 silvery submedian
stripes, extending from humeral callus on anterolateral area to posterior area; another
silvery stripe on lateral and posterior margin (Fig. 133 G). Basal sector of R and Sc
without hair. Ratio of length/width of hind basitarsus = 6. Genitalia as in S. orbitale.
Pupa: Cocoon as in S. orbitale; length basally 2.5-3.0 mm; gills 1.0 mm. Frontoclypeus
with small tubercles on basal 1/3, as in S. orbitale and on thorax only dorsally on midline.
Gills with 12 branches arranged as in S. orbitale (Fig. 134 E).
Larva: Maximum length 6.0 mm. Cephalic apotome, mandible and antenna as in S.
orbitale. Cephalic fan with 54-58 rays. Hypostomium border with teeth arranged as in S.
hirtipupa. Body cuticle with abundant trichomes racket shaped, bigger than in other
species, length 0.028 mm (Fig. 135 G 2). Anal ring with about 140 rows with 23-25 hooks
each. Rectal papillae with 5-12 diverticula on each lobe.
Distribution: Brazil: Amapá, Amazonas, Espírito Santo, Goiás, Maranhão, Mato
Grosso, Minas Gerais, Pará, Paraná, Rio de Janeiro, Roraima, Santa Catarina, São Paulo,
Tocantins; Venezuela: Amazonas, Bolivar; French Guiana; Suriname.
Bionomics: The larval stages were collected in moderately fast creeks 3-5 m wide and
0.4-0.5 m deep, but they can be found in larger streams (SHELLEY et al. 2000) with sandy
bottoms in yellowish water and attached to aquatic vegetation especially Podostemaceae.
Females are zoophilic and anthropophilic. They are important vectors of Onchocerca volvulus.
Discussion: The closest species is S. orbitale, under which the differences were
explained. Synonymy with S. pintoi was commented on by SHELLEY et al. (1997). Studies of
CHARALAMBOUS et al. (1996) showed the presence of 4 cytospecies, which could explain
the differences in feeding behavior.

Simulium (Thyrsopelma) itaunense D’ANDRETTA & GONZÁLEZ


Simulium itaunensis D’ANDRETTA & GONZÁLEZ, 1964: 106-108; VULCANO, 1967: 14; 1977: 292; PY-
DANIEL et al., 1985: 98-99.
Simulium (Thyrsopelma) itaunense: COSCARÓN, 1987: 39; 1991: 290-291.
Thyrsopelma itaunense: PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 2004: 131.
Simulium (Trichodagmia) itaunense: CROSSKEY & HOWARD, 1997: 79; 2004: 74; HERNÁNDEZ et al., 2005: 3-12.
Female: Wing length 2.8-3.1 mm. Scutum blackish, humeri brown with grey pruinosity
and recumbent grouped whitish hairs. Scutellum and metanotum black, the latter with silver
pruinosity. Pleura blackish gray pruinose. Legs brown with darkened portion dark brown.
Abdomen shiny blackish. Cibarium, wings, claws and genitalia as in S. orbitale.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 621

F
C

J L
M

Figure 134. Pupa. A-D: S. (Thyrsopelma) orbitale; A: Pupa lateral view; B: thorax, anterodorsal
portion and gill; C: frontoclypeus; D; abdominal chaetotaxy. E: S. (T.) guianense, gill. F-I: S. (T.)
hirtipupa; F: pupa; G: gill; H: gill apex enlarged view; I: frontoclypeus. J-M: S. (T.) scutistriatum; J:
lateral view; K: gill; L: frontoclypeus; M: integument tubercles enlarged.
622 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Male: Wing length 2.8-3.0 mm. Scutum velvety black, nacreous shiny on lateral
margins and prescutellum region, and 1+1 silvery rounded spots on anterior 1/3; humeri
with silver pruinosity. Scutellum, metanotum and pleurae blackish. Abdomen velvety
blackish with 1+1 lateral silvery spots on tergites II, VI-VIII, and anterior submedian are
on segments IV-V. Basal sector of R haired and Sc with hair only on base. Genitalia similar
to those of S. orbitale, but paraproct with distal membranous area more evident.
Pupa: Cocoon aperture weakly elevated anteriorly, in lateral view similar to S.
hirtipupa. Length basally 4.5-4.9 mm; dorsally 3.4-4.3 mm; gills 0.7-1.2 mm. Frontoclypeal
and thoracic tubercles and trichomes as in S. orbitale, thoracic trichomes robust. Gill with
46-52 branches, relatively thick and short, emerging from thick and elongated basal trunk
horizontally arranged; branches apically acute and well sclerotized, with spine-like aspect,
similar to arrangement in S. orbitale but with more branches.
Larva: Maximum length 6.8 mm. Antenna, hypostomial teeth and cuticular body
hair as in S. orbitale. Cephalic fan with 48 rays. Postgenal bridge slightly shorter than
hypostomium length. Lateral sclerite of thoracic proleg with about 65 teeth arranged in
14-16 groups. Anal ring with about 133 rows of 17-22 hooks each. Rectal papillae with
diverticula on each lobe.
Distribution: Brazil: Rio Grande do Sul, Santa Catarina, São Paulo.
Bionomics: This species breeds in clean creeks with fast, semi-torrential flow,
attached to stones and aquatic vegetation. Female feeding habits are unknown.
Discussion: The closest species is S. guianense. Males of S. guianense can be separated
by the elongated anterior silvery spots and the pupa by the 12 gill branches.

Simulium (Thyrsopelma) hirtipupa LUTZ (Figs. 132 G, H; 133 H, I; 134 F-I; 135 H-L.)
Simulium hirtipupa LUTZ, 1910: 200, 260, 263-264; PINTO, 1931: 702, 756; VARGAS, 1945a: 146;
VULCANO, 1967: 13, 1971: 292; MAIA-HERZOG et al., 1985: 483-490.
Simulium (Thyrsopelma) hirtipupa: COSCARÓN, 1987: 39; 1991: 292-293.
Thyrsopelma hirtipupa: PY-DANIEL & MOREIRA SAMPAIO, 1995: 119.
Simulium (Trichodagmia) hirtipupa: CROSSKEY & HOWARD, 1997: 79; 2004: 74.
Female: Wing length 2.7-2.9 mm. Scutum black with thin gray pruinosity and thin
median and 1+1 sublateral stripes extending 2/3 of scutum length, scutum hair grouped like
scales in longitudinal rows. Scutellum black, metanotum velvety blackish. Pleurae blackish
with gray pollinosity. Abdomen dark grayish brown with tergal plates blackish. Cibarium as
in S. orbitale. Fronto-ocular triangle short, higher than wide. Basal section of R and Sc
without hair. Paraproct apically continued by relatively wide and curved membranous
process with abundant hair (Fig. 132 G). Eighth sternite with 12-16 setae on each side,
gonapophysis subtriangular, about as long as wide at base with blunt apex (Fig. 132 H).
Male: Wing length 2.6-2.8 mm. Scutum blackish with much gray pollinosity and golden
pile, with frontal illumination becoming leaden gray, and with posterior light blackish with
1+1 submedian grayish stripes on posterior 2/3. Scutellum and metanotum as in female.
Abdomen black with 1+1 silvery spots on segments II, V-VIII. Gonostylus nearly 1.5 times
longer than gonocoxite, becoming thinner distally (Fig. 133 H), basally narrower than in S.
orbitale. Ventral plate with 1+1 lateral and 1 wider median processes (Fig. 133 I).
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 623

A
D
C

B 1

G
2

F
H E
I

J
K

O
M
P

Q
S
R

Figure 135. Larva. A-G: S. (Thyrsopelma) orbitale; A: larva in lateral view; B: head, dorsal view; C:
hypostomium; D: teeth of hypostomium, anterior margin; E: antenna; F: mandibular teeth; G:
integument trichomes (1: S. (T.) orbitale; 2: S. (T.) guianense). H-L: S. (T.) hirtipupa; H: larva, lateral
view; I: teeth of hypostomium, anterior margin; J: antenna; K: mandibular teeth; L: integument
trichomes. M-S: S. (T.) scutistriatum; M: postgenal cleft and hypostomium; N: teeth of hypostomium,
anterior margin; O: antenna; P: mandibular teeth; Q: dorsal posterior of head and cervical sclerites;
R: anal sclerite (ventral branches up); S: integument trichomes.
624 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Pupa: Cocoon aperture lower anteriorly (Fig. 134 F), weave thick with wrinkled
surface. Length basally 4.0-5.0 mm, dorsally 2.6-3.6 mm, gills 0.7 mm. Frontoclypeus and
thorax with long spines (Fig. 134 I), these spines also in other places on head but smaller.
Integument without tubercles. Gill with 12 branches relatively thick, with spine on apex
and annulations covered with irregular shaped spiculae (Figs. 134 G, H).
Larva: Maximum length 7.3-8.8 mm. General aspect as in figure 135 H. Cephalic
apotome brown, darker at base. Postgenal bridge slightly shorter than hypostomium
length; teeth with anterior margin of hypostomium very short (Fig. 135 I). Antennae
surpassing length of cephalic fan stem; shape as in figure 135 J, ratio of proximal, medial
and distal articles = 1:2-2.2:1-2. Mandible with long and thin first marginal tooth (Fig. 135
K). Cephalic fan with 55-60 rays. Body integument with petaloid hairs fewer in number
than in S. orbitale and S. pintoi, length about 0.021 mm (Fig. 135 L). Lateral sclerite of
thoracic proleg with 54-56 teeth arranged in 13-15 groups. Anal ring with 220-246 rows
with 28-36 hooks each. Rectal papillae with 12-14 diverticula on each lobe.
Distribution: Brazil: Bahia, Minas Gerais, São Paulo.
Bionomics: The species breeds in fast-flowing, shallow creeks with rocky bottoms.
Female feeding habits are unknown.
Discussion: Adults are similar to those of several species of the subgenus, but the
peculiar spines on the head and thorax of the pupa and the cocoon and pupa gill shape
permit recognition of S. hirtipupa.

Simulium (Thyrsopelma) perplexum SHELLEY, MAIA-HERZOG, LUNA DIAS & COUCH


Simulium (Trichodagmia) perplexum SHELLEY et al., 1989: 343. CROSSKEY & HOWARD, 1997: 79; 2004: 74;
SHELLEY et al., 2004: 446, 453, 469.
Thyrsopelma perplexa: PY-DANIEL & MOREIRA SAMPAIO, 1995: 119.
(Diagnosis based on original description).
Female: Wing length 2.3-2.8 mm. Coloration and morphology similar to that for S.
guianense, but with 1+1 lateral dark transverse spots on sternites II-VII, scutum hair
arranged regularly in small groups, absence of hair on basal sector of R, presence of short
subbasal tooth on hind leg claw and paraprocts without anterior tail-like projection.
Male: Wing length 2.1-2.6 mm. Coloration and general morphology similar to that of
S. guianense. Scutum with anterior illumination velvety black with dark longitudinal median
line on anterior 2/3, changing light to posterior position shows light grey. Pilosity arranged
in groups of hair-like scales, shiny golden to green intermixed with thin dark longer hairs on
anterior and posterior edge. Gonostylus as long as gonocoxite; gonostylus with base thicker
than in S. guianense and with apical spur pointed; ventral plate without median distal process.
Pupa: Similar to S. guianense but gill branch annulations with border continued
anteriorly as small processes instead of spiculae as in S. guianense.
Larva: Unknown.
Distribution: Guyana.
Bionomics: Unknown.
Discussion: This species is very close to S. guianense. In males of S. guianense, the
biggest differences are in male coloration, the 1+1 silvery longitudinal stripes extended
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 625

from the humeri to the posterior area, the gonostylus more elongated and the ventral plate
with a median distal process. The female of S. guianense has a single row of setae on the
basal section of R, a tail-like projection on the paraproct and the claw of the hind leg
without a subbasal tooth.

Simulium (Thyrsopelma) scutistriatum LUTZ (Figs. 132 I-O; 133 J, K; 134 J-M, 135 M-S.)
Simulium scutistriatum LUTZ, 1909: 133-134; 1910: 225-226; LUTZ, 1922: 90; PINTO, 1931: 712, 725, 726,
762; VARGAS, 1945a: 195; VULCANO, 1967: 20; 1971: 293; MAIA-HERZOG et al., 1984: 350-351.
Eusimulium scutistriatum: LANE & PORTO, 1939: 169, 173-174, LANE & VULCANO, 1943: 436.
Simulium (Thyrsopelma) scutistriatum: COSCARÓN, 1987: 39; 1991: 294-296.
Simulium (Trichodagmia) scutistriatum CROSSKEY & HOWARD, 1997: 79; 2004: 74.
Thyrsopelma scutistriata: PY-DANIEL & MOREIRA SAMPAIO, 1995: 119.
Female: Wing length 3.0-3.6 mm. Scutum dark reddish brown, without special
ornamentation, with golden to tin, small, shiny pile, grouped, simulating scales (Fig. 132
I) and arranged in longitudinal rows, not continuous. Scutellum yellowish brown and
metanotum velvety blackish brown. Pleurae dark grayish brown with gray pollinostity.
Legs brown with darker areas dark brown and pale yellowish. Abdomen brown with
tergal plates dark brown. Fronto-ocular triangle about as wide as high. Basal sector of R
haired, Sc with about 40 hair. Sensory vesicle of maxillary palp shorter than half of
palpomere length (Fig. 132 J). Claw without subbasal tooth (Fig. 132 K). Cibarium
smooth with cornuae well sclerotized (Fig. 132 L). Eighth sternite with concavity and
well-sclerotized median area and with about 26 setae on each side; gonapophysis
subovoidal, wider than long (Fig. 132 M). Paraproct subtriangular, without depression
on anterior border (Fig. 132 N). Genital fork with distal portion of anterior branches
very wide (Fig. 132 O).
Male: Wing length 2.9-3.3 mm. Coloration as in female but darker. Scutum dark brown
with dark pollinosity increased on posterior region, with 1 median and 1+1 submedian
longitudinal stripes, the last ones not reaching anterior margin; with changing illumination,
scutum shows grayish pollinosity. Abdomen blackish with 1+1 silvery spots on segment I, V-
VII. Gonostylus relatively thick and shorter than in other species of subgenus (Fig. 133 J).
Ventral plate body proportionately long and median process shorter (Fig. 133 K).
Pupa: Cocoon with aperture inclined basally (Fig. 134 J). Length basally 4.0-4.5 mm,
dorsally 3.8-4.0 mm, gill 1.2 mm. Weave resistant, threads not visible. Frontoclypeus and
exposed portion of thorax with abundant, apically acute tubercles (Figs. 134 L-M);
cephalic and thoracic trichomes single and relatively short. Gills with 12 branches,
relatively short, with thick and acute apices (Fig. 134 K).
Larva: Maximum length 10.0-10.5mm. Cephalic apotome darkened on median line and
base, with positive spots. Antenna as long as or longer than cephalic fan stem; ratio of
proximal, medial and distal articles 1:2-2.3:0.9-1. Postgenal bridge about same length as
hypostomium (Fig. 135 M); hypostomial anterior border with large lateral teeth (Fig. 135 N);
antenna as in figure 135 O; marginal teeth subequal in shape and relatively short and thick (Fig.
135 P). Cephalic fan with about 53 teeth. Cervical sclerite wide (Fig. 135 Q). Anal ring with
280-300 rows with 28-35 hooks each. Rectal papillae with 30 diverticula on each lobe. Anal
626 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

sclerite with hair and scales (Fig. 135 R). Body integument with hair racket shaped, intermixed
with filamentous hairs (Fig. 135 S). Lateral sclerite of thoracic proleg with 56 teeth.
Distribution: Brazil: Goiás, Mato Grosso, Minas Gerais, Paraná, Rio de Janeiro, São
Paulo, Tocantins; Colombia; Paraguay; Venezuela.
Bionomics: The larval stages can be found in fast-flowing creeks, attached to roots
and stones. Females were collected on horses and around humans.
Discussion: The similar species of the subgenus can be differentiated in the female
by the blackish scutum and shorter paraprocts with the distal portion membranous; in the
male by the blackish scutum with wide silvery stripes or big humeral spots and the ventral
plate body with lateral and large median processes; and in the pupa by the absence of
tubercles with acute apices on the frontoclypeus and thorax.

Species inquirenda

Simulium albopictum LANE & PORTO


Simulium albopictum LANE & PORTO, 1940: 193-194.
Simulium (Thyrsopelma) albopictum: COSCARÓN, 1987: 39; 1991: 296.
Simulium (Trichodagmia) albopictum: CROSSKEY & HOWARRD, 2004: 74.
Thyrsopelma albopicta: PY-DANIEL & MOREIRA SAMPAIO, 1995: 119.
Simulium albopictum was described based on 3 females deposited in Facultade de
Saude Publica of São Paulo University. They are externally similar to those of S. guianense
but without knowledge of the other stages, it is impossible to be sure if they are
conspecific. We prefer to maintain S. albopictum as species inquirenda.
Distribution: Brazil: Paraná.

Species not placed to subgenus because of insufficient information

Simulium costaricense SMART, 1944: 132 (n. nom. Acropogom rufidorsum ENDERLEIN, 1936:
119) (Costa Rica).
Scarce information does not permit us to place S. costacricense in any supraspecific
taxon; it probably falls into the S. (Ectemnaspis) DINELLII species group, based on the
scutum coloration.

Simulium diaznajerai VARGAS, 1943a: 136 (n. nom. Odagmia tenuifrons ENDERLEIN, 1936:
130) (Mexico).
Based on the description, it is impossible to identify the species.

Simulium nigrifemur (ENDERLEIN), 1936: 118 (Acropogon). (Chile: Cauquenes).


With an insufficient original description and without the location of the type
material, species identification is doubtful.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 627

6
MAPS OF DISTRIBUTION
628 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 629

Tlalocomyia
revelata

Mayacnephia
aguirrei
atzompensis
fortunensis
grenieri
mixensis
muzquicensis
pachecolundi
roblesi
salasi
tadai

Araucnephia
montana
iberaensis
Araucnephioides
schlingeri

Cnesia
dissimilis
ornata
pusilla

Cnesiamima
atroparva

Paraustrosimulium
anthracinum

Lutzsimulium
flavopubescens
hirticosta
pernigrum

Kempfsimulium
simplicicolor

Figure 136. Tlalocomyia, Mayacnephia, Araucnephia, Araucnephioides, Cnesia, Cnesiamima, Paraustrosimulium,


Lutzsimulium, Kempfsimulium.
630 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Simulium (Pternaspatha)

“NIGRISTRIGATUM GROUP”

S. colopaxi
S. deagostini
S. diamantinum
S. dureti
S. horcochuspi
S. limay
S. nigristrigatum
S. pichi
S. pulchrum
S. simile
S. strigidorsum

Figure 137. Simulium (Pternaspatha), nigristrigatum species group.


ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 631

Simulium (Pternaspatha)

“NEMORALE GROUP”

S. albicinctum
S. albilineatum
S. annulatum
S. barbatipes
S. bordai
S. caprii
S. hectorvargasi
S. herreri
S. luchoi
S. nemorale
S. philippii
S. prodexargenteum
S. quechuanum
S. stelliferum
S. yacuchuspi

Figure 138. Simulium (Pternaspatha), nemorale species group.


632 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Simulium (Notolepria)

S. blantoni
S. cuasiexiguum
S. exiguum
S. llutense
S. paraguayense
S. subexiguum
S. gonzalezi

Figure 139. Simulium (Notolepria).


ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 633

Simulium (Chirostilbia)

“PERTINAX GROUP”

S. bifenestratum
S. distinctum
S. empascae
S. friedlanderi
S. laneportoi
S. obesum
“SUBPALLIDUM GROUP”
S. pertinax
S. acarayense
S. riograndense
S. serranum S. papaveroi

S. spinibranchium S. subpallidum

Figure 140. Simulium (Chirostilbia), pertinax and subpallidum species group.


634 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Simulium (Inaequialium)

“INAEQUALE GROUP”

S. beaupertuyi

S. clarki

S. clavibranchium

S. diversibranchium

S. inaequale

S. leapoldense

S. mariavulcanoae

S. nahimi
“BOTULIBRANCHIUM GROUP”
S. nogueirai

S. parimaensis S. botulibranchium

S. rappae S. petropoliense

S. subclavibranchium S. souzalopesi

S. subnigrum

S. travassosi

Figure 141. Simulium (Inaequialium), inaequale and botulibranchium species group.


ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 635

Simulium (Psaroniocompsa)

“INCRUSTATUM GROUP”

S. anamariae

S. angrense

S. auripellitum

S. auristriatum
“SIOLII GROUP”
S. bonaerense

S. brevifurcatum S. damascenoi

S. fuliginis S. quaporense

S. incrustatun S. lourencoi

S. jujuyense S. siolii

S. limbatum S. tergospinosum

S. minuanum

S. schmidtmummi

Figure 142. Simulium (Psaroniocompsa), incrustatum and siolii species group.


636 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Simulium (Cerqueirellum)

S. amazonicum
S. argentiscutum
S. chaquense
S. cuneatum
S. delponteianum
S. ganalesense
S. minusculum
S. oyapockense
S. raraimense
S. sanguineum
S. venezuelense

Figure 143. Simulium (Cerqueirellum).


ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 637

Simulium (Coscaroniellum)

“QUADRIFIDUM GROUP”
S. cauchense
S. cerradense
S. dolianhanhani
S. goeldii
S. quadrifidum
S. ulyssesi

“QUADRIVITTATUM GROUP”
S. quadrivittatum

Figure 144. Simulium (Coscaroniellum), quadrifidum and quadrivittaum species group.


638 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Simulium (Ectemnaspis)

“BICOLORATUM GROUP”

S. anaimense

S. antonii

S. arcabucense

S. bicoloratum

S. bicornutum

S. cormonsi

S. furcillatum

S. ignescens

S. jaimeramirezi

S. pautense

S. pifanoi

S. roquemayu

S. rubiginosum

S. tolimaense

S. tunja

Figure 145. Simulium (Ectemnaspis), bicoloratum species group.


ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 639

Simulium (Ectemnaspis)

“ROMANAI GROUP”

S. albanense

S. gabaldoni

S. lutzianum

S. mayuchuspi

S. romanai

S. wolffhuegeli

Figure 146. Simulium (Ectemnaspis), romanai species group.


640 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Simulium (Ectemnaspis)

“PERFLAVUM GROUP”

S. kabanayense

S. maroniense

S. perflavum

S. rorotaense

S. suarezi

S. trombetense

Figure 147. Simulium (Ectemnaspis), perflavum species group.


ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 641

Simulium (Ectemnaspis)

“DINELLII GROUP”

S. antillarum
S. dinellii
S. nuneztovari
S. ochraceum
S. shewellianum

Figure 148. Simulium (Ectemnaspis), dinellii species group.


642 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Simulium (Psilopelmia)

“BLANCASI GROUP”
S. blancasi
S. penai
S. tenuipes

“ESCOMELI GROUP”
S. bivittatum
S. bobpetersoni
S. callidum
S. dandrettai
S. downsi
S. dugesi
S. escomeli
S. gonzalezherrejoni
S. haematopotum
S. iracouboense
S. longithallum
S. mangabeirai
S. mediovittatum
S. ochoai
S. panamense
S. pseudocallidum
S. samboni
S. travisi
S. trivittatum
S. veracruzanum
S. zempoalense

Figure 149. Simulium (Psilopelmia), escomeli and blancasi species group.


ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 643

Simulium (Aspathia)

S. anduzei
S. metallicum
S. bustosi S. morae
S. costalimai S. parrai
S. covagarciai S. piperi
S. hetchi S. puigi
S. horacioi S. putre
S. hunteri S. racenisi
S. iriartei S. ruizi
S. jacumbae S. sandyi
S. jobbinsi S. spilmani
S. kompi S. tricorne
S. marquezi S. wygoi

S. matteabranchium

Figure 150. Simulium (Aspathia).


644 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Simulium (Hemicnetha)

“PAYNEI GROUP”
S. bricenoi
S. paynei
S. rubrithorax
S. virgatum

“MEXICANUM GROUP”
S. biuxinisa
S. earlei
S. freemani
S. hinmani
S. keenani
S. mexicanum
S. lobatai
S. placidum
S. seriatum
S. smarti
S. hieroglyphicum
S. yepocapense

“BRACHYCLADUM GROUP”
S. brachycladum
S. cristalinum
S. guerrerense
S. pulverulentum
S. solarii

“OVIEDOI GROUP”
S. oviedoi
S. rivasi

Figure 151. Simulium (Hemicnetha), paynei, mexicanum, brachycladum and oviedoi species group.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 645

Simulium (Hearlea)
S. ayrozai S. ethelae
S. burchi S. gorirossiae
S. canadense S. johnsoni
S. capricome S. juarezi
S. carolinae S. larvispinosum
S. contrerense S. menchacai
S. chiriquiense S. microbranchium
S. dalmati S. nigricome
S. delatorrei S. paracarolinae
S. estevezi S. temascalense

Figure 152. Simulium (Hearlea).


646 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Simulium (Trichodagmia)

S. huairayacu
S. lahillei
S. muiscorum
S. nigrimanum
S. sumapazense
S. townsendi
S. wygodzinskyorum

Figure 153. Simulium (Trichodagmia).


ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 647

Simulium (Thyrsopelma)

S. quianense
S. hirtipupa
S. itaunense
S. orbitale
S. perplexum
S. scutistriatum

Figure 154. Simulium (Thyrsopelma).


648 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

7
REFERENCES
ADLER, P.H., CURRIE, D.C. & D. M. WOOD (2004): The black flies (Diptera: Simuliidae) of North
America. - Cornell University Press, Ithaca, New York, 941 pp.
ALENCAR, Y.B., VEIGA LUDVIG, T.A., SOARES, C. & N. HAMADA (2001): Stomach content analysis of
Simulium perflavum ROUBAUD 1905 (Diptera: Simuliidae) larvae from stream in Central
Amazônia, Brazil. - Memorias do Instituto Oswaldo Cruz (Rio de Janeiro) 96: 571-596.
ALVAN-AGUILAR, M.A., HAMADA, N., ADLER, P.H. & S. L.B. LUZ (2005): Cytotaxonomy of Simulium
cauchense FLOCH & ABONNENC and Simulium quadrifidum LUTZ. (Diptera: Simuliidae) in Brazilian
Amazonia. - Memorias do Instituto Oswaldo Cruz (Rio de Janeiro) 100: 249-257.
ANDRADE, C.F. (1989): Manejo integrado de borrachudos. - Seminario sobre insectos e acaros. -
Actas XI Congreso Brasileiro de Entomologia, Campinhas, Fundação Cargill: 141-157.
ANDUZE, P.J. (l947): Simulium matteabranchia. - Boletin de la Academia de Ciencias Físicas,
Matemáticas y Naturales 10 (30): 374-376.
ARTEAGA, L.T. & P. MUÑOZ DE HOYOS (1999): New cytotype in the Simulium metallicum complex
(Diptera: Simuliidae) from Cundinamarca, Colombia. - Journal of Medical Entomology 36:
133-140.
BARBOSA DE ALMEIDA, J.B. (1979): Simulium cerqueirai, nomen novum para Simulium septentrionale
CERQUEIRA & ALMEIDA, 1970 (Diptera, Simuliidae). - Acta Amazônica 9 (4): 789.
BARRETO, P. (1969): The species of black flies found in Colombia (Diptera: Simuliidae). - Journal of
the New York Entomological Society 77 (1): 31-35.
BECKERT, T. (1908): Dipteren der Kanarischen Inseln. - Mitteilungen aus dem Zoologischen
Museum in Berlin 4:1-180 + 4 plates.
BELLARDI, L. (1859): Saggio di ditterología messicana. Parte. I. - Memoire della Reale Accademia
delle Scienze di Torino, Serie 2, 19: 1-80, 2 pls.
BELLARDI, L. (1862): Saggio di ditterologia messicana. Appendice. - Memoire della Reale Accademia
delle Scienze di Torino, Serie 2, 21: 1-18, 1 pl.
BEQUAERT, J. (l934): Part III. Notes on the black-flies of Simuliidae, with special reference to those
of the Onchocerca region of Guatemala. - In: STRONG, R.P., SOUNDGROUND, J.H., BEQUAERT, J.C.
& M. MUNOZ OCHOA (eds): Onchocerciasis, with special reference to the Central American
form of the disease: l75-224. Contributions from the Department of Tropical Medicine and
the Institute for Tropical Biology and Medicine 6: 1-234.
BEQUAERT, J. (1945): Dr. LUIS VARGAS on American black-flies - a review, with critical notes
(Diptera). - Bulletin of the Brooklyn Entomological Society 40: 111-115.
BIGOT, E. (1888): Diptères. - In: Mission Scientifique du Cap Horn 1882-1883. - Gauthier-Villars et
Fils, Paris 6, Zoology: 1-45.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 649

BRICENO-IRAGORRY, L. (1941): Nota sobre Simuliidae (Diptera), con citación de una nueva especie
para el país y lista de las conocidas en Venezuela. - Boletin del Laboratorio de la Clínica “Luis
Razetti” 2 (6): 110-118.
BUENO, M.L., MONCADA L. & P. MUÑOZ DE HOYOS (1979): Simuliidae (Insecta - Diptera) de
Colombia I. Nueva especie de Simulium (Hemicnetha). - Caldasia 12 (60): 581-594.
CAMPOS, J. & C.F.S. ANDRADE (2001): Considerações sobre os simulideos (Diptera, Nematocera) e
seu controle. - Entomologia e Vectores 8 (1): 27-50.
CERQUEIRA, N. & F. BARBOSA DE ALMEIDA (1970): Simuliidae de Amazônia. V. Descrição de
Simulium septentrionale sp. n. (Diptera, Nematocera). - Boletim do INPA (Manaus), Patologia
Tropical 2: 1-11.
CERQUEIRA, N. & J.A. NUNES DE MELLO (1967): Simuliidae da Amazônia (II). Descrição de Simulium
goeldii sp. n. (Diptera, Nematocera). - Amazoniana 1 (2): 125-130.
CERQUEIRA, J. & J.A. NUNES DE MELLO (l968): Simuliidae de Amazônia (IV). Descrição de Simulium
fulvinotum sp. n. (Diptera, Nematocera) - Amazoniana 1 (3): 205-210.
CHARALAMBOUS, M., SHELLEY, A.J. & M. ARZUBE (1993a): The distribution and taxonomic status of
chromosomal forms of the onchocerciasis vector Simulium exiguum (Diptera: Simuliidae). -
Medical and Veterinary Entomology 7: 299-305.
CHARALAMBOUS, M., SHELLEY, A.J., MAIA-HERZOG, M. & A.P.A. LUNA DIAS (1996): Four new
cytotypes of the onchocerciasis vector blackfly Simulium guianense in Brazil. - Medical and
Veterinary Entomology 10: 111-120.
CHARALAMBOUS, M., LOWRY, C.A., LOWELL, S., SHELLEY, A.J. & M. ARZUBE (l997): The value of larval
head pattern for differentiating Simulium exiguum s. l. and S. gonzalezi (Diptera: Simuliidae) in
the onchocerciasis focus of Ecuador. - Bulletin of Entomological Research 87: 19-24.
CHARALAMBOUS, M., READY, P.D., SHELLEY, A.J., ARZUBE, M. & C.A. LOWRY (1993b): Cytological and
isoenzyme analysis of the Bucay and Quevedo cytotypes of the onchocerciasis vector
Simulium exiguum (Diptera: Nematocera) in Ecuador. - Memorias do Instituto Oswaldo Cruz
(Rio de Janeiro) 88: 39-48.
CONN, J., ROTHFELS, K.H., PROCUNIER, W.S. & H. HIRAI (1989): The Simulium metallicum species
complex (Diptera: Simuliidae) in Latin America: a cytological study. - Canadian Journal of
Zoology 67: 1217-1245.
COQUILLETT, D.W. (1902): New Diptera from North America. - Proceedings of the United States
National Museum 25 (1280): 83-l26.
COSCARÓN, S. (1968): Ampliación del área de distribución de Simuliidae (Diptera) en Argentina. -
Revista de la Sociedad Entomológica Argentina 30: 65-67.
COSCARÓN, S. (1971): Notas sobre Simúlidos Neotropicales I. Sobre una nueva especie del Norte
Argentino (Diptera, Simuliidae). - Revista de la Sociedad Entomológica Argentina 33: 33-41.
COSCARÓN, S. (1976a): Las especies de “jerjeles” (Simuliidae, Diptera, Insecta) de la zona de Arica.
- Idesia 4: 25-34.
COSCARÓN, S. (1976b): Notas sobre Simúlidos Neotropicales VI. Sobre dos especies nuevas de
jejenes de la provincia de Misiones, Argentina (Diptera, Insecta). - Revista de la Sociedad
Entomológica Argentina 35: 147-154.
COSCARÓN, S. (1980): Notas sobre Simúlidos Neotropicales IX. Sobre un grupo de especies
próximo al subgénero Simulium (Inaequalium) (Diptera - Insecta). - Revista de la Sociedad
Entomológica Argentina 39 (3-4): 293-302.
650 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

COSCARÓN, S. (1981a): Notas sobre Simúlidos Neotropicales XI. Sobre el subgénero Simulium
(Chirostilbia) ENDERLEIN, con la descripción de dos especies nuevas del S.E. del Brasil (Diptera
- Insecta). - Revista de la Sociedad Entomológica Argentina 40 (1-4): 157-164.
COSCARÓN, S. (1981b): Insecta, Diptera, Simuliidae. - In: RINGUELET, R. (ed.): Fauna de Agua Dulce
de la Republica Argentina. Vol. 38 (1). FECIC, Buenos Aires: 105 pp.
COSCARÓN, S. (1982): Notas sobre Simúlidos Neotropicales X. Sobre un grupo peculiar de especies
del género Simulium del S.E. de Brasil, Paraguay y N.O. de Argentina (Diptera - Insecta). -
Revista de la Sociedad Entomológica Argentina 41 (1-4): 65-76.
COSCARÓN, S. (1983): Notas sobre Simúlidos Neotropicales XII. Las especies de Simulium “grupo
amazonicum” de Argentina, Sur de Brasil y Paraguay (Simuliidae - Diptera). - Revista de la
Sociedad Entomológica Argentina 42 (1-4): 89-99.
COSCARÓN, S. (1984): Revisión del Subgénero Simulium (Ectemnaspis) ENDERLEIN (Simuliidae,
Diptera, Insecta). - Revista de la Sociedad Entomológica Argentina 43 (1-4): 238-325.
COSCARÓN, S. (1985): Sobre la larva de Cnesiamina atroparva EDWARDS y su significado para
interpretar las relaciones filogenéticas de los simúlidos notogeicos (Diptera - Insecta). -
Boletín de la Sociedad de Biología, Concepción, Chile 56: 67-70.
COSCARÓN, S. (1987): El género Simulium LATREILLE en la región Neotropical: Análisis de los grupos
supraespecíficos, especies que los integran y distribución geográfica (Simuliidae, Diptera,
Insecta). - Museu Paraense Emilio Goeldi; Col. E. Snethlage: 111 pp.
COSCARÓN, S. (1989): Los estudios ecológicos en Simúlidos Neotropicales (Diptera-Insecta). - Actas
XI Congreso Sociedade Entomológica Brasileira (Campinas 12-18 de julio de 1987). -
Fundação Cargill, Seminario Insetos e Acaros. Anais 5: 69-98.
COSCARÓN, S. (1990): Taxonomia y Distribución del subgénero Simulium (Ectemnaspis) ENDERLEIN
(Simulidae, Diptera, Insecta). - Iheringia, Sér. Zoologia (Porto Alegre) 70: 109-170.
COSCARÓN, S. (1991): Insecta, Diptera, Simuliidae. - In: CASTELLANOS, Z.A. DE (ed.): Fauna de Agua
Dulce de la República Argentina. Vol. 38 (2). FECIC, Buenos Aires: 304 pp.
COSCARÓN, S. (1994): Los Simuliidae (Diptera) y su importancia sanitaria en Latinoamérica. -
Memorias XXI Congreso Sociedad Colombiana de Entomología (Medellín): 357-366.
COSCARÓN, S. (2003): Biodiversidad de Simuliidae y Tabanidae del Iberá. - In: ALVAREZ, B. (ed.):
Fauna del Iberá. Eudene (Corrientes): 3-6.
COSCARÓN, S. & C.L. COSCARÓN ARIAS (1995): Distribution of Neotropical Simuliidae (Insecta,
Diptera) and its areas of endemism. - Revista de la Academia Colombiana de Ciencias Exactas
Físicas y Naturales (Bogotá) 19 (75): 717-732.
COSCARÓN, S. & C.L. COSCARÓN ARIAS (1996a): On Simulium (Pternaspatha), with description of a new species
(Diptera - Simuliidae). - Memorias do Instituto Oswaldo Cruz (Rio de Janeiro) 91 (6): 703-710.
COSCARÓN, S. & C.L. COSCARÓN ARIAS (1996b): Análisis cladístico de Simulium (Pternaspatha)
ENDERLEIN con datos sobre su distribución geográfica (Diptera: Simuliidae). - Acta Ento-
mológica Chilena 20: 71-82.
COSCARÓN, S. & C.L. COSCARÓN ARIAS (1997): Cladistic analysis of the subgenera Inaequalium,
Psaroniocompsa and Chirostilbia of the genus Simulium with comments on their distribution
(Simuliidae: Diptera). - Revista de la Sociedad Entomológica Argentina 56 (1-4): 109-121.
COSCARÓN, S. & C.L. COSCARÓN ARIAS (2000a): New species records for the blackflies (Diptera -
Simuliidae) fauna of Argentina with description of adults, pupa and larva of Simulium oyapockense
s. l. and S. seriatum. - Memorias do Instituto Oswaldo Cruz (Rio de Janeiro) 95 (2): 179-187.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 651

COSCARÓN, S. & C.L. COSCARÓN ARIAS (2000b): Los simúlidos (Diptera - Insecta) del río Paraná en
el área próxima a la ciudad de Corrientes (Argentina): Dinámica Poblacional de los Estados
Inmaduros. - Facena 15: 25-37.
COSCARÓN, S. & C.L. COSCARÓN ARIAS (2002): Araucnephia iberaensis n. sp., a Neotropical black fly
with a peculiar distribution (Diptera - Simuliidae). - Memorias do Instituto Oswaldo Cruz
(Rio de Janeiro) 97 (1): 81-87.
COSCARÓN, S. & S. IBÁÑEZ BERNAL (1993): Sobre una nueva especie de Simuliidae del sur de México:
Simulium (Hemicnetha) biuxinisa n. sp. (Diptera). - Folia Entomológica Mexicana 88: 61-68.
COSCARÓN, S. & S. IBÁÑEZ BERNAL (1994): Sobre la ubicación taxonómica de Simulium ganalesense VARGAS,
MARTÍNEZ PALACIOS y DÍAZ NÁJERA (Diptera: Simuliidae). - Folia Entomológica Mexicana 90: 1-7.
COSCARÓN, S. & A. MATTA (1982): Notas sobre Simúlidos Neotropicales VIII. Sobre dos especies
nuevas de “jerjeles” del género Simulium Latreille del Norte de Chile (Simuliidae, Diptera,
Insecta). - Idesia 6: 49-61.
COSCARÓN, S. & D.R. MIRANDA ESQUIVEL (1998a): Pedrowygomyia a new neotropical Prosimuliini
genus (Diptera: Simuliidae): Gigantodax s. lat. split of into two genera. - Entomologica
Scandinavica 29 (2): 161-168.
COSCARÓN, S. & D.R. MIRANDA ESQUIVEL (1998b): New synonym of Simulium (Inaequalium) inaequale
(PATERSON Y SHANNON), 1927 (Diptera: Simuliidae). - Memorias do Instituto Oswaldo Cruz,
Rio de Janeiro 93 (3): 337.
COSCARÓN, S. & P. MUÑOZ DE HOYOS (1995): Blackfly novelties from the area near the “Paramo de
los Valles” in the Department of Tolima, Colombia (Diptera: Simuliidae). - Revista de la
Academia Colombiana de Ciencias Exactas, Físicas y Naturales 19 (74): 587-592.
COSCARÓN, S. & V. PY-DANIEL (1989): Tres especies nuevas de Simulium LATREILLE de la región
Neotropical: Simulium (Hemicnetha) cristalinun sp. n., (Grenieriella) wygodzinskyorum sp.n. y
sumapazense sp. n. (Diptera, Simuliidae). - Revista de Saúde Pública (São Paulo) 23 (4): 313-321.
COSCARÓN, S. & H. TAKAOKA (1989): Description of a new species of the black fly subgenus
Simulium (Ectemnaspis) from the Andes of Ecuador (Diptera, Simuliidae). - Japan Journal of
Tropical Medicine & Hygiene 17 (2): 191-196.
COSCARÓN, S. & P. WYGODZINSKY (1960): Sobre la presencia de la familia Simuliidae (Diptera,
Insecta) en la provincia de Buenos Aires. - Orientación Médica 432: 1124-1125.
COSCARÓN, S. & P. WYGODZINSKY (1962): Simuliidae (Diptera - Insecta) de Tierra del Fuego,
Patagonia e Isla de Juan Fernández. - Acta Zoológica Lilloana 18: 281-333.
COSCARÓN, S. & P. WYGODZINSKY (1972a): Notas sobre Simúlidos Neotropicales III. Sobre tres
especies de jejenes con hembras de color claro del S.E. del Paraguay y del N.E. de la Argentina
(Simuliidae, Diptera). - Revista del Museo de La Plata (N.S.) Secc. Zoología 11: 209-231.
COSCARÓN, S. & P. WYGODZINSKY (1972b): Taxonomy and distribution of the black fly genus
Simulium (Pternaspatha) ENDERLEIN (Simuliidae, Diptera, Insecta). - Bulletin of the American
Museum of Natural History 147: 199-240.
COSCARÓN, S. & P. WYGODZINSKY (1973a): Notas sobre Simúlidos Neotropicales II. Sobre Simulium
(Psaroniocompsa) opalinifrons (ENDERLEIN) y notas sobre el subgénero (Diptera, Insecta). - Physis
32: 161-172.
COSCARÓN, S. & P. WYGODZINSKY (1973b): Notas sobre simúlidos Neotropicales IV. Sobre la presencia
de Simulium inaequale (PATERSON Y SHANNON) en el N.E. de Argentina y S.E. del Paraguay y una
nueva especie muy afín. - Revista de la Sociedad Entomológica Argentina 34: 141-149.
652 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

COSCARÓN, S. & P. WYGODZINSKY (1975): Notas sobre simúlidos Neotropicales V. Aportes para el
conocimiento del subgénero Simulium (Notolepria) ENDERLEIN (Diptera - Simuliidae). - Revista
de la Sociedad Entomológica Argentina 34: 277-288.
COSCARÓN, S. & P. WYGODZINSKY (1984): Notas sobre simúlidos Neotropicales VII. Sobre los
subgéneros Simulium (Psaroniocompsa) ENDERLEIN y Simulium (Inaequalium) nov. - Arquivos de
Zoología (São Paulo) 31 (2): 37-103.
COSCARÓN, S., CERQUEIRA, A.L.P., SATO SCHUMAKER, T.T.V. & V.L.A. SALVIA (1992): Nuevos datos
sobre la distribución de simulidos de Brasil y descripción de Simulium (Coscaroniellum) cerradense
sp. n. (Diptera, Simuliidae). - Revista Brasilelira de Entomologia 36 (1): 111-119.
COSCARÓN, S., COSCARÓN ARIAS, C.L. & G. PORCARO (2000): Simuliidae (Diptera) del río Quequén
Grande en Las Cascadas, (Buenos Aires, Argentina): variación poblacional y relación con
temperatura, caudal y precipitaciones. - Acta Entomológica Chilena 24: 29-36.
COSCARÓN, S., IBÁÑEZ BERNAL, S. & C.L. COSCARÓN ARIAS (l996): Revisión de Simulium (Psilopelmia)
ENDERLEIN en la región Neotropical y análisis cladistico de sus especies (Diptera; Simuliidae).
- Acta Zoológica Mexicana (n.s.) 69: 37-l04.
COSCARÓN, S., IBÁÑEZ BERNAL, S. & C.L. COSCARÓN ARIAS (1999): Revision of Simulium (Simulium) in
the Neotropical realm (Insecta: Diptera: Simuliidae). - Memoirs of Entomology International
14: 543-604.
COSCARÓN, S., MIRANDA-ESQUIVEL, D.R., MOULTON, K., COSCARÓN ARIAS C. L. & S. IBÁÑEZ BERNAL
(2004): Simulium (Hearlea) VARGAS, MARTÍNEZ PALACIOS & DÍAZ NÁJERA (Diptera: Simuliidae):
Taxonomic revision and cladistic analysis. - Zootaxa 396: 1-52.
COSCARÓN, S., PY-DANIEL, V. & C.L. COSCARÓN ARIAS (1995): El subgénero Simulium (Psilopelmia) ENDERLEIN
en Sudamérica. - Boletim do Museu Paraense Emílio Goeldi, Série Zoologia 1993. 9 (2): 283-311.
COSCARÓN, S., SARANDÓN, R., COSCARÓN ARIAS, C.L. & E. DRAGO (1996): Análisis de factores
ambientales que influyen en la distribución de los Simuliidae (Diptera: Insecta) en el Cono
Austral de América del Sur. - Revista de la Academia Colombiana de Ciencias Exactas, Físicas
y Naturales 20 (78): 549-573.
COSCARÓN ARIAS, C.L. (1998a): Los simúlidos y su control en el Valle del Río Negro y Neuquén,
Argentina (Diptera: Culicoidea: Simuliidae). - Revista de la Sociedad Entomológica Argentina
58 (1-2): 276-281.
COSCARÓN ARIAS, C.L. (1998b): The polytene chromosomes of Cnesia dissimilis (EDWARDS) and three
species of Gigantodax ENDERLEIN (Diptera: Simuliidae) from Lanin National Park (Argentina).
- Memorias do Instituto Oswaldo Cruz (Rio de Janeiro) 93 (4): 445-458.
COSCARÓN ARIAS, C.L. (2001): Insecta, Diptera, Simuliidae. - In: FERNÁNDEZ, H.R. & E. DOMÍNGUEZ
(eds.): Guía para la determinación de los artrópodos bentónicos sudamericanos 195-219.
Serie: Investigaciones de la UNT. Subserie: Ciencias Exactas y Naturales. Vol. 1. Editorial
Universitaria de Tucumán, Argentina.
COSCARÓN ARIAS, C.L. (2003): Clave de Simuliidae (Insecta, Diptera) de Ecuador. - Folia
Entomológica Mexicana 43 (2): 191-219.
COSTA I. DA CUNHA, M., COSCARÓN, S. & R.M. AZEVEDO DE BASSI (1998): Determinación de los
estadios larvales de Simulium (Diptera, Simuliidae) de Paraná, Brasil. - Acta Biológica
Paranaense 27: 57-66.
CROSSKEY, R.W. (1969): A re-classification of the Simuliidae (Diptera) of Africa and its islands. -
Bulletin of the British Museum (Natural History), Entomology, Suppl. 14: 1-195.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 653

CROSSKEY, R.W. (1987): An annotated checklist of the world black flies (Diptera: Simuliidae). Part.
VIII: Black fly species of the world. - In: KIM, K.C. & R.W. MERRITT (eds.): Black flies:
ecology, population management and annoted word list: 425-520. Pennsylvania State
University Press, University Park: 528 pp.
CROSSKEY, R.W. (1990): The natural history of blackflies. - J. Wiley, Chichester, 711 pp.
CROSSKEY, R.W. & T.M. HOWARD (1997): A new taxonomic and geographical inventory of world
blackflies (Diptera: Simuliidae). - The Natural History Museum, London: 144 pp.
CROSSKEY, R.W. & T.M. HOWARD (2004): A revised taxonomical and geographical inventory of
world blackflies (Diptera - Nematocera). - Internetversion: http//www.nhm.ac.uk/entomolo-
gy/projects/blackflies/Inventory-pdf (accessed 27 May 2004).Introl 8. Inventory 78. The
Natural History Museum, London: 82 pp.
DALMAT, H.T. (1949): Some species of Simuliidae (Diptera) from Guatemala 1. - Annals of the
Entomological Society of America 42 (4): 538-553.
DALMAT, H.T. (1950): New species of Simuliidae (Diptera) from Guatemala 2. - Annals of the
Entomological Society of America 43 (1): 137-151.
DALMAT, H.T. (1951): Notes on the Simuliidae (Diptera) of Guatemala, including descriptions of
three new species. - Annals of the Entomological Society of America 44 (1): 31-58.
DALMAT, H.T. (1952): Description of two new species of Simulium (Diptera, Simuliidae) from
Guatemala. - Annals of the Entomological Society of America 45 (2): 339-347.
DALMAT, H.T. (1953): Simuliidae (Diptera) of Guatemala. Description of Simulium (Dyarella) ardeni
n. sp., and Simulium (Lanea) jacobsi n. sp. - Annals of the Entomological Society of America 46
(1): 35-42.
DALMAT, H.T. (1955): The blackflies (Diptera, Simuliidae) of Guatemala and their role as vectors of
onchocerciasis. - Smithsonian Miscellaneous Collections 125 (1): vii+ 425 pp.
DAMPF, A. (1927): Un Simulium nuevo de México (Orden Diptera, Suborden Nematocera)
procedente de Tiltepec, Estado de Oaxaca. - Revista Mexicana de Biologia 7 (6): 125-129.
D’ANDRETTA, C. (1954): Sobre Simulium distinctum LUTZ (1910) (Diptera, Simuliidae). - Anais da
Faculdade de Farmacia e Odontologia da Universidade de São Paulo 12: 63-66.
D’ANDRETTA, C. (1956): Redescrição de Simulium perflavum ROUBAUD, 1908. - Anais da Faculdade de
Farmacia e Odontologia da Universidade de São Paulo 13: 33-38 (1955).
D’ANDRETTA, C. & M.A.V. D’ANDRETTA (1947): As espécies Neotropicais da família Simuliidae
SCHINER (Diptera, Nematocera). II. Lutzsimulium cruzi n. gen. e n. sp. e nova concepção de
nervação das asas dos simulideos. - Memorias do Instituto Oswaldo Cruz (Rio de Janeiro) 44:
401-411, 35 figs.
D’ANDRETTA, C. & M.A.V. D’ANDRETTA (1948): Espécies Neotropicais da família Simuliidae
SCHINER (Diptera, Nematocera). IV. Lutzsimulium cruzi ANDR. & ANDR., 1946: Descrição do
alotipo macho e da larva. - Arquivos de Zoología (São Paulo) 5: 637-647.
D’ANDRETTA, C. & M.A.V. D’ANDRETTA (1950): Espécies Neotropicais da família Simuliidae
SCHINER (Diptera, Nematocera). IV. Redescrição de Simulium pertinax KOLLAR, 1832. - Papeis
Avulsos do Departamento de Zoología (São Paulo) 9: 193-213.
D’ANDRETTA, M.A.V. & C. D’ANDRETTA (1946): As espécies Neotropicais da família Simuliidae
SCHINER, 1864 (Diptera - Nematocera). Simulium (Eusimulium) orbitale LUTZ, 1910, Simulium
(Eusimulium) pintoi n. sp. e Simulium nigrimanum MACQUART, 1837, sp. inquirendae [sic]. -
Memorias do Instituto Oswaldo Cruz (Rio de Janeiro) 45: 85-l52.
654 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

D’ANDRETTA, M.A.V. & C.D. D’ANDRETTA (1947): Espécies Neotropicais da família Simuliidae
(Diptera, Nematocera). III. Simulium botulibranchium LUTZ, 1910 e travassosi n. sp. - Papeis
Avulsos do Departamento de Zoología (São Paulo) 8: 145-180.
D’ANDRETTA, M.A.V. & C. D’ANDRETTA (1949): Espécies Neotropicais da família “Simuliidae”
SCHINER (Diptera, Nematocera). Redescrição de “Simulium auristriatum LUTZ, 1910 e “Simulium
(Eusimulium) venustum infuscata” LUTZ, 1909 var. inquirendae [sic]. - Revista Brasileira de
Biología 9 (1): 55-66.
D’ANDRETTA, M.A.V. & C. D’ANDRETTA (1952): Espécies Neotropicais da família Simuliidae (Diptera,
Nematocera). VII. - Redescrição de Lutzsimulium pernigrum (LUTZ, 1910), n. comb. e consider-
ações sôbre o gênero. - Papeis Avulsos do Departamento de Zoología (São Paulo) 10: 307-324.
D’ANDRETTA, C. & D. GONZÁLEZ (1964): Tres novas espécies Brasileiras de Simuliidae (Diptera,
Nematocera). - Memorias do Instituto Butantan 30 (960-62): 103-116.
DE ALMEIDA, F.B. (1979): Simulium cerqueirellum, nomen novum para Simulium septentrionale CERQUEI-
RA & ALMEIDA, 1970 (Diptera: Simuliidae). - Acta Amazônica 9 (4): 789.
DE LEÓN, J.R. (1943): Preliminares para la descripción de cinco nuevas especies de la región
occidental de Guatemala. - Boletín Sanitario de Guatemala 51: 94-101.
DE LEÓN, J.R. (1945): Nuevas especies de simúlidos en la región occidental de Guatemala. - Boletín
Sanitario de Guatemala 52: 66-77 (1944).
DE LEÓN, J.R. (1948): Un nuevo simúlido de la región occidental de Guatemala. - Impreso no. 56,
Instituto de Investigaciones Científicas, Universidad de San Carlos (Guatemala): 1-23 pp.
DELLOME FILHO, J. (1983): Considerações sobre os fatores físico-químicos dos criadouros de
Simulium goeldii CERQUEIRA & MELLO, 1967 (Diptera: Nematocera). - Revista Brasileira de
Entomologia 27: 155-160.
DÍAZ NÁJERA, A. (1961): Nota acerca de Simulium (Psilopelmia) antillarum JENNINGS, 1915. Nueva
especie para México (Diptera, Simuliidae). - Revista del Instituto de Salubridad y Enfer-
medades Tropicales, México 21 (1-2): 79-92.
DÍAZ NÁJERA, A. (1962): Claves para identificar las larvas del género Cnephia y descripción de dos nuevas
especies. - Revista del Instituto de Salubridad y Enfermedades Tropicales, México 22: 271-287.
DÍAZ NÁJERA, A. (1965): Acerca de un nuevo simúlido mexicano (Diptera: Simuliidae). - Revista del
Instituto de Salubridad y Enfermedades Tropicales, México 15 (1): 25-45.
DÍAZ NÁJERA, A. (1969): Simulidos de Aguascalientes y Zacatecas. Descripción de seis especies del
subgénero Psilopelmia y datos de dimorfismo. - Revista de Investigaciones de Salud Pública,
México 29 (1): 21-63.
DÍAZ NÁJERA, A. (1971): Descripción de una nueva especie del género Cnephia del norte de México.
- Revista de Investigaciones de Salud Pública, México 31 (4): 239-247.
DÍAZ NÁJERA, A. (1979): Una nueva especie del género Simulium (Diptera: Simuliidae) del Estado de
Coahuila, México. - Anales del Instituto Biológico, Universidad Autónoma (Mexico), ser.
Zoologia 50 (1): 553-561.
DÍAZ NÁJERA, A. & M.A. VULCANO (1961): Descripción de Simulium (Psilopelmia) longithallum n. sp.
(Diptera - Simuliidae). - Revista del Instituto de Salubridad y Enfermedades Tropicales,
México 21 (3-4): 221-235.
DÍAZ NÁJERA, A. & M.A. VULCANO (1962): Claves para identificar las larvas de Simúlidos del
subgénero Hearlea, con descripción de dos nuevas especies. - Revista del Instituto de
Salubridad y Enfermedades Tropicales, México 22 (1-2): 91-144.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 655

DUMBLETON, L.J. (1960): Larval Simuliidae (Diptera) from Navarino Island, Tierra del Fuego. - New
Zealand Journal of Science 3: 543-548.
DYAR, H.G. & R.C. SHANNON (1927): The North American two-winged flies of the family
Simuliidae. - Proceedings of the United States National Museum 69 (10): 1-54.
EDMAN, J.D. & K.R. SIMMONS (1985a): Rearing and colonization of blackflies (Diptera: Simuliidae).
- Journal of Medical Entomology 22: 1-17.
EDMAN, J.D. & K.R. SIMMONS (1985b): Simuliids (mainly Simulium decorum WALKER): 145-152. - In:
SINGH, P. & R.F. MOORE (eds.): Handbook of insect rearing Vol. 2, Elsevier, Amsterdam: 514 pp.
EDWARDS, F.W. (1931): Simuliidae. - In: Diptera of Patagonia and South Chile. British Museum
(Natural History), London 2 (4): 121-154.
EDWARDS, F.W. (1933): Die Ausbeute der deutschen Chaco-Expedition.1925-26. XLIII. Simuliidae
(Melusinidae). - Konowia l2 (3-4): 210-211.
ELLIOT, M.J. & C. POTTER (1978): The future of pyrethroides in insect control. - Annual Review of
Entomology 23: 443-469.
ENDERLEIN, G. (1921a): Das System Kriebelmücken (Simuliidae). - Deutsche Tierärztliche Wochen-
schrift 29: 197-200.
ENDERLEIN, G. (1921b): Neue aussereuropäische Simuliden. - Sitzungsberichte der Gesellschaft
Naturforschender Freunde zu Berlin 4-5: 77-81.
ENDERLEIN, G. (1922): Weitere Beiträge zur Kenntnis der Simuliiden. - Konowia 1: 67-76.
ENDERLEIN, G. (1925): Weitere Beiträge zur Kenntnis der Simuliiden und Verbreitung. - Zoologi-
scher Anzeiger 62: 201-211.
ENDERLEIN, G. (1929): Über einige neotropische Simuliiden des Genus Friesia (Dipt.). - Deutsche
Entomologische Zeitschrift 4: 327-328.
ENDERLEIN, G. (1930): Der heutige Stand der Klassifikation der Simuliiden. - Archiv für
klassifikatorische und phylogenetische Entomologie 1: 77-97.
ENDERLEIN, G. (l934a): Aussereuropäische Simuliiden aus dem Wiener Museum. - Sitzungsberichte
der Gesellschaft Naturforschender Freunde (Berlin): 190-195.
ENDERLEIN, G. (1934b): Weiterer Ausbau des Systems der Simuliiden. - Deutsche Entomologische
Zeitschrift 2-3: 273-292 (l933).
ENDERLEIN, G. (l935): Neue Simuliiden, besonders aus Afrika. - Sitzungsberichte der Gesellschaft
Naturforschender Freunde (Berlin) 1934: 358-364.
ENDERLEIN, G. (1936): Simuliologica. - Sitzungsberichte der Gesellschaft Naturforschender Freun-
de (Berlin): 113-130.
FAIRCHILD, G.B. (1940): Notes on the Simuliidae of Panama (Diptera, Nematocera). - Annals of the
Entomological Society of America 33 (4): 701-719.
FAIRCHILD, G.B. & L. BRICEÑO IRAGORRY (1942): Nota sobre Simulium lutzianum. - Boletin del
Laboratorio de la Clínica «Luis Razetti» 2:158-159 (3:187-188).
FIELD, G. (1967a): Studies on black flies of Panama (Diptera: Simuliidae). I. Two new species of Simulium
of the subgenus Notolepria. - Annals of the Entomological Society of America 60 (1): 188-194.
FIELD, G. (1967b): Studies on black flies of Panama (Diptera: Simuliidae). II. A new species of
Simulium of the subgenus Hearlea. - Annals of the Entomological Society of America 60 (1):
l94-l99.
FIELD, G. (1969a): Studies of black flies of Panama. III. Two new species of Simulium of the
subgenus Hemicnetha. - Annals of the Entomological Society of America 62 (1): 157-163.
656 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

FIELD, G. (1969b): Studies of the black flies of Panama. IV. A new species of Simulium. - Annals of
the Entomological Society of America 62 (2): 281-284.
FLOCH, H. & E. ABONNENC (1946a): Simulides de la Guyane Française. I. S. guianense WISE, 1911, S.
rorotaense n. sp., S. maroniense n. sp. - Publication de l’Institut Pasteur de la Guyane et du
Territoire de l’Inini 136: 1-20.
FLOCH, H. & E. ABONNENC (1946b): Simulides de la Guyane Française. II. S. cauchense n. sp., S.
oyapockense n. sp., S. iracouboense n. sp. - Publication de l’Institut Pasteur de la Guyane et du
Territoire de l’lnini 137: 1-19.
FOX, I. (1953): Notes on Puerto Rican Simuliidae from light traps (Diptera). - Proceedings of the
Entomological Society of Washington 55 (3): 135-140.
FRIES, B.F. (1824): Monographia Simuliarum Sveciae: 5-20. - Observationes Entomologicae part I,
Lundae, Sweden: 20 pp.
GIL-ACEVEDO, L.H., FIGUEIRO, R. & M. MAIA-HERZOG (2005): Simulium (Psaroniocompsa) stellatum
(Diptera: Simuliidae), a new black fly from high mountain range in southeastern Brazil. -
Zootaxa 922: 1-12.
GIL-ACEVEDO, L.H., SANTOS MALLET, J.R. & M. MAIA-HERZOG (2004): Caracteres diagnosticos de
Simulium (Chirostilbia ) pertinax KOLLAR (Diptera: Simuliidae). - Neotropical Entomology 33
(4): 433-437.
GOELDI, E.A. (1905): Os mosquitos no Pará. - Memorias do Museu de Historia Natural
Ethnographico (Museu E. Goeldi) (Belém) 4: 1-154.
GRAY, E.W., ADLER, P.H. & R. NOBLET (1997): Economic impact of black flies (Diptera: Simuliidae)
in South Carolina and development of a localized suppression program. - Journal of the
American Mosquito Control Association 12: 676-678.
GRAY, E.W., ADLER, P.H., COSCARÓN ARIAS, C.L., COSCARÓN, S. & R. NOBLET (1999): Development of
the first black fly (Diptera: Simuliidae) management program in Argentina and comparison with
other programs. - Journal of the American Mosquito Control Association 15 (3): 400-406.
GRILLET, M.E. & R. BARRERA (1997): Spatial and temporal abundance, substrate partitioning and
species co-occurrence in a guild of Neotropical blackflies (Diptera: Simuliidae). - Hydrobiolo-
gia 345 (2-3): 197-208.
HAGEN, H.A. (1880): A new species of Simulium with a remarkable nympha case. - Proceedings of
the Boston Society of Natural History 20: 305-307.
HAMADA, N. (1998): Bionomics of Simulium perflavum ROUBAUD (Diptera: Simuliidae) in Central
Amazonia, Brazil. - Revista Brasileira de Entomologia 41 (2-4): 523-526
HAMADA, N. (2000): Simulium (Psaroniocompsa) tergospinosum new species (Diptera: Simuliidae) in siolii
group from southern part of the State of Amazonas, Brazil. - Memorias do Instituto
Oswaldo Cruz (Rio de Janeiro) 95 (6): 819-828.
HAMADA, N. & P.H. ADLER (1998a): A new species of Simulium (Diptera: Simuliidae) from open areas in
Central Amazonia, Brazil. - Memorias do Instituto Oswaldo Cruz (Rio de Janeiro) 93 (3): 3l7-325.
HAMADA, N. & P.H. ADLER (1998b): Taxonomy of the Simulium perflavum species-group (Diptera:
Simuliidae), with description of a new species from Brazil. - Insecta Mundi 12 (3-4): 207-226.
HAMADA, N. & P.H. ADLER (1999): Cytotaxonomy of four species in the Simulium perflavum species
group (Diptera: Simuliidae) from Brazilian Amazonia. - Systematic Entomology 24: 273-288.
HAMADA, N. & P.H. ADLER (2001): Bionomía e chave para imaturos e adultos de Simulium (Diptera:
Simuliidae) na Amazônia Central, Brasil. - Acta Amazônica 3l (1): 109-132.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 657

HAMADA, N. & C.A. AZEVEDO (2002): Descrição do macho e fêmea de Simulium lourencoi e lista de
espécies de Simuliidae (Diptera: Nematocera) de Carolina e municipios arredores, Maranhão,
Brasil. - Livro de Resumos do 19° Congresso Brasileiro de Entomología, Manaus, AM: 232-233.
HAMADA, N. & F. FOURQUE (2001): Black flies (Diptera: Simuliidae) of French Guiana: Cytotaxon-
omy and preliminary list of species. - Memorias do Instituto Oswaldo Cruz (Rio de Janeiro)
96 (7): 955-959.
HAMADA, N. & M.E. GRILLET (2001): Black flies (Diptera: Simuliidae) of the Gran Sabana
(Venezuela) and Pacaraima Region (Brazil): Distributional data and identification keys for
larvae and pupae. - Entomotropica l6 (1): 29-49.
HAMADA, N. & J.W. MCCREADIE (1999): Environmental factors associated with the distribution of
Simulium perflavum group (Diptera: Simuliidae) among streams in Brazilian Amazonia. -
Hydrobiologia 397: 71-78.
HAMADA, N. & M. PEPINELLI (2004): Simulium (Chirostilbia) bifenestratum (Diptera: Simuliidae), a new
black-fly species from the Atlantic Forest, State of São Paulo, Brazil. - Memorias Instituto
Oswaldo Cruz (Rio de Janeiro) 99 (1): 45-52.
HAMADA, N., ALE-ROCHA, R. & S.J. BESSA-LUZ (2003): Description of Simulium damascenoi (Diptera:
Simuliidae) male and the black-fly species from the state of Amapa, Brazil. - Memorias do
Instituto Oswaldo Cruz, Rio de Janeiro 98 (3): 353-360
HAMADA, N., BESSA-LUZ, S.J. & S. COSCARÓN (2004): New morphological information on Simulium
cerradense adults and new records of black fly species (Diptera: Simuliidae) in the western
region of Bahia state, Brazil. - Zootaxa 693: 1-8.
HEARLE, E. (1932): The blackflies of British Columbia (Simuliidae: Diptera). - Proceedings of the
Entomological Society of British Columbia 29: 5-19.
HERNÁNDEZ, L.M. & A.J. SHELLEY (2005): New specific synonymic and taxonomic notes on
Neotropical blackflies (Diptera: Simuliidae). - Zootaxa 853: l-46.
HERNÁNDEZ, L.M., SHELLEY, A.J., LUNA DIAS, A.P.A. & M. MAIA-HERZOG (2005): Description of the
female of Simulium itaunense D’ANDRETTA & GONZÁLEZ and redescription of the male and
pupa (Diptera: Simuliidae). - Zootaxa 896: 1-14.
HOFFMANN, C.C. (1930a): Un Simulium nuevo de la zona cafetera de Chiapas. - Anales del Instituto
de Biología, México 1: 51-53.
HOFFMANN, C.C. (1930b): Los simúlidos de la region onchocercosa de Chiapas (con descripción de
nuevas especies). - Anales del Instituto de Biologia, México 1 (4): 293-306.
HOFFMANN, C.C. (1931): Los simulidos de la region oncocercosica de Chiapas, segunda parte; los
estados larvales. - Anales del Instituto de Biología, México 2: 207-218.
IBÁÑEZ BERNAL, S. (l992): Las especies mexicanas de Simulium (Hemicnetha) y S. (Notolepria) (Diptera:
Simuliidae). - M.Sc.-thesis. Universidad Autónoma de México. México D.F: 302 pp.
IRIARTE, D.R. (1943): Contribución a la fauna Venezolana. - Boletin del Laboratorio de la Clínica
“Luis Razetti” 3: 189-203.
JENNINGS, A. (1915): Two new species of Simuliidae from tropical America. - Proceedings of the
Entomological Society of Washington 17 (4): 199-200.
JOAN, T. (1912): Nota sobre un díptero ponzoñoso. - Boletin del Ministerio de Agricultura, Buenos
Aires 14: 363-385.
KERTÉSZ, K. (1902a): Catalogus dipterorum hucusque descriptorum 1. - Leipzig: 339 pp.
KERTÉSZ, K. (1902b): Catalogus dipterorum hucusque descriptorum 2. - Leipzig: 359 pp.
658 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

KNAB, F. (1911): Dr. A. LUTZ‘s studies of Brazilian Simuliidae. - Proceedings of the Entomological
Society of Washington 13: 172-179.
KNAB, F. (1913): A note on some American Simuliidae. - Insecutor Inscitiae Menstruus 1 (12): 154-156.
KNAB. F. (1914a): Simuliidae de Chile septentrional. - Anales de Zoologia Aplicada, Chile 1 (7): 17-22.
KNAB. F. (1914b): Simuliidae of Peru. - Proceedings of the Biological Society of Washington 27: 81-86.
KNAB, F. (1914c): Supplementary notes on Peruvian Simuliidae. - Proceedings of the Biological
Society of Washington 27: 123-124.
KNAB, F. (1915a): A new Simulium from Texas (Diptera: Simuliidae). - Insecutor Inscitiae Menstruus
3 : 77-78.
KNAB, F. (1915b): New data and species in Simuliidae (Diptera). - Insecutor Inscitiae Menstruus 2
(12): 177-180 (19l4).
KNAB, F. (1915c): Some new Neotropical Simuliidae. - Bulletin of Entomological Research 6 (3): 279-282.
KOLLAR, V. (1832): Die vorzüglich lästigen Insecten Brasiliens. - In: POHL, J.E. (ed.): Reise im Innern
von Brasilien, Vol. 1: 101-119. Wien: 448pp.
LACEY, L.A. (1981): Simulideos antropofilicos no Parque Nacional da Amazônia (Tapajos), Brasil, com
referencia aos efeitos no homen. - Boletin de la Oficina Sanitaria Panamericana 90 (4): 326-338.
LACEY, L.A. & J.D. CHARLWOOD (1980): On the biting activities of some anthropophilic Amazonian
Simuliidae (Diptera). - Bulletin of the Entomological Research 70: 495-509.
LACEY, L.A. & A.H. UNDEEN (1988): The biological control potential of pathogens and parasite of
black flies. - In: KIM, K.C. & R.W. MERRITT (eds.): Black flies: ecology, population management
and annoted world list: 327-340 (1987). Pennsylvania State University, University Park: 528 pp.
LANE, J. & C.E. PORTO (1939): Simulideos da região neotropical: o gênero Eusimulium. - Boletim
Biológico (n. s.) (São Paulo) 4 (2): 168-176.
LANE, J. & C.E. PORTO (1940): Simulideos da região Neotrópica III. Descrição de novas especies dos
gêneros “Simulium “Eusimulium”. - Arquivos do Instituto Biológico (São Paulo) 11: 189-195.
LANE, J. & M.A. VULCANO (1943): A armadura bucal dos simuliideos e seu valor taxonômico
(Diptera: Simuliidae). - Revista de Entomologia (Rio de Janeiro) 14 (3): 430-440.
LATREILLE, P.A. (1802): Histoire naturelle, générale et particulière des Crustacés et des Insectes. 3.
Familles naturelles des genres. - F. Dufart, Paris: 467 pp.
LEÓN L.A. & P. WYGODZINSKY (1953): Los simúlidos de Ecuador, su importancia en medicina
humana. - Boletin de Informaciones Científicas Naciónales (Quito) 6 (57): 269-288.
LEWIS, D.J. (1963): Simuliidae (Diptera) from the human onchocerciasis area of Venezuela. -
Proceedings of the Royal Entomological Society of London 32: 53-62.
LEWIS, D.J. & R. IBÁÑEZ ALDECOA (1962): Simuliidae and their relation to human onchocerciasis in
northern Venezuela. - Bulletin of World Health Organization 27: 449-464.
LEWIS, D.J. & J.P. LEE POTTER (l964): Simuliidae (Diptera) from the Sierra Nevada de Santa Marta,
Colombia. - Annals and Magazine of Natural History, Ser. 13, 7: 95-100.
LOEW, H. (1862): Monographs of the Diptera of North America. Part I. - Smithsonian
Miscellaneous Collections 6 (1) xxiv + 221 pp.
LUNA DIAS, A.P.A., HERNÁNDEZ, L.M., MAIA-HERZOG, M. & A.J. SHELLEY (2004): Simulium lobatoi,
new species of blackfly, (Diptera: Simuliidae) from the States of Mato Grosso and Goias,
Central Brazil. - Memorias do Instituto Oswaldo Cruz (Rio de Janeiro) 99 (1): 37-43.
LUTZ, A. (1909): Contribuição para o conhecimento das espécies Brasileiras do gênero “Simulium”.
- Memorias do Instituto Oswaldo Cruz (Rio de Janeiro) 1: 124-146.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 659

LUTZ, A. (1910): Segunda contribução para o conhecimento das espécies Brasileiras do gênero
“Simulium”. - Memorias do Instituto Oswaldo Cruz (Rio de Janeiro) 2: 213-267.
LUTZ, A. (1917): Terceira contribução para o conhecimento das espécies Brasileiras do gênero
Simulium. O piúm do norte Simulium amazonicum . - Memorias do Instituto Oswaldo Cruz (Rio
de Janeiro) 9 (1): 63-67.
LUTZ, A. (1922): Zoología Medica. Nematoceros hematophagos não pertencentes aos culicídeos. -
A Folha Medica (Rio de Janeiro) 3: 89-92.
LUTZ, A. & A. MACHADO (1915): Viajem pelo São Francisco e por alguns dos seus afluentes entre
Pirapora e Joazeiro. - Memorias do Instituto Oswaldo Cruz (Rio de Janeiro) 7 (1-2): 5-49.
LUTZ, A. & M. NUÑEZ TOVAR (1928): Estudios de zoología e parasitología venezuelanas. - Rio de
Janeiro: 133 pp., 26 pls.
LUTZ, A., DE SOUZA ARAUJO, H.C. & O. DE FONSECA (1918): Viagem scientífico no Rio Paraná e à
Asunción com volta por Buenos Aires, Montevideo e Rio Grande. - Memorias do Instituto
Oswaldo Cruz (Rio de Janeiro) 10 (2): 104-173.
MACQUART, J. (1838): Diptères exotiques nouveaux ou peu connus.1er. partie. - Mémoires de la
Société Royale des Sciences, de l’Agriculture et des Arts de Lille 1838 (2): 9-225.
MACQUART, J. (1845): Diptères exotiques nouveaux ou peu connus. Supplèment. - Mémoires de la
Société Royal des Sciences, de l’Agricúlture et des Arts de Lille 1844: 133-364. (also published
separately with pagination 5-238, Paris).
MAIA-HERZOG, M., SHELLEY, A.J. & A.P.A. LUNA DIAS (1985): Simulium hirtipupa LUTZ, 1910
(Diptera: Simuliidae) - Descrição dos adultos e larva e rediscrição da pupa. - Memorias do
Instituto Oswaldo Cruz (Rio de Janeiro) 80 (4): 483-490.
MAIA-HERZOG, M., SHELLEY, A.J., LUNA DIAS, A.P.A. & R. MALAGUTI (1984): Comparação entre
Simulim brachycladum e Simulium rubrithorax, suas posiçoes no subgênero Hemicnetha e nota
sobre uma espécie próxima S. scutistriatum (Diptera: Simuliidae). - Memorias do Instituto
Oswaldo Cruz (Rio de Janeiro) 79: 342-356.
MALLOCH, J.R. (1912): One new genus and eight new species of dipterous insects in the United States
National Museum collection. - Proceedings of the United States National Museum 43: 649-658.
MALLOCH, J.R. (1913): A new species of Simulium from Texas. - Proceedings of the Entomological
Society of Washington 15: 133-134.
MALLOCH, J.R. (1914): American black flies or buffalo gnats. - United States Department of
Agriculture, Bureau of Entomology, Technical Series, Washington 26: 1-83.
MILLEST, A.L. (1990): Differences in the larval head patterns and body coloration of members of
Simulium metallicum complex (Diptera: Simuliide) from Mexico. - Bulletin of Entomological
Research 80: 191-l94.
MIRANDA-ESQUIVEL, D.R. (1999): Análisis filogenético de la tribu Simuliini (sensu CROSSKEY 1987)
para las regiones zoogeográficas Neotropical, Etiópica y Australiana. - Ph.D.-thesis, Facultad
de Ciencias Naturales y Museo, Universidad Nacional de La Plata, Argentina: 238 pp.
MIRANDA-ESQUIVEL, D.R. & S. COSCARÓN (2001): Cladistic analysis of Simulium (Trichodagmia) and
Simulium (Thyrsopelma) (Diptera: Simuliidae). - Zoological Journal of the Linnean Society 132
(4): 429-439.
MIRANDA-ESQUIVEL, D.R. & S. COSCARÓN (2004): Distributional patterns of Neotropical, Afrotropi-
cal and Australian-Oriental Simulium LATREILLE subgenera (Diptera: Simuliidae). - Cimbebasia
19: 165-174 (2003).
660 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

MORENO RAMÍREZ, C. (1997): Estudio morfológico y citologico integrado de tres especies del
Subgénero Simulium (Ectemnaspis) sensu COSCARÓN, 1997. - Magister-thesis in Biology (Sistema-
tics), Facultad de Ciencias, Universidad Nacional de Colombia, Bogotá, Colombia: 146 pp.
MOREIRA, G.R.P., PEGORARO, R.A. & G. SATO (1994): Influencia de fatores abioticos sobre o
desenvolvimento de Simulium nogueirai D’ANDRETTA e GONZÁLEZ en um corrego da mata
atlántica. - Anais da Sociedade Entomológica do Brasil 23 (3): 525-542.
MOULTON, J.K. (1996): A new species of Gigantodax ENDERLEIN (Diptera: Simuliidae) from the
United States. - Proceedings of the Entomological Society of Washington 98 (4): 741-751.
MOULTON, J.K. (1998): Reexamination of Simulium (Psilopelmia) ENDERLEIN (Diptera: Simuliidae) of
America North of Mexico. - Proceedings of the Entomological Society of Washington 100
(1): 50-71.
MUHAMMAD, A. (1988): A cytological description of Simulium virgatum COQUILLET and related
species. - M.Sc.-thesis, Sul Ross State University, Alpine, USA: 94 pp., 18 pls.
MUÑOZ DE HOYOS, P. & S. COSCARÓN (1999): Claves para la identificación de simulidos (Diptera:
Simuliidae) presentes entre las vertientes magdalenense y orinocense, en un sector al centro de
Colombia. - Revista de la Academia Colombiana de Ciencias Exactas, Físicas y Naturales 23
(Suplemento Especial): 181-214.
NATHAN, M.B., TIKASINGH, E.S. & P. MUNROE (1982): Filariasis in Amerindian of Western Guyana
with observations of Mansonella ozzardi by a Simulium species of the amazonicum group. -
Tropenmedizin und Parasitologie 33: 219-222.
NEIVA, A. & B. PENNA (1916): Viagem cientifica pelo Norte de Bahia, Sudoeste de Pernambuco, Sul
de Piauí e de Norte a Sul de Goiaz. - Memorias do Instituto Oswaldo Cruz (Rio de Janeiro) 8
(3): 1-224.
NUNES DE MELLO, J.A.S. (1974): Simulideos (Diptera, Nematocera) do Territorio Federal de
Roraima (Brasil). - Ph.D.-thesis, Doctoral Faculdade de Medicina, Universidade de Sorocaba,
São Paulo: VII+56 pp.
NUNES DE MELLO, J.A. & F. BARBOSA DE ALMEIDA (1974): Simuliidae da Amazônia VII. Descrição
de Simulium pseudoexiguum sp. n. (Diptera, Nematocera). - Acta Amazônica 4 (1): 65-68.
NUNES DE MELLO, J.A. & E. VIEIRA DA SILVA (1974): Simuliidae da Amazônia VIII. Descrição de
Simulium mutucuna sp. n. (Diptera, Nematocera). - Acta Amazônica 4 (2): 69-71.
NUNES DE MELLO, J.A., BARBOSA DE ALMEIDA, F. & J. DELLOME (1973): Simuliidae da Amazônia VI.
Descrição do Simulium meruoca sp. n. (Diptera, Nematocera). - Acta Amazônica 3 (3): 59-62.
OCHOA, J.O. & H. TAKAOKA (l993): A new blackfly species of Mayacnephia from Guatemala (Diptera:
Simuliidae). - Japanese Journal of Tropical Medicine and Hygiene 21 (1): 5-7.
OKAZAWA, T. & O. ONISHI (1980): Description of a new species of Simulium (Simulium) LATREILLE
and redescription of Simulium (Simulium) metallicum BELLARDI from Guatemala (Diptera:
Simuliidae). - Japanese Journal of Sanitary Zoology 32 (3): 167-179.
ORFILA, R.N. (1939): Simuliidae (Dipt.) de la República Argentina. - Novena Reunión de la
Sociedad Argentina de Patologia Regional 3: 1525-1534.
ORTIZ, I. (1957): Nuevos representantes hematófagos de los géneros Culicoides (Diptera: Cerato-
pogonidae) y Simulium (Diptera: Simuliidae) de Venezuela. - Boletin Venezolano de Laborato-
rio Clínico 2 (3-4): 161-168.
PATERSON, G. & R.C. SHANNON (1927): Los simulidos del noroeste argentino. - Revista del Instituto
Bacteriológico del Departamento Nacional de Higiene 4 (7): 737-742.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 661

PEPINELLI, M., TRIVINHO-STRIXINO, M. & N. HAMADA (2003): New records of Simuliidae (Diptera,
Nematocera) in the State of São Paulo, Brazil. - Revista Brasileira de Entomologia 47 (4): 653-655.
PERRUOLO LANETI, G. (1971): Descripción del macho y la hembra del Simulium matteabranchia ANDUZE,
1947 (Diptera, Simuliidae). - Kasmera Universidad de Zula, Maracaibo 3 (4): 397-403.
PETERSEN, J.L. (1985): Mayacnephia fortunensis (Diptera: Simuliidae), a new black fly species from
Panama. - Proceedings of the Entomological Society of Washington 87 (1): 80-84.
PETERSON, B.V (1981): Simuliidae. - In: MCALPINE, J.F, PETERSON, B.V., SHEWELL, G.E., TESKEY, H.J.,
VOCKEROTH, J.R. & D.M. WOOD (eds.): Manual of Nearctic Diptera I: 355-391. Biosystematic
Research Institute, Canadian Government Publication Center, Hull: 674 pp.
PETERSON, B.V. (1993): The black flies of the genus Simulium, subgenus Psilopelmia (Diptera: Simuliidae)
in the contiguous United States. - Journal of the New York Entomological Society 10 (3): 301-390.
PETERSON, B.V. & B.C. KONDRATIEFF (1995): The black flies (Diptera: Simuliidae) of Colorado: An
annoted list with keys, illustrations and descriptions of three new species. - Memoirs of the
American Entomological Society 42: 1-121 (1994).
PETERSON, B.V., VARGAS, M. & J. RAMÍREZ PÉREZ (1988): Simulium (Hemicnetha) hieroglypicum (Diptera:
Simuliidae), a new black fly species from Costa Rica. - Proceedings of the Entomological.
Society of Washington 90 (1): 76-86.
PHILIPPI, R.A. (1865): Aufzählung der chilenischen Dipteren. - Verhandlungen der Kaiserlich-
Königlichen Zoologisch-Botanischen Gesellschaft in Wien 15: 595-782.
PINTO, G. (1931): Simulidae da America Central e do Sul (Diptera). - Séptima Reunión de la
Sociedad Argentina de Patologia Regional del Norte: 661-763.
PINTO-SANCHEZ, N.R., MIRANDA-ESQUIVEL, D.R. & P. MUÑOZ DE HOYOS (2005): Phylogenetic
analysis of Gigantodax (Diptera: Simuliidae). - Insect Systematics and Evolution 36 (2): 1-25.
PORTO, C. (1939): Simulideos da região Neotropical II (Género Simulium). - Boletim Biológico (Nr.)
4 (3): 369-373.
POST, R.J., ADAMS, Z., SHELLEY, A.J., MAIA-HERZOG, M., LUNA DIAS, A.P.A. & S. COSCARÓN (2003):
The morphological discrimination of microfilariae of Onchocerca volvulus from Mansonella
ozzardi. - Parasitology 127: 21-27.
PY-DANIEL, V. (1981): Algumas considerações sôbre Simuliidae (Diptera: Nematocera) II. - Acta
Amazônica 11 (1): 171-181.
PY-DANIEL,V. (1982): Prosimuliini (Diptera: Simuliidae) Neotropical I. Kempfsimulium V. PY-DANIEL
& J.A. NUNES DE MELLO n. gen.; K. simplicicolor (LUTZ, 1910) n. comb.; Lutzsimulium
flavopubescens (LUTZ, 1910) n. comb.; Maycnephia muzquicensis (DÍAZ NÁJERA, 1971) n. comb. -
Amazoniana 7 (3): 293-333.
PY-DANIEL, V. (1983): Caracterização de dois novos subgéneros em Simuliidae (Diptera, Culicomor-
pha) Neotropicais. - Amazoniana 8 (2): 159-223.
PY-DANIEL, V. (1984): Simuliidae (Diptera. Culicomorpha) no Brasil II. Sobre o Simulium nahimi, sp.
n. - Boletim do Museu Paraense Emilio Goeldi, Série Zoologia 1 (1): 125-137.
PY-DANIEL, V. (1987): Simuliidae (Diptera, Culicomorpha) no Brasil V. Sôbre o Simulium (Chirostil-
bia) friedlanderi sp. n. e revisão do Simulium (C.) laneportoi VARGAS, 1941. - Revista de Saúde
Pública (São Paulo) 21 (4): 331-341.
PY-DANIEL, V. (1988): Simuliidae (Diptera, Culicomorpha) no Brasil VI. Sôbre Simulium (Psaronio-
compsa) siolii sp. n., Simulium (P.) lourençoi. sp. n., Simulium (P.) damascenoi sp. n. - Revista de Saúde
Pública (São Paulo) 22 (4): 292-310.
662 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

PY-DANIEL, V. (1989a): Novas sinonimias e correções em Simulium com a revalidação de S. pruinosum


LUTZ, 1904 (Culicomorpha, Simuliidae). - Revista de Saúde Pública (São Paulo) 23 (3): 254-257.
PY-DANIEL, V. (1989b): Simuliidae (Diptera, Culicomorpha) no Brasil X. Descrição de Simulium
(Psaroniocompsa) guaporense sp. n. - Revista de Saúde Pública (São Paulo) 23 (6): 502-508.
PY-DANIEL, V. & S. COSCARÓN (1982): Simuliidae (Diptera: Nematocera) no Brasil I. Sôbre o
Simulium rappae sp. n. - Revista Brasileira de Biologia 42 (1): 155-163.
PY-DANIEL, V. & S. COSCARÓN (2001): Simuliidae (Diptera: Culicomorpha) no Brasil III. Sôbre o
Coscaroniellum ulyssesi sp. n. - Entomologia e Vectores (Río de Janeiro) 8 (1): 51-64.
PY-DANIEL, V. & G.R.P. MOREIRA (1988): Simuliidan (Diptera, Culicomorpha) no Brasil VII. Sôbre
o Simulium (Chirostilbia) empascae sp. n. - Iheringia, Sér. Zoologia 67: 77-86.
PY-DANIEL,V. & G.R.P. MOREIRA (1989): Simuliidae (Diptera, Culicomorpha) no Brasil. IX. Simulium
(Inaequalium) nogueirai D’ANDRETTA & GONZÁLEZ, 1964. - Iheringia, Ser. Zoologia 69: 85-95.
PY-DANIEL,V. & R.T. MOREIRA SAMPAIO (1994): Jalacingomyia gen. n. (Culicomorpha); a resurreção de
Gymnopaidinae; a eliminação do nivel tribal; apresentação de novos caracteres e a redescrição
dos estagios larval e pupal de Simulium columbachensis (FABRICIUS, 1787) (Diptera: Simuliidae). -
Memorias del CAÍCET 4 (1-2): 101-148.
PY-DANIEL,V. & R.T. MOREIRA SAMPAIO (1995): Gêneros e espécies de Simuliidae (Diptera:
Culicomorpha) assinalados para o Brasil até 1995. - Entomologia e Vectores (Rio de Janeiro)
2 (50):117-121.
PY-DANIEL,V. & A.J. SHELLEY (1980): Revisão do Simulium spinibranchium LUTZ, 1910 (Diptera:
Simuliidae), com a primeira descrição dos adultos e larva, e redescrição da pupa. - Acta
Amazônica 10 (1): 213-223.
PY-DANIEL,V., KONRAD, H.G. & H.A.O. GASTAL (1985): Simuliidae (Diptera, Culicomorpha) no
Brasil. IV. Descrição das larvas de Simulium (Psaroniocompsa) brevifurcatum LUTZ, 1910; S. (P.)
anamariae VULCANO, 1962; S. (Thyrsopelma) itaunense D’ANDRETTA & DOLORES GONZÁLEZ, 1962.
- Iheringia, Sér. Zoologia 65: 95-102.
PY-DANIEL,V., SOUZA, M.A.T. & E.P. CALDAS (1988): Simuliidae (Diptera, Culicomorpha) no Brasil.
III. Sôbre o Simulium (Chirostilbia) riograndense sp. n. e revisão do Simulium (Chirostilbia)
distinctum LUTZ, 1910. - Iheringia, Sér. Zoologia 67: 37-57.
RAMBAJAN, I. (1979): A new species of (Simuliidae: Diptera) from the Lethen area, Rupununi
District, Guyana, South America. - Journal Guyana Museum and Zoo of the Royal
Agricultural and Commercial Society 45: 63-77.
RAMÍREZ PÉREZ, J. (1971): Distribución geográfica y revisión taxonómica de los simulidos (Diptera:
Nematocera) de Venezuela con descripción de diez especies nuevas. - Acta Biológica
Venezolana 7 (3): 271-372.
RAMÍREZ PÉREZ, J. (1980a): Descripción de dos nuevas especies del género Gigantodax ENDERLEIN
(Diptera, Simuliidae) del estado de Mérida (Venezuela). - Boletín de la Dirección de
Malariologia y Saneameto Ambiental 20 (1-4): 52-58.
RAMÍREZ PÉREZ, J. (1980b): Descripción de cuatro nuevas especies de Simulium LATREILLE, 1802:
(Diptera: Simuliidae) de la región amazónica de Venezuela. - Boletín de la Dirección de
Malariologia y Saneamiento Ambiental 20 (1-4): 59-69.
RAMÍREZ PÉREZ, J. (1983): Los jejenes de Venezuela. - Simposio de Oncocercosis Americana,
CAICIT, Puerto Ayacucho (ed. Mimeograf.): 156 pp.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 663

RAMÍREZ PÉREZ, J. & B.V. PETERSON (1981a): Estudio del complejo Simulium amazoncum- sanguineum
en Venezuela. Descripción de tres nuevas especies. - Boletín de la Dirección de Malariologia y
Saneamiento Ambiental 21: 151-160.
RAMÍREZ PÉREZ, J. & B.V. PETERSON (1981b): Simulium santaelenae (Diptera: Simuliidae) nueva especie
de jejen del Estado de Bolívar (Venezuela). - Boletín Dirección de Malariologia y Saneamiento
Ambiental 21: 161-164.
RAMÍREZ PÉREZ, J. & M.A. VULCANO (1973): Descripción y redescripciones de algunos simulidos de
Venezuela (Diptera: Simuliidae). - Archivo Venezolano de Medicina Tropical y Parasitologia
Médica 5 (2): 375-399.
RAMÍREZ PÉREZ, J., PETERSON, B.V. & M. VARGAS (1988): Mayacnephia salasi (Diptera: Simuliidae), a
new black fly species from Costa Rica. - Proceedings of the Entomological Society of
Washington 90 (1): 66-75.
RAMÍREZ PÉREZ, J., RASSI, E. & A. RAMÍREZ (1977): Cinco especies nuevas do Simulium LATREILLE,
1802 (Diptera, Simuliidae) de la región amazónica de Venezuela. - Boletín de la Dirección de
Malariologia y Saneamiento Ambiental, Venezuela 17 (2): 162-174.
RAMÍREZ PÉREZ, J., YARZABAL, L. & B.V. PETERSON (1982): La simuliofauna del Territorio Federal
Amazonas (Venezuela). - Publicacion Científica No. 1. Ediciones Proicet Amazonas: 104 pp.
RAMÍREZ PÉREZ, J., YARZABAL, L., TAKAOKA, H., TADA, I. & A. RAMÍREZ (1986): Simulium parimaensis
(Diptera: Simuliidae), nueva especie de jejen en el Territorio Federal Amazonas, Venezuela. -
Boletín de la Dirección de Malariologia y Saneamiento Ambiental 26 (1-4): 61-64.
RILEY, C.V. (1887): Report of the entomologist. - U.S. Department of Agriculture, Report l886: 459-592.
RÍOS-VELÁSQUEZ, C., HAMADA, N. & P. H. ADLER. 2002. Cytotaxonomy of Simulium goeldii CERQUEIRA
& NUNES DE MELLO and Simulium ulyssesi (PY-DANIEL & COSCARÓN) (Diptera: Simuliidae) in
Central Amazonia, Brazil. - Journal of Insect Systematics and Evolution 33: 113-120.
ROIG JUÑENT, S. & S. COSCARÓN (2001): Biogeographical history of the Neotropical and Neantarctic
Simuliidae (Diptera). - Revista del Museo Argentino de Ciencias. Naturales Bernardino
Rivadavia (Buenos Aires) 3 (2): 119-134.
ROMAÑA, C. & J.W. ABALOS (1948): Control larvario de simulidos y culicidos con Gammexane y
DDT. - Anales del Instituto de Medicina Regional, Tucuman 2 (2):107-146.
ROUBAUD, M.E. (1906a): Aperçus nouveaux, morphologiques et biologiques sur les diptères
piqueurs du groupe des Simulies. - Comptes Rendus Hebdomadaires de séances de
l’Académie Royal des Sciences (Paris) 143: 519-521.
ROUBAUD, M.E. (1906b): Simulies nouvelles de l’Amerique du Sud. - Bulletin du Museum d’Histoire
Naturelle 12: 106-110.
ROUBAUD, M.E. (1909): Description d’une simulie nouvelle du Pérou. - Bulletin de la Societé de
Pathologie exotique et de ses Filiales 2: 428-430.
RUBTZOV, I.A. (1940): Blackflies, Simuliidae. - Fauna of the USSR, New Series No. 23, Insects,
Diptera 6 (6), Akademii Nauk SSSR Moscow-Leningrad: 533 pp.
RUBTZOV, I.A. & I. GARCÍA ÁVILA (1972): Los simúlidos de Cuba (Diptera: Simuliidae). - Serie
Poeyana 96: 1-39.
SAWYER, J. (1991) : A comparison of taxonomic techniques in the identification of sibling species of
South American Simuliidae. - Ph.D.- thesis, University of Salford, UK: 216 pp.
SAY, T. (1823): Descriptions of dipterous insects of the United States.- Journal of the Academy of
Natural Sciences of Philadelphia 3: 9-54.
664 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

SCHROTTKY, C. (1909): Drei neue blutsaugende Dipteren aus Paraguay. - Zeitschrift für wissen-
schaftliche Insektenbiologie, Wien 5: 61-63.
SCOPOLI, J.A. (1780): Zwölfter Brief. Über eine Gattung schädlicher Insekten, die man die
kolombachischen Mücken nennet, welche unter den Ochsen, Schafen, Ziegen, Schweinen und
Pferden in Servien, dem Temeswarer Bannat, und der abendländischen Wallachen großen
Schaden anrichten. - In: GRISELINI, F.: Versuch einer politischen und natürlichen Geschichte
des Temeswarer Bannats in Briefen an Standespersonen und Gelehrte. Zweyter Theil: 123-
135. J.P. Krauss, Wien: 135 pp, 9 pls.
SHELLEY, A.J. (1982): Simulium amazonicum GOELDI, 1905 (Diptera, Simuliidae); proposed suppres-
sion of syntypes and designation of neotype. Z.N(S.) 2364. - Bulletin of Zoologic
Nomenclature 39 (1): 67-70.
SHELLEY, A.J. (1988): Biosystematic and medical importance of the Simulium amazonicum group and
S. exiguum complex in Latin America: 203-220. - In: SERVICE, M.W. (ed.): Biosystematics of
Haematophagous Insects. Systematics Association, Special volume 37. Oxford University
Press, Oxford: 363 pp.
SHELLEY, A.J. & M. ARZUBE (1985): Studies on the biology of Simuliidae (Diptera) at the Santiago
onchocerciasis focus in Ecuador, with special reference to the vectors and disease transmis-
sion. - Transactions of the Royal Society of Tropical Medicine and Hygiene 79: 328-338.
SHELLEY, A.J. & S. COSCARÓN (2001): Simuliid blackflies (Diptera: Simuliidae) and ceratopogonid
midges (Diptera: Ceratopogonidae) as vectors of Mansonella ozzardi (Nematoda: Onchocercidae)
in Northern Argentina. - Memorias do Instituto Oswaldo Cruz (Rio de Janeiro) 96 (4): 451-458.
SHELLEY, A.J. & A.P.A. LUNA DIAS (1980): Simulium argentiscutum sp. nov. (Diptera: Simaliidae) a
member of the S. amazonicum group of species: description of adults, pupa and larva. -
Memorias do Instituto Oswaldo Cruz (Rio de Janeiro) 75 (3-4): 105-111.
SHELLEY, A.J. & V. PY-DANIEL (1981): Simuliidae of Goias and the Federal District (Brasilia) I. A
description of Simulium (Chirostillbia) dekeyseri, new species. - Memorias do Instituto Oswaldo
Cruz (Rio de Janeiro) 76 (1): 23-32.
SHELLEY, A.J., ARZUBE, M. & C.A. COUCH (1989): Simuliidae (Diptera) of Santiago onchocerciasis
focus of Ecuador. - Bulletin of the British Museum (Natural History) (Entomology) 58: 79-130.
SHELLEY, A.J., HERNÁNDEZ, L.M. & J.B. DAVIES (2004): Blackflies (Diptera: Simulidae) of Southern
Guyana with keys for the identification of adults and pupae, a review. - Memorias do Instituto
Oswaldo Cruz (Rio de Janeiro) 99 (5): 443-470.
SHELLEY, A.J., HERNÁNDEZ, L.M. & M. PENN (2002): A biosystematic revision of the blackflies
(Diptera: Simuliidae) of Belize, Central America. - Bulletin of Natural History Museum,
London (Entomology) 72 (2): 135-271.
SHELLEY, A.J., LUNA DÍAS, A.P.A. & M. MAIA-HERZOG (1984): New specific synonymy in
Neotropical Simulium s. l. (Diptera: Simuliidae). - Memorias do Instituto Oswaldo Cruz (Rio
de Janeiro) 79 (2): 143-161.
SHELLEY, A.J., LUNA DIAS, A.P.A. & M.A.P. MORAES (1980): Simulium species of the amazonicum-
group as vectors of Mansonella ozzardi in the Brazilian Amazon. - Transactions of the Royal
Society of Tropical Medicine and Hygiene 74: 784-788.
SHELLEY, A.J., PINGER, R.R. & M.A.P. MORAES (1982): The taxonomy, biology and medical
importance of Simulium amazonicum GOELDI (Diptera: Simuliidae), with a review of related
species. - Bulletin of the British Museum (Natural History) (Entomology) 44 (1): 1-29.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 665

SHELLEY, A.J., LOWRY, C.A., MAIA-HERZOG, M., LUNA DIAS, A.P.A. & M.A.P. MORAES (1997):
Biosystematics studies on the Simuliidae (Diptera) of the Amazonia onchocerciasis focus. -
Bulletin of the Natural History Museum, London (Entomology) 66 (1): 1-121.
SHELLEY, A.J., LUNA DIAS, A.P.A., MAIA-HERZOG, M., PROCUNIER, W.S. & M.A.P. MORAES (1987):
Identification of vector species (Diptera: Simuliidae) of human onchocerciasis in the
Amazonia focus of Brazil and Venezuela. - Memorias do Instituto Oswaldo Cruz (Rio de
Janeiro) 82 (4): 461-465.
SHELLEY, A.J., LUNA DIAS, A.P.A., MAIA-HERZOG, M., LOWRY, C.A., GARRITANO, P.R., PENN, M. & M.
CAMARGO (2001): Simulium cuasiexiguum, a new blackfly species (Diptera: Simuliidae) from the
Minaçu area in the State of Goias, Central Brazil. - Memorias do Instituto Oswaldo Cruz (Rio
de Janeiro) 96 (4): 483-496.
SHELLEY, A.J., MAIA-HERZOG, M., LUNA DIAS, A.P.A. & C.A. COUCH (1989): Description of the
adults and pupa of Simulium (Trichodagmia) perplexum, new species (Diptera: Simuliidae). -
Memorias do Instituto Oswaldo Cruz (Rio de Janeiro) 84: 343-349
SHELLEY, A.J., MAIA-HERZOG, M., LOWRY, C.A., LUNA DIAS, A.P.A., GARRITANO, P., CAMARGO, M. & H.G.
CARTER (2000): The Simuliidae (Diptera) of the secondary onchocerciasis focus at Minaçu in Central
Brasil. - Bulletin of the Natural History Museum, London (Entomological Series) 69 (2): 171-221.
SHELLEY, A.J., PINGER, R.R., MORAES, M.A.P., CHARLWOOD, J.D. & J. HAYES (1979): Vectors of
Onchocerca volvulus at the river Toototobi, Brazil. - Journal of Helminthology 53: 41-43.
SILVA FIGUEROA, C. (1917): Dos nuevos simulidos de Chile. - Boletin del Museo Nacional de Chile 10: 28-35.
SMART, J. (1940): Simuliide (Diptera) from British Guiana and the Lesser Antilles. - Transactions of
the Royal Entomological Society of London 90: 1-11.
SMART, J. (1944): Notes on Simuliidae (Diptera) II. - Proceedings of the Royal Entomological
Society of London, Ser. B, 13: 131-136.
SMART, J. (1945): The classification of the Simuliidae (Diptera). - Transactions of the Royal
Entomological Society of London 95: 463-528.
SMITH, C.N. (1973): Pyrethrum for control of insects affecting man and animals. - In: CASIDA, J.E.
(ed.): Pyrethrum, the natural insecticide. Academy Press, London: 329 pp.
SPEISER, P. (1904): Zur Nomenclatur blutsaugender Dipteren Amerikas. - Insekten-Börse 21: 148
STAINS, G.S. & G.F. KNOWLTON (1943): A taxonomic and distributional study of Simuliidae of
western United States. - Annals of the Entomological Society of America 36 (2): 259-280.
STONE, A. (1948): Simulium virgatum COCQUILLET and a new related species (Diptera: Simuliidae). -
Journal of the Washington Academy of Sciences 38: 399-404.
STONE, A. (1962): Notes on the types of some Simuliidae (Diptera) described by ENDERLEIN. -
Annals of the Entomological Society of America 55: 206-209.
STONE, A. (1963): An annotated list of genus-group names in the family Simuliidae (Diptera). -
United States Department of Agriculture, Technical Bulletin 1284: 1-28.
STONE, A. (1969): The black flies of Dominica (Diptera: Simuliidae). - Proceedings of the
Entomological Society of Washington 61: 312-318.
STONE, A. & M.M. BOREHAM (1965): A new species of Simulium from the southwestern United
States (Diptera: Simuliidae). - Journal of Medical Entomology 2: 164-170.
STRIEDER, M.N. (1998): Revisão e analise cladistica de espécies de Inaequalium (COSCARÓN &
WYGODZINSKY, 1984): (Diptera: Simuliidae). - Ph.D.-thesis, Pontificia Universidade Catolica de
Rio Grande do Sul, Porto Alegre, Brasil: 160 pp.
666 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

STRIEDER, M.N. (2004): Espécies de simulídeos (Diptera, Nematocera, Simuliidae) no Rio Grande do
Sul, Brasil: distribuição geográfica. - Entomologia e Vectores (Rio de Janeiro) 11 (1): 113-143.
STRIEDER, M.N. & S. COSCARÓN (2000): El estado de Simulium (Psaroniocompsa) (Diptera, Simuliidae)
en la región Sur de Brasil y Argentina, con descripción de Simulium minuanum sp. n. -
Entomologia e Vectores (Rio de Janeiro) 7 (1): 103-117.
STRIEDER, M.N. & V. PY-DANIEL (1999): Especies de Inaequalium (Diptera, Simuliidae): dados
bionômicos e chaves para sua identificação. - Biociencias (Porto Alegre) 7 (2): 43-72.
STRIEDER, M.N. & V. PY-DANIEL (2000): Revisão de Inaequalium (Diptera, Simuliidae), com
redescrição das formas imaturas e descrição de uma nova espécie. - Entomologia e Vectores
(Rio de Janeiro) 7 (2): 91 pp.
STRIEDER, M.N. & V. PY-DANIEL (2002): A cladistic análisis of Inaequalium COSCARÓN & WYGODZIN-
SKY, 1984 with information on geographical distribution (Diptera: Simuliidae). - Memorias do
Instituto Oswaldo Cruz (Rio de Janeiro) 97 (1): 65-72.
STRIEDER, M.N., CORSEUIL, E. & V. PY-DANIEL (1992): Espécies do gênero Simulium (Diptera,
Simuliidae) ocorrentes no Rio Grande do Sul - Brasil, com chaves para sua identificação. -
Acta Biológica Leopoldense 14 (2): 53-74.
STUARDO ORTIZ, C. (1946): Catálogo de los dípteros de Chile. - Ministerio de Agricultura, Santiago:
250 pp.
SURCOUF, J.M. & R. GONZÁLEZ RINCONES (1911): Essai sur les diptères vulnérants du Venezuela. -
Diptères Nématocères vulnérants, Vol. 1, Malone, Paris: 320 pp.
TAKAOKA, H. (1982): Observations on the bionomics of larval and man-biting female populations
of Simulium horacioi, a new potencial vector of Onchocerca volvulus in Guatemala. - Japan Journal
of Tropical Medicine and Hygiene 10: 49-62.
TAKAOKA, H. & H. TAKAHASI (1982): A new species of black fly (Diptera: Simuliidae) from upland
areas of Guatemala. - Journal of Medical Entomology 19: 63-67.
TAKAOKA, H., HIRAI, H. & I. TADA (1988): New species of Gigantodax (Diptera: Simuliidae) from
the Andean area of Ecuador. - Journal of Medical Entomology 25 (6): 541-549.
TIDWELL, M.A., PETERSON, B.V., RAMÍREZ PÉREZ, J., TIDWELL, M.A. & L.A. LACEY (1980): Notas y claves
preliminaries de los jejenes neotropicales pertenecientes a los grupo Simulium amazonicum y S.
sanguineum (Diptera: Simuliidae) incluyendo los vectores de Onchocerca volvulus y Mansonella ozzardi.
- Boletin de la Dirección de Malariologia y Saneamiento Ambiental 21 (2): 79-89.
TIDWELL, M.A., TIDWELL, M.A., MUÑOZ DE HOYOS, P., CORREDOR, A. & P. BARRETO (1980): Vectores
de Onchocerca volvulus y Mansonella ozzardi en Colombia. - Colombia Médica 11 (4): 119-127.
TIDWELL, M.A., TIDWELL, M.A. & B.V. PETERSON (1981): A redescription of the female of Simulium
sanguineum KNAB and descriptions of the male, pupa, and larva (Diptera: Simuliidae). -
Proceedings of the Entomological Society of Washington 83 (1): 13-27.
TOWNSEND, C.H.T. (1897): Diptera from the Lower Rio Grande or Taumalipan region of Texas. -
Journal of the New York Entomological Society 5: 171-178.
TWINN, C.R. (1938): Blackflies from Utah and Idaho, with descriptions of new species (Simuliidae,
Diptera). - Canadian Entomologist 70: 48-55.
VARGAS, L. (1941a): Simulium lane-portoi n. sp. (Simuliidae, Dipt.) y lista de simúlidos mexicanos. -
Revista del Instituto de Salubridad y Enfermedades Tropicales, México 2 (1): 110-122.
VARGAS S, L. (1941b): Notas sobre la importancia sanitaria de los simúlidos y de su morfologia
interna. - Revista del Instituto de Salubridad y Enfermedades Tropicales, México 2: 213-236.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 667

VARGAS, L. (1942a): Notas sobre la terminalia de algunos simúlidos de México. S. (E.) paynei n.sp. VARGAS,
1942. - Revista del Instituto de Salubridad y Enfermedades Tropicales, México 3 (3): 229-249.
VARGAS, L. (1942b): Simulium fairchildi VARGAS, 1942 n. sp. de Panamá (Diptera, Simuliidae). - Revista
Médica Mexicana 22 (420): 458-459.
VARGAS, L. (1943a): Nombres y datos nuevos de simúlidos del Nuevo Mundo. - Revista de la
Sociedad Mexicana de Historia Natural 4 (3-4): 135-136.
VARGAS, L. (1943b): Nuevos datos sobre simulidos mexicanos. - Revista del Instituto de Salubridad
y Enfermedades Tropicales, México 4 (4): 359-370.
VARGAS, L. (1945a): Simúlidos del Nuevo Mundo. - Monografía del Instituto de Salubridad y
Enfermedades Tropicales, México, Vol. 1: vi+ 241 pp., 10 pls.
VARGAS, L. (1945b): Cuatro nuevas especies y otros datos sobre simúlidos de México.- Revista de la
Sociedad Mexicana de Historia Natural 6 (1-2): 71-82.
VARGAS, L. (1948): Notas sobre oncocercosis. VII. Infección experimental de Simulium (Lanea)
mangabeirai con Onchocerca volvulus. - Revista del Instituto de Salubridad y Enfermedades
Tropicales, México 9: 309-311.
VARGAS, L. & A. DÍAZ NÁJERA (1946): Simúlidos de México. Datos sobre sistemática y morfología.
Descripción de nuevos subgéneros y especies. - Revista del Instituto de Salubridad y
Enfermedades Tropicales, México 7 (3): 101-192.
VARGAS, L. & A. DÍAZ NÁJERA (l948a): Simulium clarki (FAIRCHILD, 1940) y Simulium lutzianum (PINTO,
1931): Estructuras genitales. - Revista del Instituto de Salubridad y Enfermedades Tropicales,
México 9 (1): 75-79.
VARGAS, L. & A. DÍAZ NÁJERA (1948b): Nuevas especies de simúlidos de México y consideraciones
diversas sobre especies ya descritas. - Revista del Instituto de Salubridad y Enfermedades
Tropicales, México 9 (4): 321-369.
VARGAS, L. & A. DÍAZ NÁJERA (1949): Claves para identificar las pupas de los simúlidos de México.
Descripción de Simulium (Dyarella) freemani n. sp., de Simulium (Neosimulium) encisoi n. sp. y
referencias adicionales sobre S. anduzei y S. ruizi. - Revista del Instituto de Salubridad y
Enfermedades Tropicales, México 10: 283-319.
VARGAS, L. & A. DÍAZ NÁJERA (1951): Notas sobre sistemática y morfologia de simúlidos. - Revista
de la Sociedad Mexicana de Historia Natural 12: 123-208.
VARGAS, L. & A. DÍAZ NÁJERA (1952): Descripción de la larva de Cnephia aguirrei (DALMAT, 1949). -
Revista de la Sociedad Mexicana de Historia Natural 12: 53-56.
VARGAS, L. & A. DÍAZ NÁJERA (1953a): Nota sobre simúlidos de Ecuador. - Revista Ecuatoriana de
Entomologia y Parasitologia 1 (4): 17-21.
VARGAS, L. & A. DÍAZ NÁJERA (1953b): Notas sobre el examen de los tipos de simúlidos descritos
por el Prof. G. ENDERLEIN. - Revista del Instituto de Salubridad y Enfermedades Tropicales,
México 13: 137-152.
VARGAS, L. & A. DÍAZ NÁJERA (1953c): Simulium (Notolepria) gonzalezi n. sp. (Insecta, Diptera). -
Revista del Instituto de Salubridad y Enfermedades Tropicales, México 13 (3): 235-239.
VARGAS, L. & A. DÍAZ NÁJERA (1954): Algunas consideraciones morfológicas y de nomenclatura
relativas a simúlidos americanos. - Revista del Instituto de Salubridad y Enfermedades
Tropicales, México 14: 57-72.
VARGAS, L. & A. DÍAZ NÁJERA (1956): Simulium (Hemicnetha) guerrerense n. sp. (Diptera: Simuliidae). -
Revista del Instituto de Salubridad y Enfermedades Tropicales, México 16 (3): 51-56.
668 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

VARGAS, L. & A. DÍAZ NÁJERA (1957a): Simúlidos de Durango y Simulium (Simulium) marquezi n. sp. (Diptera,
Simuliidae). - Revista del Instituto de Salubridad y Enfermedades Tropicales, México 17 (1): 33-41.
VARGAS, L. & A. DÍAZ NÁJERA (1957b): Simúlidos Mexicanos. - Revista del Instituto de Salubridad
y Enfermedades Tropicales, México 17 (4): 143-399.
VARGAS, L. & A. DÍAZ NÁJERA (1958): Simulium (Psilopelmia) bivittatum MALLOCH, 1914 (Diptera:
Simuliidae). - Revista del Instituto de Salubridad y Enfermedades Tropicales, México 18 (1): 13-30.
VARGAS, L., DÍAZ NÁJERA, A. & A. MARTÍNEZ PALACIOS (1943): Tres simúlidos nuevos para México.
- Revista del Instituto de Salubridad y Enfermedades Tropicales, México 4 (3): 287-291.
VARGAS, L., MARTÍNEZ PALACIOS, A. & A. DÍAZ NÁJERA (1946): Simúlidos de México. Datos sobre
sistemática y morfología. Descripción de nuevos subgéneros y especies. - Revista del Instituto
de Salubridad y Enfermedades Tropicales, México 7 (3): 95-192.
VARGAS, M. & J. RAMÍREZ PÉREZ (1988): Gigantodax bierigi & G. willei (Diptera: Simuliidae), two new
black fly species from Costa Rica. - Revista de Biología Tropical 36 (2B): 457-469.
VARGAS, M., VARGAS, J.V. & J. RAMÍREZ PÉREZ (1993): Simulium (Psilopelmia) travisi (Diptera:
Simuliidae), a new black fly species from Costa Rica. - Proceedings of the Entomological
Society of Washington 95 (2): 175-181.
VARGAS, M., TRAVIS, B.V., DÍAZ NÁJERA, A. & F. FALLAS (1977): Bionomics of black flies (Diptera:
Simuliidae) in Costa Rica. VII, genus Simulium, subgenus Hearlea. - Revista de Biología
Tropical 25 (1): 137-149.
VULCANO, M.A. (1958): Redescrição do Simulium rubrithorax LUTZ, 1909 e descrição do alotipo macho
(Diptera, Simuliidae). - Papeis Avulsos do Departamento de Zoología (São Paulo) 13: 227-240.
VULCANO, M.A. (1959): Descrição de Simulium obesum, sp. n. (Diptera, Simuliidae). - Papeis Avulsos
do Departamento de Zoología (São Paulo) 13: 351-360.
VULCANO, M.A. (1962): Simulideos da Serra de Bocaina. Observações bionômicas, revalidação de
Simulium brevifurcatum LUTZ e descrição de uma espécie nova, S. anamariae (Diptera, Simuli-
idae). - Papeis Avulsos do Departamento de Zoología (São Paulo) 15: 239-272.
VULCANO, M.A. (1967): Family Simuliidae. - A catalogue of the Diptera of the Americas South of
the United States. Departamento de Zoologia, Secretaria de Agricultura (São Paulo) 16: 1-43.
VULCANO, M.A. (1971): Simuliidae. - In: HURLBERT, S.H. (ed.): Biota Acuática de Sudamérica
Austral: 285-293. San Diego State University: 342 pp.
VULCANO, M.A. (1981): Simuliidae, Part I. - In: HURLBERT, S.H., RODRIGUEZ, G. & N. SANTOS (eds.)
Aquatic biota of tropical South America: 275-285. San Diego State University: 342 pp.
WALKER, F. (1861): Characters of undescribed Diptera in the collection of W.W. SAUNDERS. -
Transactions of the Entomological Society of London (n.s.) 5 (1858-1861): 297-334.
W.H.O. (1982): Division of vector biology and control onchocerciasis control programme in the
Volta river basin. A review of the literature concerning the feeding behaviour and general
ecology of Simulium. - Technical Report: 40 pp.
WILLISTON, S.W. (1893): List of Diptera of the Death Valley Expedition. - In: RILEY, C.V. (ed.): The
Death Valley Expedition. A Biological survey of parts of California, Nevada, Arizona and
Utah, Part II. 4. Report of a small collection of insects made during the Death Valley
Expedition. North America Fauna 7: 235-268.
WISE, K.S. (1911): The Simuliidae of British Guiana. - Timehri 1: 248-254.
WISE, K.S. (1912): The Itanimi fly. A new Simulium in British Guiana. - Journal of Tropical Medicine
and Hygiene 15: 43.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 669

WYGODZINSKY, P. (1949): Contribuciones al conocimiento de los Simuliidae Argentinos. I.


Introducción. Redescripción de Simulium lahillei PATERSON & SHANNON, 1927 (Diptera). -
Anales del Instituto de Medicina Regional, Tucumán 2: 303-319.
WYGODZINSKY, P. (1950): Contribución al conocimiento de los Simulidos Argentinos. III. Simulium
dinellii (JOAN, 1912) y S. wolffhuegeli (ENDERLEIN, 1920). - Anales del Instituto de Medicina
Regional, Tucumán 3 (1): 75-97.
WYGODZINSKY, P. (1951a): Sobre Simulium jujuyense PATERSON & SHANNON, 1927, Simulium exiguum
ROUBAUD, 1906 y Simulium opalinifrons (ENDERLEIN, 1934). - Anales del Instituto de Medicina
Regional, Universidad Nacional deTucumán 3 (2): 207-220.
WYGODZINSKY, P. (1951b): Sobre Simulium ignescens ROUBAUD, 1906, Simulium romanai sp. n. y Simulium
adolfolutzi sp. n. (Diptera, Simuliidae). - Anales del Instituto de Medicina Regional, Tucumán 3:
221-236.
WYGODZINSKY, P. (1953a): Sobre algunos simúlidos argentinos (Diptera). - Anales del Instituto de
Medicina Regional, Tucumán 3 (3): 293-320.
WYGODZINSKY, P. (1953b): Sobre algunos simúlidos de los países andinos (Diptera). - Anales del
Instituto de Medicina Regional, Tucumán. 3 (3): 321-337.
WYGODZINSKY, P. (1958): Notas y descripciones de Simuliidae patagónicos y andinos (Diptera). -
Acta Zoológica Lilloana 16: 121-148.
WYGODZINSKY, P. (1961): Sobre una nueva especie de Simulium de la provincia de Buenos Aires
(Simuliidae, Diptera). - Acta Kravsi, Cuadernos del Instituto Nacional de Microbiologia
(Buenos Aires) 2: 30-36.
WYGODZINSKY, P. (1971): Descriptions and redescriptions of species of the black-fly genus Simulium
from northern Andes of Colombia (Simuliidae-Diptera). - American Museum Novitates
2447: 1-38.
WYGODZINSKY, P. (1973): A black fly of the subgenus Simulium (Psaroniocompsa) (Simuliidae, Diptera)
from the High Andes of Colombia. - American Museum Novitates 2513: 1-11.
WYGODZINSKY, P. & S. COSCARÓN (1962): On the relationships and zoogeographical significance of
Austrosimulium anthracinum (BIGOT), a black fly from southern South America (Diptera,
Simuliidae). - Pacific Insects 44: 1-244.
WYGODZINSKY, P. & S. COSCARÓN (1967): A review of Simulium (Pternaspatha) ENDERLEIN (Simuliidae,
Diptera). - Bulletin of the Amertican Museum of Natural History 136: 42-116.
WYGODZINSKY, P. & S. COSCARÓN (1970): A new species-group of the black fly genus Simulium from
Western South America (Simuliidae - Diptera - Insecta). - American Museum Novitates 2433: 1-20.
WYGODZINSKY, P. & S. COSCARÓN (1973a): A review of the Mesoamerican and South America black
flies of the tribe Prosimuliini (Simuliinae - Simuliidae). - Bulletin of the American Museum of
Natural History 151: 133-199.
WYGODZINSKY, P. & S. COSCARÓN (1973b): Notes on South American black flies of the tribe
Prosimuliini (Simuliinae, Simuliidae, Diptera). - American Museum Novitates 2529: 1-9.
WYGODZINSKY, P. & S. COSCARÓN (1979): Description of a black fly of the subgenus Simulium
(Pternaspatha) from the high Andes of Ecuador (Simuliida, Diptera). - American Museum
Novitates 2670: 1-9.
WYGODZINSKY, P. & S. COSCARÓN (1982): Description of three unusual species of black fly subgenus
Simulium (Ectemnaspis) from the Andes of Colombia (Diptera, Simuliidae). - American
Museum Novitates 2736: 1-13.
670 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

WYGODZINSKY, P. & S. COSCARÓN (1989): A revision of the blackfly genus Gigantodax ENDERLEIN
(Simuliidae, Diptera, Insecta). - Bulletin of the American Museum of Natural History 189: 269 pp.
WYGODZINSKY, P. & A. DÍAZ NÁJERA (1970): Un nuevo género de simulido de la Republica Mexicana
(Diptera: Simuliidae). - Revista de Investigación en Salud Pública, México 30: 83-110.
ZETTERSTADT, J.W. (1838): Sectio tertia, Diptera. Dipterologis scandinaviae amicis et popularibus
carissimis: 478-868. - In: Insecta Lapponica. - L.Voss, Leipzig: 1140 pp. (1840).
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 671

8
SUBJECT INDEX *

Acropogon 139 Gigantodax 24, 33, 39, 41, 42, 44, 45, 47, 48,
barbatipes 191 50, 67, 90
edwardsi 159 adleri 91
Araucnephia 40, 41, 42, 44, 45, 47, 48, 49, 50, antarcticus 92
51, 66, 76, 94, 629 bierigi 91
iberaensis 67, 68, 69, 70, 71 bolivianus 90
montana 68, 69, 72, 73, 74 brophyi 92
montanum 66, 72, 108 clandestinum 29
Araucnephioides 40, 41, 42, 44, 45, 47, 48, 49, conviti 91
50, 51, 67, 76, 94, 629 dryadicaudicis 92
schlingeri 76, 77, 79, 80 femineum 22, 90
Archicnesia 90 herreri 32
Astega 96 igniculus 30, 92
Austrosimulium 99 lazoi 91
Cnephia 56, 57, 59, 60, 61, 72 marginalis 30, 92
Cnephia (Stegopterna) 84 minor 92
Cnesia 24, 40, 41, 42, 45, 47, 48, 50, 51, 81, rufidulus 92
94, 629 willei 91
dissimile 81 Jalacingomyia 471
dissimilis 83, 84, 85, 86, 88, 90 Kempfsimulium 40, 41, 42, 45, 47, 48, 50, 51,
gynandra 84 110, 111, 629
gynandrum 81,84 hebeticolor 111
ornate 83, 84, 88, 89, 90 simplicicolor 110, 111, 112, 113
pusilla 83, 84, 85, 88 Lutzsimulium 39, 40, 41, 35, 45, 47, 48, 50,
Cnesiamima 39, 41, 42, 43, 44, 45, 48, 49, 50, 51, 102, 110, 111, 629
51, 81, 93, 94, 97, 629 cruzi 101, 106
atroparva 95, 96, 98 flavopubescens 104, 105, 109
atroparvum 93, 96 hirticosta 104, 105, 106, 107, 109
Dasypelmoza 139, 216 montanum 108
varipes 216 pernigrum 103, 104, 105, 108, 109
Eusimulium 109 Mayacnephia 40, 41, 42, 44, 45, 47, 48, 49, 50,
Friesia 108, 216 51, 52, 54, 63, 67, 629

*
Figures in bold
672 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

alticola 65 bustosi 473, 474, 476, 477, 478, 480, 482,


aguirrei 53, 55, 56, 58 483, 484, 496
atzopensis 55, 56, 58 costalimai 473, 474, 476, 477, 480, 481,
fortunensis 55, 56, 58 482, 483, 484, 485, 488, 489, 497, 499,
grenieri 55, 56, 58, 59 502
mixensis 55, 56, 58, 59 covagarciai 472, 474, 475, 477, 478, 479,
muzquicensis 55, 56, 58, 60 481, 482, 485, 503, 511
pachecolunai 52, 55, 56, 58, 60, 62 guatemalense 489
roblesi 53, 55, 56, 58, 61 hechti 472, 474, 476, 477, 478, 479, 482,
salasi 55, 56, 58 483, 486, 499, 502, 506, 507, 517
tadai 55, 56, 58 hirtipes 488
Odagmia 626 horacioi 473, 474, 476, 477, 478, 480, 482,
Oestrus 114 483, 487, 490, 498
columbacensis 114 hunteri 471, 473, 474, 476, 477, 479, 480,
Paraustrosimulium 39, 41, 42, 44, 45, 47, 49, 481, 483, 485, 487, 489, 496, 502, 507
50, 51, 97, 99, 629 iriartei 473, 474, 476, 477, 479, 480, 481,
anthracinum 22, 94, 99, 100, 101 483, 488, 500, 502, 507, 515
moorei 101 jacumbae 473, 474, 476, 477, 479, 480, 482,
Pedrowygomyia 24, 33, 39, 41, 42, 44, 45, 47, 485, 489, 496, 502, 504, 507
49, 50, 51 jobbinsi 473, 474, 476, 477, 479, 482, 483,
chacabamba 93 490, 498
cortesi 91, 92, 93 knowltoni 503
jatunchuspi 92, 93 kompi 472, 474, 475, 477, 478, 479, 480,
punapi 92, 93 482, 491, 514
Prosimulium 87 lassmanni 487
Simulium 39, 43, 45, 49, 51, 114 marquezi 473, 474, 475, 477, 478, 480, 481,
costaricense 626 482, 492, 499, 500, 502, 506, 510, 512,
diaznajerai 626 515, 516
nigrifemur 626 matteabranchia 493
spinifer 237 matteabranchium 473, 475, 477, 479, 480,
tenuifrons 626 481, 482, 486, 493, 494, 496, 497, 500,
Simulium (Acropogon) 626 502, 503, 506, 510, 511
Simulium (Anasolen) 518, 612 metallicum 366, 473, 474, 476, 477, 478,
Simulium (Aspathia) 116, 117, 120, 122, 123, 480, 482, 483, 487, 490, 491, 494, 495,
124, 126, 127, 128, 131, 133, 134, 135, 496, 497, 499, 500, 501, 502, 506, 507,
471, 643 508, 509, 511, 515, 516
A-K (cytoforms) 495, 498 morae 473, 474, 476, 477, 478, 480, 482, 501
H (cytoform) 487 nitidum 495
L (cytoform) 495 parnassum 517
X (cytoform) 495 parrai 473, 474, 476, 477, 479, 480, 481,
anduzei 471, 473, 474, 476, 477, 478, 480, 483, 495, 496, 499, 501, 502, 510, 511
482, 483 patziciaense 483
avidum 495 piperi 473, 474, 476, 477, 479, 482, 485,
490, 494, 496, 499, 502, 503
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 673

puigi 473, 474, 476, 477, 478, 479, 482, cuneatum 327, 328, 329, 330, 331, 332, 333,
483, 499, 502, 504, 507 334, 335, 337, 339, 340, 342, 343, 344,
putre 472, 475, 476, 477, 478, 480, 481, 345, 346, 347, 348
482, 494, 496, 497, 499, 500, 502, 505, delponteianum 326, 327, 328, 330, 331, 332,
506, 507, 515, 516 333, 334, 337, 347, 348
racenisi 473, 474, 476, 477, 478, 479, 482, exiguum 348
498, 501, 508, 509 ganalesense 327, 328, 329, 330, 331, 332,
riveti 495 333, 334, 335, 339, 340
ruizi 472, 475, 477, 480, 481, 483, 502, lutzi 348
509, 510 minuscula 332
sandyi 472, 474, 475, 477, 478, 479, 481, minusculum 326, 327, 328, 329, 330, 331, 332,
482, 496, 510, 512 333, 334, 335, 337, 339, 340, 345, 348
sayi 486, 503 minutum 348
sp. G 508 oyapockense 327, 328, 329, 330, 331, 332,
spilmani 473, 474, 476, 477, 478, 480, 481, 333, 335, 337, 340, 341
483, 484, 500, 502, 507, 512 pseudoamazonicum 336
stonei 503 pseudosanguineum 335
tescorum 484 quadristrigata 348
tricorne 474, 478, 480, 481, 492, 506 quadristrigatum 348
tricornis 513 roraimense 326, 327, 328, 329, 330, 33l,
vandalicum 492 332, 334, 335, 339, 340
versicolor 495 sanchezi 335
violacescens 495 sanguineum 327, 328, 329, 330, 331, 332,
wygoi 472, 474, 476, 477, 478, 480, 482, 333, 334, 337, 339, 340, 341, 343
483, 487, 496, 497, 500, 502, 506, 507, tallaferroae 341
514, 515, 516 venezuelense 326, 327, 328, 330, 331, 339,
Simulium (Astega) 96 345
Simulium (Byssodon) 117, 120, 123, 125, 126, Simulium (Chirostilbia) 115, 118, 119, 120, 121,
127, 133, 134, 136, 138, 365 122, 123, 124, 125, 126, 128, 130, 131,
forbesi 138 132, 133, 134, 135, 136, 633
meridionale 138 acarayense 240, 241, 242, 243, 244, 245, 246,
tamaulipense 138 259, 260, 261, 262, 263, 264, 266
Simulium (Cerqueirellum) 117, 120, 123, 124, bifenestratum 239, 240, 241, 242, 243, 244,
131, 133, 134, 135, 296, 325, 335, 360, 245, 246, 257
364, 366, 458, 636 cerqueirai 247
amazonicum 325, 326, 327, 328, 329, 330, dekeyseri 249
331, 332, 333, 339, 343, 344, 346 distincta 255
argentiscutum 326, 327, 328, 329, 331, 332, distinctum 239, 240, 241, 242, 243, 244,
333, 334, 337, 346, 348, 361 245, 246, 248, 250, 252, 254, 255, 256,
chaquense 327, 328, 329, 330, 331, 332, 333, 257, 259, 265
334, 337, 339, 340, 341, 342, 343, 344, empascae 240, 242, 243, 244, 246, 250, 252, 254
345, 348 flavifemur 238, 247
cuasisanguineum 335 friedlanderi 239, 240, 241, 242, 243, 244,
cuneata 344 245, 246, 252, 257
674 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

guarani 259 mallochi 365


inexorabile 247 metallicum 365
infuscatum 247 quadrifidum 349, 350, 351, 352, 353, 354,
laneportoi 239, 240, 241, 242, 243, 244, 355, 357, 358, 359, 360, 361, 362, 363,
245, 246, 248, 249, 250, 251, 253, 254, 365
256, 266 quadrivittatum 326, 349, 350, 351, 352, 353,
lutziana 247 354, 355, 357, 358, 364, 365, 449
obesa 258 rangeli 361
obesum 239, 240, 241, 242, 243, 244, 245, rasii 354
246, 248, 250, 252, 257, 258 scorzai 362
papaveroi 240, 241, 242, 243, 244, 245, 246, sextobecium 361
259, 260, 261, 262, 263, 264, 265 sp. A 360
pertinax 239, 240, 241, 242, 243, 244, 245, sp. B1 363
246, 247, 248, 249, 250, 251, 252, 253, torrealbai 354
255, 265, 266 ulyssesi 349, 350, 351, 352, 353, 354, 358,
pilosum 249 359, 361, 363, 364
prumirimense 255 Simulium (Cnephia) 81
riograndense 239, 240, 241, 242, 243, 244, gynandrum 81
245, 246, 250, 252, 256, 259 Simulium (Dasypelmoza) 400, 401, 413
septentrionale 247 Simulium (Dyarella) 517, 531, 537, 539, 546,
serranus 239, 240, 241, 242, 243, 244, 245, 552, 553, 554, 555, 581
246, 248, 249, 250, 251, 252, 265 Simulium (Ectemnaspis) 118, 120, 121, 122,
spadicidorsum 255 123, 125, 129, 131, 132, 133, 134, 136,
spinibranchia 253 294, 366, 420, 461, 462, 463, 518, 626,
spinibranchium 239, 240, 241, 242, 243, 638, 639, 640, 641
244, 245, 246, 248, 249, 250, 251, 253, adolfolutzi 403
254, 255, 265 albanense 28, 367, 371, 374, 375, 377, 379,
striginotum 266 382, 384, 386, 389, 391, 393, 395, 399,
subpallidum 28, 239, 240, 241, 242, 243, 244, 403, 406, 415
245, 246, 259, 260, 262, 263, 264, 265 alirioi 404
subviride 259 anaimense 367, 370, 375, 378, 382, 385,
venustum 247 395, 396, 399, 402
Simulium (Coscaroniellum) 117, 118, 120, 122, antillarum 368, 370, 372, 374, 375, 378,
123, 124, 128, 131, 133, 134, 135, 296, 380, 383, 384, 386, 393, 417, 418, 419
326, 349, 364, 637 antonii 367, 368, 370, 372, 375, 377, 378,
benjamini 365, 449 381, 383, 388, 392, 397
cauchense 349, 350, 351, 352, 353, 354, 355, antunesi 409
356, 361 arcabucense 367, 368, 371, 375, 377, 380,
cerradense 349, 350, 351, 352, 353, 355, 382, 384, 386, 390, 391, 392, 393, 394,
356, 357, 358, 359, 360, 361 395, 396, 399, 405
daltanhani 349, 350, 351, 352, 353, 357, 363 bicolorata 385, 400
fairchildi 365 bicoloratum 367, 371, 374, 375, 377, 379,
goeldii 349, 350, 351, 352, 353, 354, 355, 382, 384, 385, 386, 389, 394, 395, 400,
357, 358, 359, 362, 363, 364 401, 402, 403, 404, 405
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 675

bicornutum 367, 368, 371, 375, 377, 379, ochraceum 368, 370, 372, 374, 375, 378,
381, 384, 388, 391, 395, 397, 399, 400 380, 383, 384, 386, 389, 391, 393, 412,
bipunctatum 416 413, 414, 416, 418
cormonsi 367, 375, 377, 379, 382, 383, 401, pautense 367, 368, 371, 375, 377, 380, 382,
402 384, 386, 389, 390, 391, 392, 393, 394,
dinellii 368, 370, 372, 374, 375, 378, 380, 395, 396, 399, 402, 406
383, 384, 386, 389, 393, 395, 414, 416, perflava 409
419 perflavum 367, 372, 375, 378, 380, 382, 384,
flavipictum 414, 415 386, 389, 391, 393, 396, 399, 409, 414
fulvinotum 411 pifanoi 367, 368, 371, 374, 375, 377, 380,
furcillatum 367, 368, 371, 375, 377, 379, 381, 382, 384, 388, 389, 390, 391, 394, 395,
384, 388, 389, 391, 394, 395, 397, 398 396, 399
gabaldoni 367, 371, 375, 377, 380, 382, 383, pseudoantillarum 418
386, 390, 393, 405, 415 romanai 25, 367, 371, 374, 375, 377, 379,
gaudeatum 400 382, 385, 386, 391, 393, 395, 396, 399,
haarupiana 407 400, 403, 405, 406, 408
ignacioi 411, 412 roquemayu 367, 370, 374, 375, 377, 378,
ignescens 367, 368, 371, 375, 377, 379, 381, 383, 387, 388, 389, 390, 396
384, 385, 386, 387, 394 rorotaense 367, 370, 372, 375, 378, 380,
iguazuense 404 381, 383, 386, 389, 391, 393, 396, 399,
jaimeramirezi 367, 371, 375, 377, 379, 382, 410, 411, 412, 413
383, 400, 402 rubiginosum 367, 368, 371, 374, 375, 377, 379,
kabanayense 367, 370, 372, 375, 378, 380, 383, 384, 386, 387, 391, 393, 395, 396
381, 383, 386, 389, 391, 393, 394, 396, scutellatum 416, 417
410 shewellianum 368, 370, 372, 374, 375, 378,
kavanayense 410 380, 383, 385, 386, 389, 391, 393, 395,
lewisi 404 396, 399, 412, 415, 417, 418
lutziana 404 simplex 397
lutzianum 367, 371, 374, 375, 377, 379, sp. A 390
382, 385, 386, 389, 395, 396, 400, 403, sp. B 390
404, 406, 415, 416 suarezi 367, 370, 372, 375, 378, 380, 381,
macca 400, 401 383, 386, 389, 395, 396, 410, 411, 412
maroniense 367, 370, 372, 375, 378, 380, 381, tolimaense 367, 368, 370, 374, 377, 378,
383, 386, 395, 396, 410, 411, 412, 413 381, 383, 395, 397
martinezi 414 trombetense 367, 370, 372, 375, 378, 380,
mauense 409 381, 383, 396, 413
mayuchuspi 25, 367, 370, 374, 375, 377, tunja 367, 368, 371, 375, 377, 380, 381,
378, 383, 385, 386, 389, 391, 393, 395, 385, 389, 391, 393, 394, 395, 399
399, 408 varians 419
miniata 414 wolcotti 416, 417
molli 400 wolffhuegeli 367, 371, 374, 375, 377, 379,
nilesi 409 383, 384, 386, 389, 391, 393, 395, 396,
nuneztovari 368, 370, 372, 374, 375, 378, 399, 407
380, 383, 418 wuayaraka 411, 412
676 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Simulium (Eusimulium) 115, 119, 123, 124, microbranchium 563, 564, 566, 567, 568,
130, 133, 136, 137, 266, 461, 531, 533, 570, 571, 575, 577, 578, 585, 586
597, 616 nigricorne 562, 564, 565, 568, 569, 577, 586,
aureum 137 595
diazi 137 paracarolinae 562, 563, 567, 569, 590, 593,
donovani 137 595, 596
Simulium (Freemaniellum) 518, 612 temascalense 562, 563, 565, 568, 570, 590,
Simulium (Grenieriella) 597, 603, 605, 607, 609 591, 594, 596
Simulium (Hearlea) 33, 117, 121, 122, 123, 127, Simulium (Hemicnetha) 116, 117, 119, 120, 121,
131, 132, 134, 518, 560, 589, 645 122, 124, 127, 129, 131, 133, 135, 517,
ayrozai 562, 564, 566, 568, 570, 571, 577, 549, 573, 607, 612, 644
583, 586 acatenangoensis 531
burchi 562, 564, 566, 567, 569, 570, 572, ardeni 555
580, 581 aureopunctatum 548
canadense 562, 564, 566, 567, 568, 570, 573, bilimekae 531
577, 578, 581 biuxinisa 519, 521, 522, 524, 526, 528, 530,
capricorne 562, 564, 566, 567, 568, 570, 532, 533, 535, 536, 538, 540, 542
572, 574, 575, 576, 577, 578, 579, 581, brachycladum 519, 520, 521, 523, 524, 526, 527,
582, 583, 584, 585, 593, 596 528, 529, 532, 534, 536, 538, 540, 542,
carolinae 562, 563, 565, 567, 568, 569, 586, 543, 544, 545, 547, 553, 554, 555, 556
590, 592, 595 brevibranchium 540
chiriquiense 562, 564, 568, 578, 584 bricenoi 519, 521, 522, 524, 526, 527, 528,
contrerense 562, 564, 565, 567, 568, 570, 530, 533, 535, 536, 537, 538, 539, 549,
579, 585 551, 552, 555, 556, 560
dalmati 563, 564, 566, 567, 569, 570, 572, chiapanense 537
574, 578, 579, 580, 583 cinereum 537
delatorrei 562, 564, 566, 567, 569, 570, 572, clavipes 560
575, 576, 580, 581 coffeae 560
deleonii 574 conviti 531
estevezi 562, 564, 566, 569, 570, 578, 580, cristalinum 519, 520, 522, 523, 524, 526,
582 527, 528, 530, 532, 534, 536, 538, 541,
ethelae 563, 564, 566, 567, 568, 570, 577, 542, 544, 545, 546, 547, 556
578, 581, 583, 584, 586 dehnei 553, 554
gorirossiae 563, 565, 567, 568, 570, 587, earlei 519, 521, 522, 523, 524, 526, 528,
588, 591, 592, 593, 594, 596 529, 534, 536, 538, 553, 554, 556, 560
gorirossii 587 falculatum 560
johnsoni 562, 564, 565, 567, 568, 569, 588, freemani 519, 520, 522, 523, 524, 525, 527,
590, 591, 592, 597 528, 529, 549, 552
juarezi 562, 564, 566, 567, 568, 570, 575, guerrerense 519, 521, 522, 523, 524, 526,
576, 580, 584 527, 529, 544, 546, 547, 553, 555
larvispinosum 563, 564, 565, 567, 568, 569, hieroglyphicum 519, 520, 522, 523, 524, 526,
575, 576, 581, 587, 588, 589, 590, 591, 527, 529, 530, 538, 540, 551, 552
593, 594, 595, 596, 597 hinmani 462, 519, 521, 522, 524, 526, 527,
menchacai 563, 565, 567, 569, 588, 590, 594 530, 554, 557
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 677

hippovorum 537, 539 turgidum 548


keenani 519, 520, 521, 523, 524, 526, 529, virgatum 519, 521, 522, 525, 526, 527,
553, 554 528, 530, 531, 534, 535, 536, 537, 538,
laticalx 549 547, 560
latidigitus 549 virgatum chiapanense 537
lobatoi 519, 521, 523, 524, 526, 527, 528, yepocapense 519, 521, 522, 523, 524, 526,
529, 545 528, 529, 530, 532, 553, 554, 555
lugubre 548 Simulium (Inaequalium) 115, 120, 122, 123,
magnum 533 125, 126, 130, 132, 134, 136, 266, 294,
major 533 634
mathesoni 531 argentata 273
mexicanum 517, 518, 519, 520, 522, 523, baiensis 273
525, 527, 529, 530, 531, 532, 536, 538, beaupertuyi 267, 281, 282
547, 549, 550, 551, 552, 560 botulibranchium 267, 268, 269, 270, 271,
mutucuna 531 273, 288, 289, 290, 291, 292
oviedoi 518, 519, 520, 521, 523, 524, 525, clarki 267, 268, 269, 271, 272, 275, 282,
526, 528, 530, 556, 557, 558, 559 283
paynei 517, 519, 521, 522, 524, 526, 527, clavibranchium 267, 268, 269, 270, 271, 272,
528, 530, 531, 532, 535, 536, 537, 539, 274, 276, 277, 278, 279, 281, 283, 284,
540, 560 285, 287, 288
pictipes 560 diversibranchium 267, 268, 269, 270, 271,
placidum 548 272, 274, 276, 277, 279, 280, 281, 285,
pulverulentum 519, 521, 522, 523, 524, 526, 286, 287, 288, 289
527, 528, 529, 532, 534, 536, 541, 544, diversifurcatum 278
546, 547, 554, 555, 556 hauseri 267, 268, 269, 271, 272, 275, 281,
ribasi 557 283
rivai 557 inaequale 267, 268, 269, 271, 272, 273, 274,
rivasi 518, 519, 520, 521, 523, 524, 525, 275, 276, 277, 278, 279, 280, 281, 282,
526, 528, 530, 557, 558, 559 283, 284, 285, 287, 291
rubicundulum 537 inaequalis 273
rubrithorax 519, 521, 522, 524, 526, 527, jundiaiense 273
528, 530, 531, 532, 533, 534, 536, 537, leopoldense 267, 268, 271, 272, 279, 281,
538, 539, 540, 541, 542, 545, 601 282
seriatum 519, 520, 522, 523, 525, 527, 529, luribayae 294
530, 532, 534, 536, 538, 542, 549, 551, manicata 273
552, 553, 555, 556 mariavulcanoae 267, 268, 269, 270, 271,
smarti 519, 520, 522, 523, 524, 525, 527, 272, 276, 277, 279, 286
529, 530, 532, 534, 536, 538, 549, 550, mbarigui 278
551, 552 missionum 287
solarii 519, 521, 523, 525, 526, 527, 529, nahimi 267, 268, 270, 271, 273, 276, 279,
530, 546, 553, 554 285
tarsale 560 nogueirai 267, 268, 269, 270, 271, 272, 275,
tarsatum 549 279, 281, 282
tephrodes 537 parimaensis 294
678 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

petropoliense 267, 268, 270, 271, 273, 290, urubambanum 228


292, 293 Simulium (Odagmia) 400, 409, 626
pseudoexiguum 294 Simulium (Pliodasina) 324
rappae 267, 268, 269, 271, 276, 277, 280, 287 guttata 313, 324
souzalopesi 267, 268, 269, 270, 271, 273, Simulium (Psaroniocompsa) 117, 120, 122, 123, 125,
290, 291, 292, 294 131, 133, 134, 136, 325, 335, 364, 635
subclavibranchium 267, 268, 269, 271, 272, aequifurcatum 314, 315
274, 276, 277, 278, 279, 283, 284, 289 anamariae 296, 297, 298, 299, 301, 304,
subnigrum 23, 267, 268, 269, 270, 271, 272, 308, 309, 313, 315, 316, 317, 318, 319
274, 275, 276, 278, 279, 280, 281, 282, angrense 296, 297, 298, 300, 301, 305, 308,
283, 284, 285, 289 309, 310
travassosi 267, 268, 269, 270, 271, 273, 274, angustitarse 306
276, 277, 280, 281, 288 auripellita 302
tulibranchium 289 auripellitum 296, 297, 298, 299, 300, 301,
Simulium (Lanea) 419, 437, 441, 444, 448, 451, 302, 303, 304, 305, 307, 308, 309, 310,
454, 456, 457, 458, 461 311, 315, 320
Simulium (Morops) 407 auristriata 316
Simulium (Neosimulium) 138 auristriatum 296, 297, 299, 300, 303, 304,
Simulium (Nevermania) 115, 119, 123, 124, 305, 308, 3l2, 313, 316, 318
130, 133, 136, 137 bonaerense 296, 297, 298, 300, 301, 302,
annulipes 137 303, 304, 305, 307, 308, 309, 310, 311,
carbunculum 138 312, 313, 314, 315, 318, 319
pugetense 138 brevifurcata 317
ruficorne 137 brevifurcatum 296, 297, 298, 299, 301, 303,
Simulium (Notolepria) 116, 120, 123, 126, 130, 308, 313, 316, 317, 319
134, 136, 317, 319, 360, 361, 632 catarinense 324
blantoni 237 damascenoi 296, 297, 298, 300, 303, 305,
cuasiexiguum 225, 226, 227, 228, 233, 234, 308, 309, 320, 322, 323
313 fuliginis 296, 299, 315
delpontei 228 guaporense 296, 298, 301, 320, 322, 323
exigua 228 guttata 324
exiguum 29, 224, 226, 227, 228, 230, 231, guttatum 324
232, 233, 234, 313 hoffmanni 306
glaucophthalmum 228 incaicum 306
gonzalezi 225, 226, 227, 228, 229, 230, 232, incrustata 311
233, 234, 237 incrustatum 296, 297, 298, 299, 301, 302,
incertum 237 303, 305, 307, 308, 309, 310, 311, 312,
llutense 225, 226, 227, 228, 235, 236 314, 315, 317
paraguayense 226, 227, 228, 229, 230, 231, infuscatum 316
232, 233, 234, 237, 313 jujuyense 296, 297, 298, 300, 301, 302, 305,
paranense 237 306, 307, 308, 309, 310
sp. A 226, 227, 228, 229, 233, 234, 235 limbata 306
subexiguum 225, 226, 227, 228, 229, 233, limbatum 296, 297, 298, 299, 301, 302, 303,
234, 237 304, 305, 308, 309, 313, 314, 315, 317
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 679

lourencoi 296, 298, 301, 320, 321, 322 escomeli 419, 422, 424, 428, 430, 432, 435,
machadoallisoni 314 436, 438, 440, 443, 444, 446, 447, 448,
machadoi 314 450, 451, 452, 453, 454, 455, 457, 458
mendozana 306 gonzalezherrejoni 422, 424, 426, 428, 430,
meruoca 314 433, 434, 436, 440, 443, 446, 450, 456,
minuana 305 457, 458, 459
minuanum 296, 297, 298, 299, 300, 301, haematopotum 366, 419, 421, 424, 426, 428,
303, 305, 308, 310 430, 432, 434, 436, 438, 439, 440, 441,
opalinifrons 311 446, 447, 448, 470
schmidtmummi 296, 297, 298, 299, 300, 304, idahoense 442
305, 308, 309, 318, 319, 321, 322 iracouboense 422, 423, 426, 428, 430, 431,
sicuani 306 434, 436, 438, 440, 443, 445, 446, 447,
siolii 296, 297, 298, 300, 301, 308, 312, 452
320, 321, 322, 323, 324 jacobsi 470
spinifer 306 jerezense 459
stellatum 296, 297, 298, 299, 300, 319, 322 letrasense 459
tergospinosum 296, 297, 298, 300, 304, 305, longithallum 422, 424, 426, 427, 430, 433,
308, 320, 321, 322 434, 436, 450, 456, 457, 458, 459
yarzabali 311, 313 mangabeirai 422, 424, 427, 430, 433, 435, 438,
Simulium (Psilopelmia) 116, 118, 120, 121, 122, 443, 446, 447, 450, 452, 456, 457, 458
123, 124, 125, 127, 129, 131, 132, 133, marathrumi 448, 449
134, 135, 136, 360, 361, 365, 419, 642 mazzottii 455
bivittatum 422, 424, 428, 430, 432, 435, mediovittatum 421, 423, 428, 429, 432, 435,
437, 438, 440, 442, 443, 446, 447, 448, 436, 439, 440, 441, 442, 447, 449, 452
449, 450 mooseri 461
blancasi 420, 422, 425, 427, 428, 431, 433, netteli 459
436, 464, 465, 467, 469, 470 ochoai 421, 424, 426, 427, 429, 432, 435,
bobpetersoni 422, 424, 426, 428, 430, 432, 436, 439, 441, 442, 447, 449, 450, 452
435, 437, 440, 446, 447, 450, 456, 457, olympicus 459
458, 459 panamense 420, 421, 423, 425, 428, 429,
boydi 448 431, 433, 436, 438, 440, 446, 447, 450,
callidum 420, 421, 423, 425, 428, 429, 431, 452, 460
437, 438, 440, 443, 446, 450, 452, 453, penai 421, 423, 425, 427, 429, 431, 433,
461, 462 435, 464, 467, 469, 470
colvini 444 pseudocallidum 420, 421, 423, 425, 428, 429,
dandrettai 422, 424, 425, 427, 430, 432, 431, 434, 437, 438, 450, 452, 461, 462
433, 436, 438, 440, 441, 446, 447, 450, pseudohaematopotum 448
453, 454, 455 rufidorsum 419, 449
distinctum 455 samboni 422, 423, 426, 428, 430, 431, 434,
downsi 422, 423, 426, 428, 430, 431, 435, 437, 438, 440, 443, 444, 445, 447, 449,
437, 440, 446, 447, 450, 451, 459 452, 461, 470
dugesi 421, 424, 426, 427, 429, 432, 435, santaelenae 445
436, 437, 438, 439, 440, 441, 442, 443, sp. near dandrettai 425, 434, 447, 450
446, 447, 449, 450, 452 sucamense 445
680 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

tenuipes 421, 423, 425, 427, 429, 431, 433, figueroa 173
435, 464, 467, 468, 470 hectorvargasi 142, 144, 146, 149, 151, 154, 155,
travisi 462 158, 206, 207, 208, 209, 210, 212, 213
trivittatum 422, 424, 425, 428, 430, 433, herreri 143, 144, 145, 147, 148, 152, 154,
434, 437, 438, 440, 441, 446, 449, 450, 155, 156, 158, 183, 196, 197, 200, 204,
452, 455, 456, 457, 458 206, 208, 210, 212, 214
veracruzanum 422, 424, 426, 428, 430, 432, horcochuspi 141, 143, 146, 149, 150, 152,
434, 437, 440, 443, 446, 447, 450, 456, 156, 159, 172, 173, 181
457, 458, 462 huemul 163
vulcanoae 459 illiesi 173
zempoalense 422, 424, 425, 427, 430, 432, limay 141, 144, 146, 148, 150, 153, 155,
434, 436, 443, 447, 450, 452, 454, 457 158, 163, 164, 168, 169
Simulium (Psilozia) 117, 120, 123, 127, 131, luchoi 142, 144, 145, 147, 151, 154, 155,
134, 136, 138 157, 212
argus 139 nemorale 142, 143, 144, 145, 148, 151, 152,
encisoi 139 153, 155, 158, 166, 168, 169, 171, 172,
groenlandica 138 174, 176, 179, 188, 189, 192, 194, 208
kamloopsi 139 nigristrigatum 141, 144, 146, 148, 150, 153,
vittatum 139 155, 157, 159, 161, 165, 196
Simulium (Pternaspatha) 115, 116, 119, 123, philippii 142, 144, 147, 148, 152, 153, 157,
125, 126, 127, 130, 134, 136, 630, 631 158, 194, 195
albicinctum 142, 144, 147, 151, 154, 157, pichi 143, 145, 146, 149, 152, 154, 156,
202, 210, 211, 213 158, 165, 167, 171, 181, 183, 184
albicinctus 210 prodexargernteum 141, 142, 143, 144, 145, 148,
albilineatum 143, 145, 147, 149, 152, 154, 150, 151, 153, 154, 155, 157, 213, 215
157, 159, 199, 202, 204, 205, 206 pulchrum 142, 145, 146, 148, 152, 154, 155,
annulatum 141, 143, 147, 149, 150, 153, 157, 174, 176, 177
157, 159, 216, 217, 235 punctativentris 173, 174
bachmanni 160, 163 quechuanum 143, 144, 147, 149, 152, 154,
barbatipes 142, 144, 147, 149, 152, 153, 156, 158, 206, 207, 208, 213
156, 159, 174, 191, 193, 207 schoenemanni 140, 143, 145, 147, 150, 152,
bordai 140, 143, 147, 150, 152, 157, 205 154, 157, 221, 224
caprii 141, 144, 146, 149, 151, 153, 155, simile 142, 145, 148, 152, 154, 155, 158,
159, 218, 220, 224 173, 175, 176, 177
cotopaxi 142, 143, 147, 149, 151, 153, 157, stelliferum 142, 144, 147, 149, 151, 153,
159, 185, 187 156, 158, 159, 194, 221, 223, 224
deagostinii 141, 143, 144, 146, 148, 150, 152, strigidorsum 29, 141, 143, 145, 147, 150,
153, 156, 158, 171, 172, 181, 183, 184 152, 154, 157, 181, 182
diamantinum 141, 144, 146, 148, 150, 153, varipes 139, 216
156, 158, 168, 171, 178, 180, 181, 183 walterwittmeri 163, 165
dureti 143, 145, 146, 147, 149, 152, 154, yacuchuspi 143, 145, 147, 148, 152, 154,
156, 159, 166, 168, 170, 172, 181, 183, 156, 158, 199, 201, 204, 207, 210, 211
184, 214 Simulium (Thyrsopelma) 116, 118, 121, 123,
edwardsi 160 126, 130, 131, 132, 134, 518, 612, 647
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 681

A, B, C, D (cytoforms) 620 chalcocoma 601


albimanum 616 ecuadoriense 601
albopicta 626 huairayacu 598, 599, 600, 601, 602, 603,
albopictum 626 604, 605, 606, 607, 608, 609
brasiliense 612, 616 lahillei 30, 597, 598, 599, 600, 602, 603,
guianense 613, 614, 615, 616, 617, 619, 621, 604, 605, 606, 607, 608, 610
622, 623, 624 latitarse 597, 601
hirtipupa 613, 614, 615, 616, 618, 619, 620, muiscorum 598, 599, 600, 601, 603, 604,
622, 623 608
itaunense 613, 614, 615, 616, 620 nigrimanum 598, 599, 600, 601, 602, 604,
itaunensis 614, 620 606, 609, 610
nigrimanum 616 pruinosa 609
orbitale 612, 613, 614, 615, 616, 618, 619, pruinosum 609, 611
620, 621, 622, 623, 624 spadicidorsum 609
ortizi 620 strigata 601
paraguayense 616 sumapazense 598, 599, 600, 601, 603, 605,
perplexa 624 606, 609
perplexum 613, 614, 615, 616, 624 townsendi 597, 598, 599, 600, 601, 602, 607
pinto 620, 624 wygodzinskyorum 598, 599, 600, 601, 604,
scutistriata 625 605, 606, 608
scutistriatum 613, 614, 615, 616, 618, 619, Simulium (Wilhelmia) 228, 365
621, 623, 625 Simulium (Xenosimulium) 518, 612
Simulium (Trichodagmiia) 116, 118, 119, 121, Tlalocomyia 24, 39, 41, 42, 44, 45, 48, 49, 51,
122, 124, 126, 129, 130, 132, 133, 135, 54, 63, 67, 629
518, 597, 612, 616, 617, 622, 624, 625, alticola 54, 65
626, 646 revelata 63, 64, 65
682 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

9
ADDENDUM

Simulium (Chirostilbia) jefersoni HAMADA, HERNÁNDEZ, LUZ & PEPINELLI


Simulium (Chirostilbia) jefersoni HAMADA et al., 2006: 23-35.
Diagnosis based on original description.
Female: Wing length 1.3-1.6 mm. Scutum black with recumbent whitish setae,
showing 1+1 submedian and 1+1 sublateral silver stripes, changing to blackish with
different light position. Pleura dark brown. Legs yellowish with apical portion dark brown;
Sc with hair, basal section of R bare. Claw with subbasal tooth. Paraproct subtriangular,
about 1.5 times longer than cercus. Gonapophysis subovoidal, elongated, lightly sclero-
tized on internal margin.
Male: Wing length 1.6-2.5 mm. Scutum black with golden hairs, showing with some
light 1+1 submedian silver pruinose spots on anterior one third; scutellum and metano-
tum dark brown. Ventral plate very short with ventral margin mostly straight and well
sclerotized. Gonostylus finger-like, slightly shorter than gonocoxite, with longitudinal
ridge and 2 small apical spiniform setae.
Pupa: Cocoon shoe shaped with loop-like fenestrations protecting gill branches;
length ventrally 3.8-5.4 mm, and dorsally 2.8-3.8 mm. Gill length about 1.9 mm.
Frontoclypeus and thorax dorsally with rounded tubercles and multiramous trichomes.
Gill with 8 branches very open, with stout primary branches.
Larva: Maximum length 5.8-6.6 mm. Head pale brown; cephalic apotome slightly
darkened on median basal longitudinal area. Postgenal cleft subtriangular; postgenal bridge
0.7 times length of hypostomium. Antenna as long as cephalic fan stem; ratio of proximal,
medial and distal articles = 1: 0.4-0.5: 1.5-1.6. Cephalic fan with 43-47 rays. Anal ring with
about 100 rows with 13-14 hooks each. Rectal papillae with 7-9 diverticula on each lobe.
Distribution: Brazil: Bahia.
Bionomics: This species breeds in fast-flowing streams of tropical areas on trailing
vegetation and rocks, at a median altitude of about 1600 m. Female feeding habits are
unknown. Females have not been collected biting humans.
Discussion: Adults are externally similar to other species of the S. (Chirostilbia)
pertinax species group, but females can be differentiated by the relatively short paraproct,
and males by the subovoidal finger-like gonostylus and short ventral plate. The pupa has 8
gill branches and a shoe-shaped cocoon, as in S. empascae, but can be differentiated by the
gill primary and secondary branches grouped like a bundle, contrary to the isolated and
open condition in S. jeffersoni. Larvae are similar to those of S. laneportoi.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 683

Simulium (Inaequalium) maranguapense COSTA PESSOA, RÍOS-VÉLASQUEZ & PY-DANIEL


Inaequalium maranguapense COSTA PESSOA et al., 2005: 5-13.
Diagnosis based on original description.
Female (Pharate): Wing length 1.7 mm. General coloration dark brown to black;
scutum without evident silver stripes; legs pale yellowish, with distal ¼ of fore leg tibia
and tarsi blackish; coxa, trochanter and distal portion of femur and tibia of median and
posterior leg blackish. Cibarium not elevated medially, similar to that of S. souzalopesi.
Gonapophysis subtriangular, divergent, with internal border little sclerotized. Paraproct
relatively short, subtriangular.
Male: Unknown.
Pupa: Cocoon slipper shaped. Length at base 2.6-3.1 mm, dorsally 2.1 mm.
Frontoclypeus with abundant rounded tubercles and bifid trichomes. Gill with 4 thick
branches, 2 dorsal and 2 ventral, anteriorly directed and becoming gradually thin distally;
surface with small rounded tubercles. Thorax with rounded tubercles and filiform
trichomes with 1-5 branches.
Larva: Maximum length 4.1-4.8 mm. Cephalic apotome yellowish brown, pattern
similar to that of S. diversibranchium. Antenna longer than cephalic fan stem; ratio of
hypostomium/postgenal bridge = 0.6-0.7. Cephalic fan with 39-41 rays. Lateromandibu-
lar process simple, not bifurcated. Anal ring with 81-83 rows with 9-l3 hooks each. Rectal
papillae with 0-3 diverticula on each lobe.
Distribution: Brazil: Ceará.
Bionomics: This species breeds on deciduous leaves and rocks in small, rocky
streams at an altitude of about 600 m. Female feeding habits are unknown.
Discussion: The female is similar to that of S. souzalopesi, with similar cibarium
morphology, but it can be differentiated by the longer paraproct of S. souzalopesi. The
pupa of S. travassosi is similar, but has only 3 gill branches. The larval rectal papillae,
with 0-3 diverticula on each lobe, is a unique character of S. maranguapense among
species of S. (Inaequalium).

Simulium (Inaequalium) margaritatum PEPINELLI, HAMADA & LUZ


Simulium (Inaequalium) margaritatum PEPINELLI et al., 2006: 48-58.
Female: Wing length 2.0 mm. General coloration blackish; Scutum uniformly black
without silver stripes in any light, and golden green hairs. Cibarium with teeth arranged on
median elevation. Sc with setae; basal section of R bare. Coxa, trochanter, distal portion
of tibia and tarsomeres dark brown. Gonapophysis subtriangular with divergent internal
margins sclerotized. Paraproct relatively short and with apical edge rounded.
Male: Wing length 1.8 mm. Scutum black with 1+1 rounded silver spots, and golden
hairs. Sc and basal section of R bare. Gonostylus subtrapezoidal and ventral plate
relatively elongated, both similar to S. travassosi
Pupa: Cocoon slipper shaped. Length at base 3.0-3.2 mm. Dorsally 2.4-2.7 mm.
Frontoclypeus and thorax with abundant rounded tubercles, sometimes acuminate on
thorax; cephalic trichomes with 2-3 and thoracic trichomes with 3-8 branches. Gill
684 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

length 3.2-3.7 mm, with 4 filiform branches of same length, arising near base from 2
short primary branches.
Larva: Maximum length 4.5-6.0 mm. Cephalic apotome pale dark brown, with
pattern similar to that of S. diversibranchium. Postgenal cleft subtriangular with similar
shape to that of S. diversibranchium. Postgenal bridge 0.7 times as long as hypostomium
length. Subesophageal ganglion pigmented. Antenna as long as cephalic fan stem; ratio of
proximal, medial and distal articles = 1: 0.7-08 : 1.3-1.4. Mandible with 1-2 lateromandib-
ular processes surpassing teeth border. Cephalic fan with 44-48 rays. Rectal papillae with 5
or 6 diverticula on each lobe. Anal ring with about 80 rows with 12-13 hooks each.
Distribution: Brazil: Bahia.
Bionomics: The species breeds on deciduous leaves and rocks in small, rocky-
bottomed creeks, with moderate temperature and slow flow, between 800 and 1700 m of
altitude. Female feeding habits are unknown.
Discussion: The adults of S. margaritatum are similar to those of other species of
the subgenus S. (Inaequalium), particularly S. diversibranchium, which has the same female
scutal coloration and male gonostylus morphology. The number of pupal gill branches
separates S. margaritatum from other species of the subgenus. Simulium maranguapense,
which also has 4 gill branches, has thicker branches. Another unique character of S.
margaritatum, among species of S. (Inaequalium), is the bare basal section of R.

Simulium (Thyrsopelma) duodenicornium PEPINELLI, HAMADA & TRIVINHO-STRIXINO


Simulium (Thyrsopelma) duodenicornium PEPINELLI et al., 2005: 19-27.
Diagnosis based on original description.
Female: Wing length 2.5 mm. Coloration and morphology as for S. guianense and S.
orbitale, except paraproct internally concave in medial area, forming 2 short projections.
Male: Wing length 2.4 mm. Scutum black without silver or gray spots. Genitalia
similar to those of S. orbitale.
Pupa: Similar to that of S. guianense, but gill branches shorter, thicker and without
spicules in annular pattern.
Larva: Maximum length 6.2-7.5 mm. Similar to other species of the subgenus; S.
scutistriatum is most similar, but has more anal hook rows (280-300); S. duodenicornium has
174-184 rows with 26-31 hooks each.
Distribution: Brazil: São Paulo.
Bionomics: This species breeds on stems, leaves and roots of trailing vegetation in
streams with fast flow and warm water at an altitude of about 900 m. Female feeding
habits are unknown.
Discussion: Simulium duodenicornium is similar to S. guianense, but can be differentiated
by its thicker branches of the pupal gill, absence of silver spots on the male scutum, and
medially concave region of the inner margin of the paraproct in the female.
ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales 685

Simulium (Thyrsopelma) jeteri PY-DANIEL, DARWICH, MARDINI, STRIEDER & COSCARÓN


Thyrsopelma jeteri PY-DANIEL et al., 2005: 465-471.
Diagnosis based on original description.
Female and Male: Available material pharate, not showing characters that permit
differentiation from other adults of Thyrsopelma species.
Pupa: Cocoon shoe shaped, not completely protecting gill branches. Length at base
2.5-2.9 mm, dorsally 2.1-2.5 mm. Frontoclypeus and thorax with or without tubercles on
different areas. Gill with 15 thick branches with rounded spicules on pseudoannulations.
Larva: Maximum length 5.2-5.6 mm. Body cuticle with abundant racket-shaped
scales. Cephalic apotome with uniform coloration. Antenna surpassing cephalic fan apex;
ratio of proximal, medial and distal articles 1: 1.7-1.8: 1.1-1.5. Cephalic fan with 51-56
rays. Ratio of postgenal bridge/ hypostomium length = 1.0-1.2; postgenal cleft submiter
shaped. Anal ring with 130-164 rows with 14-19 hooks each. Rectal papillae with 10-15
diverticula on each lobe.
Distribution: Brazil: Rio Grande do Sul.
Bionomics: The species breeds in rocky streams with crystalline water and marginal
vegetation. Female feeding habits are unknown.
Discussion: This species is difficult to differentiate from other Thyrsopelma species;
only the pupal gill branches, numbering 15, permit identification. As in S. duodenicornium, it
has thick gill branches.

REFERENCES
COSTA PESSOA, F.A., RÍOS-VELÁSQUEZ, C.M. & V. PY-DANIEL (2005): First survey of Simuliidae
(Diptera) from North Ceará State, Brazil, with descriptions of a new species and identifica-
tion keys for the immature stages. - Zootaxa 1051: 1-18.
HAMADA, N., HERNÁNDEZ, L.M., BESSA LUZ, S.L. & M. PEPINELLI (2006): Simulium (Chirostilbia)
jeffersoni, a new species of black fly (Diptera: Simuliidae) from the state of Bahia, Brazil. -
Zootaxa: 1123: 21-37.
PEPINELLI, M., HAMADA, N. & S. TRIVINO-STRIXINO (2005): Simulium (Thyrsopelma) duodenicornium, a
new black fly species (Diptera: Simuliidae) from the southeast Region of Brazil. - Zootaxa:
1040: 19-27.
PEPINELLI, M., HAMADA, N. & S.L. BESSA LUZ (2006): Simulium (Inaequalium) margaritatum, a new
species of black fly (Diptera: Simuliidae) from the central region of Bahia State, Brazil. -
Zootaxa 1167: 47-60.
PY-DANIEL, V., DARWICH, S.M., MARDINI, L.B., STRIEDER, M.N. & S. COSCARÓN (2005): Simuliidae
(Diptera, Culicomorpha) no Brasil - XII sobre o gênero Thyrsopelma ENDERLEIN, 1934:
descrição de T. jeteri sp.n, considerações sobre o cibário e chave para as pupas. - Acta
Amazonica 35 (4): 463-472.

You might also like