Professional Documents
Culture Documents
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
Volume 3
Neotropical Simuliidae (Diptera: Insecta)
Volumen 3
Simuliidae Neotropicales (Diptera: Insecta)
2 ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae
Volumen 3
SIMULIIDAE
NEOTROPICALES
(DIPTERA: INSECTA)
por
Sixto Coscarón & Cecilia L. Coscarón Arias
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
Volume 3
NEOTROPICAL
SIMULIIDAE
(DIPTERA: INSECTA)
by
Sixto Coscarón & Cecilia L. Coscarón Arias
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
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)
© 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
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.
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
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
4S TUDY OF MATERIAL 36
7R EFERENCES 648
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).
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
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
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
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
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
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
– 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
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
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
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.
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.
C
A
B
E F
H
G
I J
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.
(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.
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.
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.
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.
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.
– 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
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
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
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.
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.
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.
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
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.
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.
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.
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.
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.
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.
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).
(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.
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
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.
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.
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.
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
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
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.
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.
Differences in the male, pupa and larva are given in the generic keys
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).
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
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
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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
– 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
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
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
– 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
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) 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.
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.
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.
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
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.
– 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
Pupae
Simulium (Pt.) albicinctum, Simulium (Pt.) schoenemanni, and Simulium (Pt.) strigidorswn are not
included in the key below because their pupae are not known.
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
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
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
Pupas
En Simulium (Pt.) albicinctum, Simulium (Pt.) schoenemanni y Simulium (Pt.) strigidorsum no se
conoce la pupa.
– 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
Larvas
En Simulium (Pt.) albicinctum, Simulium (Pt.) bordai, Simulium (Pt.) luchoi, Simulium (Pt.)
schoenemani y Simulium (Pt.) strigidorsum sus larvas son desconocidas.
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
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
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.
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).
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
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) 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
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.)
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.
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.
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
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
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
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
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.
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
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.
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.
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
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.
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
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.
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.
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
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.
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
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.
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.
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
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.
(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
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.
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
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
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
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
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
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.
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.
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
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.
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
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
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.
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
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
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.
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.
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.
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.
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.
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.
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
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
Females
Simulium hauseri and S. leopoldense females are unknown.
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
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
Machos
Simulium hauseri, S. leopoldense y S. petropoliense con machos desconocidos.
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
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
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
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
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.
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.
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.
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.
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.
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.
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.
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.
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.
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
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.
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
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
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
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.
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).
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.
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 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
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.
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.
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.
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.
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.
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.
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.
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
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
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).
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
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
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.
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.
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).
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.
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.
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.
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.
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.
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.
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.
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.
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.
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
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
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
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
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
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
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.
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.
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.
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.
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.
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.
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.
– 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
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
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
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
– 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
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.
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.
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.
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
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.
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.
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.
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.
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.
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.
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.
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.
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.
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
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).
of S. kabanayense can be distinguished by the yellowish brown abdominal tergites I-II, and
the pupa by the gill of 14-23 branches.
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.
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.
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.
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.
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.
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
Species inquirenda
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.
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
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
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
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
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
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
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.
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
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.
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
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.
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.
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).
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.
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.
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).
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.
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.
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.
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.
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).
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.
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
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.
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
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.
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).
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
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 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.
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
Larvas
No incluidas Simulium costalimai, S. covagarciai, S. kompi, S. matteabranchium, S. morae, S.
racenisi y S. sandyi.
– 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
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) 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.
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.
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.
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.
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.
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
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
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).
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.
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.
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
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
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.
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.
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.
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.
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
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
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
Larvae
Not included: Simulium freemani, S. keenani and S. lobatoi.
– 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
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
Larvas
No incluidas: Simulium freemani, S. keenani y S. lobatoi.
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
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.
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.
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.
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.
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.
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.
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
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).
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.
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
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) 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.
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.
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.
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.
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 (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.
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
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
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
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.
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.
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) 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.
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) 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.
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.
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
III
III
V
V
VII
VII
IX
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.
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) 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.
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).
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
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.
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
Larvas
No se presenta clave por escaso material y caracteres.
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
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).
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.
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.
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.
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
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.
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.
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.
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.
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.
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 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.
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
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
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
Simulium (Notolepria)
S. blantoni
S. cuasiexiguum
S. exiguum
S. llutense
S. paraguayense
S. subexiguum
S. gonzalezi
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
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
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
Simulium (Cerqueirellum)
S. amazonicum
S. argentiscutum
S. chaquense
S. cuneatum
S. delponteianum
S. ganalesense
S. minusculum
S. oyapockense
S. raraimense
S. sanguineum
S. venezuelense
Simulium (Coscaroniellum)
“QUADRIFIDUM GROUP”
S. cauchense
S. cerradense
S. dolianhanhani
S. goeldii
S. quadrifidum
S. ulyssesi
“QUADRIVITTATUM GROUP”
S. quadrivittatum
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
Simulium (Ectemnaspis)
“ROMANAI GROUP”
S. albanense
S. gabaldoni
S. lutzianum
S. mayuchuspi
S. romanai
S. wolffhuegeli
Simulium (Ectemnaspis)
“PERFLAVUM GROUP”
S. kabanayense
S. maroniense
S. perflavum
S. rorotaense
S. suarezi
S. trombetense
Simulium (Ectemnaspis)
“DINELLII GROUP”
S. antillarum
S. dinellii
S. nuneztovari
S. ochraceum
S. shewellianum
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
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
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
Simulium (Trichodagmia)
S. huairayacu
S. lahillei
S. muiscorum
S. nigrimanum
S. sumapazense
S. townsendi
S. wygodzinskyorum
Simulium (Thyrsopelma)
S. quianense
S. hirtipupa
S. itaunense
S. orbitale
S. perplexum
S. scutistriatum
7
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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
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
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
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
9
ADDENDUM
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.
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.