Professional Documents
Culture Documents
net/publication/221940759
CITATIONS READS
47 1,488
5 authors, including:
Some of the authors of this publication are also working on these related projects:
All content following this page was uploaded by José L Luque on 08 February 2014.
ZOOTAXA
3082
Magnolia Press
Auckland, New Zealand
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Materials and methods. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Key to Adult Nematoda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Key to Larval Nematoda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Parasite-Host List . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
Host–Parasite List . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76
Abstract
Seventy-four genera comprising approximately 140 named and 75 undetermined species of adult and larval nematodes are
associated with about 330 named and 50 undetermined species of marine, brackish and freshwater fishes from Brazilian waters.
These are listed with information on their hosts, habitat, distribution and records. A key to these parasites at the generic level is
provided.
Key words: Biodiversity, Nematoda, fish parasites, checklist, key, Neotropics, Brazil
Introduction
The phylum Nematoda represent a taxonomically, ecologically and geographically diverse helminth group, with an
enormous number of species, both parasitic and freeliving forms, a huge economic and ecological impact, and a
number of individuals estimated at 1022 (Hodda, 2007). Because of the enormous abundance and diversity of the
group, only an estimated 2–10% of the world fauna has been formally described, despite its importance (Hodda
2007). Nematodes represent a conspicuous component of the parasite assemblages in fishes. According to Luque
& Poulin (2007), nematodes are the second group, after digeneans, in terms of the number of parasite associations
recorded in fishes from the Neotropics.
Studies on nematodes parasitic in Brazilian fishes began with the description of Trichocephalus gibbosus
Rudolphi [now Oncophora melanocephala (Rudolphi)] by Rudolphi (1819) based on specimens collected from
Thunnus thynnus (L.) off the coast of Brazil by Pohl in 1817. The first freshwater records appear to be of Lecano-
cephalus spinulosus Diesing [now Goezia spinulosa (Diesing)] and Cheiracanthus gracilis Diesing [now Gnathos-
toma gracile (Diesing)], both of which Diesing (1839) recorded from Arapaima gigas (Schinz). However, not until
the work of Travassos et al. (1928), which recorded numerous nematodes from freshwater fishes, were detailed
studies undertaken. Subequently, a large number of papers by many authors, especially in recent years, have added
to our knowledge of this fauna. Compilations of records have been published by Thatcher (1991, 2006), restricted
to the Amazon River basin, and by Vicente et al. (1985), supplemented by Vicente & Pinto (1999). Moravec
(1998), in his monograph on nematodes parasites of Neotropical freshwater fishes, also included numerous records
from Brazil. Some of these compilations have included taxonomic keys, descriptions and/or aspects of the biology
of the different taxa. Most recently, Muniz-Pereira et al. (2009) included records of nematode species in a checklist
of helminth parasites of threatened vertebrate species from Brazil, and Eiras et al. (2010) published an account of
the nematode species associated with Brazilian freshwater fishes.
Here we provide a checklist of the nematode species associated with both marine and freshwater fishes from
Brazil, including detailed information on locality, habitat, site of infection and life cycle stage. The information is
presented as parasite-host and host-parasite lists, with a key to the generic level. We have attempted to include
information from all published reports and have added a small number of original records.
This bibliographic review of the nematode species reported from fishes in Brazilian waters was based on informa-
tion collected from two main sources. Firstly, searches of the Host–Parasite Catalogue of the Natural History
Museum in London for 1920 to 1997 and, for the years 1988–2003, the Host–Parasite Database, a partial imple-
mentation of which is now available on–line (Gibson et al., 2005). Secondly, searches of the Zoological Record,
Biological Abstracts and Helminthological Abstracts, Web of Knowledge, Google Scholar, Aquatic Sciences and
Fisheries Abstracts, Biological and Agricultural Index Plus and the Scopus databases were undertaken up to July,
2010 (with an update from Web of Knowledge in September, 2011). Original data from our own studies are also
included. The checklist follows the parasite classification and systematic arrangement of Moravec (1998, 2006),
Anderson et al. (2009) and Gibbons (2010), with some modifications based on recent records. The nematodes are
arranged according to class, order, suborder, superfamily, family and subfamily, within which the species are pre-
sented in alphabetical order. Host species are given followed by their predominant habitat (marine—MAR, fresh-
water—FW or brackish water—BW) and site of the infection; when possibe these are grouped. Localities are given
in alphabetical order of Brazilian states (AM—Amazonas, AP—Amapá, BA—Bahia, CE—Ceará, ES—Espírito
Santo, MA—Maranhão, MG—Minas Gerais, MS—Mato Grosso do Sul, MT—Mato Grosso, PA—Pará, PB—
Paraíba, PI—Piauí, PR—Paraná, RJ—Rio de Janeiro, RO—Rondônia, RN—Rio Grande do Norte, RS—Rio
Grande do Sul, and SP—São Paulo) preceded by the localities when possibly and, records in chronological
sequence.
The fish species names were updated according to Froese & Pauly (2010), supplemented by Eschmeyer
(2010). The nematode species names have been updated to agree with the recent literature, but inclusion in the par-
asite or host lists does not imply that the authors necessarily agree with their validity. The species of fish in the
host–parasite list are arranged according to the higher level classification of Nelson (2006).
Results
The nematodes parasitic in Brazilian fishes listed below are distributed in 74 genera comprising 142 named and 75
undetermined species of adult and larval nematodes. The total number of host-parasite associations listed is 1,688.
These worms were recorded from 324 named marine, brackish and freshwater fish species, and from 50 undeter-
mined host species.
Key to Nematoda associated with Brazilian fishes (based on Moravec 1998, Anderson et al. 2009, and Gibbons
2010)
1. Oesophagus cylindrical or forming stichosome or trophosome. Phasmids and deirids absent. Eggs with plug at either pole or
hatching in utero .............................................................................................................Class Adenophorea, Order Enoplida (2)
- Oesophagus never forming stichosome. Phasmids and deirids present. Eggs without polar plugs. Third larva stage infective to
final host ..................................................................................................................................................... Class Secernentea (7)
2. Oesophagus cylindrical and well developed; stichosome or trophosome absent. Male tail modified to form ventral sucker .........
. ............................................................................. Superfamily Dioctophymatoidea, Family Dioctophymatidae, Eustrongylides
- Oesophagus with oesophageal glands forming stichosome or trophosome. Male tail not modified to form ventral sucker...........
. ....................................................................................................................Superfamily Trichinelloidea, Family Trichuridae (3)
3. Spicule absent. Male distal end rounded, laterally expanded ............................................................................ Freitascapillaria
- Spicule present. Male distal end not laterally expanded ...........................................................................................................(4)
4. Spicular sheath with spines . ......................................................................................................................................... Capillaria
- Spicular sheath without spines .................................................................................................................................................. (5)
5. Pseudobursal membrane absent or very reduced ............................................................................................... Pseudocapillaria
- Pseudobursal membrane well developed ...................................................................................................................................(6)
6. Dorsolateral projection of pseudobursal margin narrow or digitiform ...................................................................Paracapillaria
- Dorsolateral projections of pseudobursal margin very wide (in ventral view) .............................................Capillostrongyloides
1. Oral aperture surrounded by 12 or 18 papillae arranged in two or three circles of six papillae each. . . . . . . . . . . Eustrongylides
- Oral aperture without these characteristics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (2)
2. Oral aperture without pseudolabia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (3)
- Oral aperture provided with pseudolabia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .(11)
3. Ventriculus with several short anterior and posterior appendices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Brevimulticaecum
- Ventriculus without appendices or with only one, usually long, posterior appendix . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .(4)
4. Anterior intestinal caecum present and posterior ventricular appendix absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (5)
- Anterior intestinal caecum and posterior ventricular appendix present, or both caecum and appendix absent . . . . . . . . . . . . .(7)
5. Excretory pore at about level of nerve ring or just posterior to it . . . . . . . . . . . . . . . . . . . . . . . . . . . . “Porrocaecum-like” larvae
- Excretory pore near ventral interlabium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (6)
6. Tail with mucron . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudoterranova
- Tail without mucron . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Terranova
7. Cuticular rings of older larvae provided with numerous small posterior spines. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Goezia
- Cuticular rings always lack spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .(8)
8. Both posterior ventricular appendix and anterior intestinal caecum present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .(9)
- Only posterior ventricular appendix or only anterior intestinal caecum present, or both absent . . . . . . . . . . . . . . . . . . . . . . .(10)
9. Excretory pore at level of nerve ring or just posterior . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hysterothylacium
- Excretory pore at base of ventral interlabium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Contracaecum
10. Posterior ventricular appendix and anterior intestinal caecum absent. Excretory pore at base of ventral interlabium . . Anisakis
- Posterior ventricular appendix present; anterior intestinal caecum absent. Excretory pore at or posterior to level of nerve ring .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Raphidascaris
11. Mouth provided with two large, lateral, triangular pseudolabia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Spiroxys
- Cephalic end bears helmet-like cuticular structure with thickened, free posterior region . . . . . . . . . . . . . . . . . Pseudoproleptus
Amplicaecum sp.—FW
Hosts and site of infection: Brycon orthotaenia (larva—intestine, swim bladder wall), Hoplias malabaricus
(unspecified—intestine), Leporinus copelandii (unspecified—intestine), Pseudoplatystoma sp. (larva—
swim bladder wall), Salminus hilarii (unspecified—intestine), Steindachneridion parahybae (larva—
intestine, swim bladder).
Locality: Pirassununga—SP.
References: Travassos et al. (1928), Travassos & Kohn (1965), Vicente et al. (1985), Kohn & Fernandes
(1987), Vicente & Pinto (1999), Muniz-Pereira et al. (2009).
Angusticaecum sp.—larva—MAR—intestine
Host: Centropomus parallelus
Locality: Rio de Janeiro—RJ
References: Lamothe-Argumedo et al. (1996)
Remark: Toxocarines are typically parasites of terrestrial higher vertebrates and the Brazilian records of Porrocae-
cum spp. are doubtful (Moravec, 1994, 1998). So, all records of Porrocaecum spp. from fishes are likely to be mis-
identifications of other ascaridiod, probably anisakid taxa, and, thus, can be considered as ‘Porrocaecum-like’
species.
References: Pinto & Noronha (1976), Vicente & Pinto (1981), Vicente et al. (1985), Rodrigues et al. (1991),
Thatcher (1991, 2006), Moravec (1998), Eiras et al. (2010).
Remark: Thatcher (2006) suggested that the “Jeju” may be an erythrinid.
Procamallanus (Spirocamallanus) pereirai Annereaux—adult—MAR
Hosts: Atlantoraja castelnaui (spiral valve), Menticirrhus americanus (intestine), Micropogonias furnieri
(intestine), Nebris microps (intestine), Paralonchurus brasiliensis (intestine), Porichthys porosissimus
(intestine), Stellifer brasiliensis (intestine), Symphurus tessellatus (intestine).
Localities: Araruama, Rio de Janeiro—RJ; Coast—RS.
References: Pinto et al. (1984), Rodrigues et al. (1991), Santos et al. (1999), Knoff et al. (2001), Ribeiro et
al. (2002), Luque et al. (2003).
Procamallanus (Spirocamallanus) pexatus Pinto, Fábio, Noronha & Rolas—adult—FW—intestine
Host: Pygidium brasiliensis.
Locality: Santa Tereza—ES.
Spiroxys sp.—larva—FW
Hosts and site of infection: Astyanax altiparanae (intestine), Cichla kelberi (body cavity, intestine),
Gymnotus carapo, Gymnotus sp. (intestine), Pygocentrus piraya (body cavity, intestinal caecum, intestine,
gall bladder), Serrasalmus brandtii (body cavity, stomach, intestine).
Localities: Três Marias Reservoir, São Francisco River—MG; Baía River—MS; Upper Paraná River—PR.
References: Isaac et al. (2004), Pavanelli et al. (2004), Santos et al. (2009), Takemoto et al. (2009), Eiras et
al. (2010).
Final remarks:
Two records founded in our bibliographic searches are clearly misidentified or could be accidental associations.
Kohn & Macedo (1984) recorded the oxyurid Aspiculurus tetraptera (Nitzsch, 1821), a common parasite of
rodents, in Haemulon sciurus off Rio de Janeiro. Another record, that of Raphidascaris (Raphidascaris) acus
(Bloch, 1779) recorded by Guimarães (1975) in Lutjanus vivanus off the Salvador (Bahia), is considered a misiden-
tification, because this is a Holarctic freshwater species (Anderson, 2000).
Order Anguilliformes
Family Muraenidae
Echidna catenata (Bloch) Heliconema heliconema
Gymnothorax moringa (Cuvier) Hysterothylacium sp.
Raphidascaris sp.
Terranova sp.
Family Ophichthidae
Ophichthus ophis (Linnaeus) Heliconema heliconema
O. gomesii (Castelnau) Heliconema heliconema
Order Batrachoidiformes
Family Batrachoididae
Porichthys porosissimus (Cuvier) Procamallanus (S.) pereirai
Order Beloniformes
Family Belonidae
Tylosurus acus acus (Lacépède) Hysterothylacium sp.
Terranova sp.
Order Clupeiformes
Family Clupeidae
Clupeidae gen. sp. Procamallanus (S.) barroslimai
Brevoortia aurea (Spix & Agassiz) Anisakis sp.
Contracaecum sp.
Procamallanus sp.
Terranova sp.
Harengula clupeola (Cuvier) Hysterothylacium fortalezae
Raphidascaris (I.) vicentei
Raphidascaris sp.
Sardinella sp. Raphidascaris (I.) vicentei
Cosmocercinae gen. sp.
Family Pristigasteridae
Pellona castelnaeana Valenciennes Nematoda gen. sp.
Order Characiformes
Family Acestrorhynchidae
Acestrorhynchus falcatus (Bloch) Paracapillaria (P.) piscicola
Procamallanus (S.) inopinatus
Acestrorhynchus lacustris (Lütken) Anisakidae gen. sp.
Contracaecum sp.
Contracaecum sp. Type 1
Contracaecum sp. Type 2
Eustrongylides sp.
Philometridae gen. sp.
Philonema sp. A
Procamallanus (S.) sp.
Procamallanus (S.) saofranciscensis
Procamallanus sp.
Travassosnema travassosi paranaensis
Travassosnema travassosi travassosi
Acestrorhynchus microlepis (Jardine) Procamallanus (S.) hilarii
Acestrorhynchus sp. Paracapillaria (P.) piscicola
“Peixe-cachorro” Procamallanus sp.
Procamallanus (S.) sp.
Family Anostomidae
Anostominae gen. sp. Capillostrongyloides sentinosa
Procamallanus (S.) iheringi
“Chimboré” Procamallanus (S.) sp.
“Ferreirinha” Procamallanus (S.) sp.
Leporellus vittatus (Valenciennes) Ancyracanthus schubarti
Capillaria sp.
Contracaecum sp.
Cucullanus sp.
Cucullanus sp. A
Filocapsulariinae gen. sp.
Goezia spinulosa
Nematoda gen. sp.
Paracapillaria (P.) piscicola
Procamallanus (S.) inopinatus
Rhabdochona acuminata
Family Characidae
Aphyocharax anisitsi Eigenmann & Kennedy Nematoda gen. sp.
Astyanax altiparanae Garutti & Britski Contracaecum sp.
Contracaecum sp. Type 1
Procamallanus (S.) caballeroi
Procamallanus (S.) inopinatus
Spiroxys sp.
Astyanax bimaculatus (Linnaeus) Camallanidae gen. sp.
Contracaecum sp.
Contracaecum sp. Type A
Contracaecum sp. Type B
Cosmoxynemoides aguirrei
Cucullanus sp.
Nematoda gen. sp.
Paraseuratum albidum
Procamallanus (S.) hilarii
Procamallanus (S.) inopinatus
Procamallanus (S.) sp.
Procamallanus (S.) sp. (= "P. laticonchus")
Procamallanus sp.
Rhabdochona acuminata
Spinitectus rodolphiheringi
Travnema travnema
Astyanax eigenmanniorum (Cope) Nematoda gen. sp.
Astyanax fasciatus (Cuvier) Ascarididae gen. sp.
Capillaria sp.
Capillostrongyloides sentinosa
Contracaecum sp.
Contracaecum sp. Type 1
Contracaecum sp. Type 2
Paraseuratum albidum
Procamallanus (S.) hilarii
Procamallanus (S.) iheringi
Procamallanus (S.) inopinatus
Procamallanus (S.) saofranciscensis
Procamallanus (S.) sp.
Family Curimatidae
Curimata sp. Contracaecum sp.
Philonema sp. B
Curimatella lepidura (Eigenmann & Eigenmann) Cosmoxynemoides aguirrei
Cyphocharax gilbert (Quoy & Gaimard) Camallanidae gen. sp.
Cosmoxynema vianai
Cosmoxynemoides aguirrei
Guyanema sp.
Nematoda gen. sp.
Philonema sp. B
Raphidascaris sp.
Travnema araujoi
Travnema travnema
Travnema sp.
Family Cynodontidae
Rhaphiodon vulpinus Spix & Agassiz Ascarididae gen. sp.
Contracaecum sp.
Contracaecum sp. Type 1
Contracaecum sp. Type 2
Eustrongylides ignotus
Eustrongylides sp.
Goezia sp.
Guyanema raphiodoni
Hysterothylacium sp.
Nematoda gen. sp.
Family Erythrinidae
Erythrinus erythrinus (Bloch & Schneider) Paraseuratum albidum
Procamallanus (S.) paraensis
Hoplerythrinus unitaeniatus (Spix & Agassiz) Contracaecum sp.
Contracaecum sp. Type 1
Contracaecum sp. Type 2
Eustrongylides sp.
Nematoda gen. sp.
Hoplias lacerdae Miranda Ribeiro Procamallanus (S.) hilarii
Hoplias malabaricus (Bloch) Amplicaecum sp.
Ascarididae gen sp.
Ascaridoidea fam. gen. sp.
Brevimulticaecum sp.
Capillostrongyloides sentinosa
Family Hemiodontidae
Bivibranchia fowleri (Steindachner) Procamallanus (S.) sp.
Bivibranchia velox (Eigenmann & Myers) Procamallanus (S.) sp.
Family Prochilodontidae
Prochilodus argenteus Spix & Agassiz Spinitectus asperus
Prochilodus lineatus (Valenciennes) Anisakidae gen. sp.
Contracaecum sp.
Contracaecum sp. Type 2
Procamallanus (S.) inopinatus
Procamallanus (S.) sp.
Procamallanus sp.
Raphidascaris sp.
Spinitectus asperus
Order Cypriniformes
Family Rhamphichthyidae
Rhamphichthys rostratus (Linnaeus) Cucullanus (T.) rhamphichthydis
Rhamphichthys hahni (Meinken) Nematoda gen. sp.
Family Rivulidae
Austrolebias bellottii (Steindachner) Hedruris iheringi
Order Gadiformes
Family Merluccidae
Merluccius hubbsi Marini Anisakis sp.
Hysterothylacium sp.
Raphidascaris sp.
Family Phycidae
Urophycis brasiliensis (Kaup) Ascarophis marina
Contracaecum sp.
Cucullanus sp.
Hysterothylacium sp.
Procamallanus (S.) macaensis
Raphidascaris sp.
Urophycis mystacea Miranda Ribeiro Anisakis sp.
Contracaecum sp.
Cucullanus cirratus
Hysterothylacium sp.
Urophycis sp. Procamallanus (S.) macaensis
Order Gymnotiformes
Family Gymnotidae
Gymnotus carapo Linnaeus Capillariinae gen. sp.
Contracaecum sp.
Contracaecum sp. Type 1
Contracaecum sp. Type 2
Eustrongylides sp.
Hysterothylacium sp.
Nematoda gen. sp.
Procamallanus (P.) peraccuratus
Spiroxys sp.
Family Sternopygidae
Sternopygus macrurus (Bloch & Schneider) Spinitectus rodolphiheringi
Order Lampriformes
Family Trachipteridae
Trachipterus jacksonensis (Ramsay) Lappetascaris lutjani
Order Lophiiformes
Family Lophiidae
Lophius gastrophysus Miranda Ribeiro Hysterothylacium sp.
Nematoda gen. sp.
Raphidascaris sp.
Order Ophidiiformes
Family Ophidiidae
Genypterus brasiliensis Regan Anisakidae gen. sp.
Anisakis physeteris
Anisakis simplex
Anisakis sp.
Contracaecum sp.
Cucullanus genypteri
Hysterothylacium sp.
Nematoda gen. sp.
Procamallanus sp.
Pseudoterranova decipiens
Pseudoterranova sp.
Raphidascaris sp.
Terranova sp.
Order Osteoglossiformes
Family Arapaimidae
Arapaima gigas (Schinz) Camallanus (C.) tridentatus
Capillostrongyloides arapaimae
Eustrongylides sp.
Gnathostoma gracile
Goezia spinulosa
Nilonema senticosum
Family Osteoglossidae
Osteoglossum bicirrhosum (Cuvier) Camallanus (C.) acaudatus
Camallanus (C.) tridentatus
Order Perciformes
“Acará-açú” Goezia spinulosa
“Cará-cachimbo” Procamallanus sp.
Procamallanus (S.) sp.
Caranx hippos (Linnaeus) Contracaecum sp.
Cucullanus pulcherrimus
Philometra sp.
Raphidascaris sp.
Terranova sp.
Caranx latus Agassiz Anisakis sp.
Contracaecum sp.
Hysterothylacium sp.
Terranova sp.
Caranx lugubris Poey Cucullanus pulcherrimus
Oligoplites palometa (Cuvier) Contracaecum sp.
Oligoplites saliens (Bloch) Contracaecum sp.
Oligoplites saurus (Bloch & Schneider) Contracaecum sp.
Porrocaecum secundum
Terranova trichiuri
Parona signata (Jenyns) Contracaecum sp.
Hysterothylacium sp.
Raphidascaris sp.
Selene setapinnis (Mitchill) Anisakis sp.
Contracaecum sp.
Hysterothylacium sp.
Raphidascaris sp.
Terranova sp.
Selene vomer (Linnaeus) Porrocaecum sp.
Raphidascaris sp.
Trachinotus carolinus (Linnaeus) Ascarophis brasiliensis
Trachurus lathami Nichols Contracaecum sp.
Terranova sp.
Uraspis secunda (Poey) Hysterothylacium sp.
Raphidascaris sp.
Scombroides occidentalis (Bloch & Schneider) Raphidascaris sp.
Family Centrarchidae
Micropterus salmoides (Lacépède) Goezia spinulosa
Family Cichlidae
Apistogramma cacatuoides Hoedeman Camallanus cotti
Astronotus ocellatus (Agassiz) Camallanus sp.
Contracaecum sp.
Contracaecum sp. Type 2
Goezia spinulosa
Nematoda gen. sp.
Procamallanus (S.) inopinatus
Australoheros facetus (Jenyns) Procamallanus (P.) peraccuratus
Procamallanus sp.
Cichla kelberi Kullander & Ferreira Contracaecum sp.
Contracaecum sp. Type 1
Eustrongylides sp.
Hysterothylacium sp.
Procamallanus (S.) inopinatus
Spiroxys sp.
Cichla monoculus Spix & Agassiz Contracaecum sp.
Eustrongylides sp.
Cichla ocellaris Bloch & Schneider Contracaecum sp.
Eustrongylides sp.
Procamallanus (P.) peraccuratus
Cichla piquiti Kullander & Ferreira Contracaecum sp. Type 1
Eustrongylides sp.
Cichlasoma facetum (Jenyns) Procamallanus (P.) peraccuratus
Cichlasoma sp. Ichthyouris ovofilamentosa
Ichthyouris sp.
Crenicichla britskii Kullander Nematoda gen. sp.
Crenicichla haroldoi (Luengo & Britski) Procamallanus (S.) inopinatus
Crenicichla lepidota Heckel Contracaecum sp. Type 1
Contracaecum sp.
Hysterothylacium sp.
Procamallanus (P.) peraccuratus
Crenicichla niederleinii (Holmberg) Nematoda gen. sp.
Geophagus brasiliensis (Quoy & Gaimard) Contracaecum sp.
Cosmoxynemoides aguirrei
Nematoda gen. sp.
Procamallanus (P.) peraccuratus
Raphidascaris (S.) sp.
Rhabdochona acuminata
Geophagus proximus (Castelnau) Raphidascaris (S.) sp.
Family Coryphaenidae
Coryphaena hippurus Linnaeus Nematoda gen. sp.
Family Ephippidae
Chaetodipterus faber (Broussonet) Porrocaecum sp.
Procamallanus (S.) macaensis
Family Gempylidae
Thyrsitops lepidopoides (Cuvier) Nematoda gen. sp.
Procamallanus (S.) macaensis
Raphidascaris sp.
Family Gerreidae
Diapterus rhombeus (Cuvier) Contracaecum sp.
Cucullanus sp.
Family Haemulidae
Haemulon plumierii (Lacépède) Philometra lateolabracis
Family Labridae
Halichoeres radiatus (Linnaeus) Dichelyne travassosi
Family Lutjanidae
Lutjanus synagris (Linnaeus) Raphidascaris (I.) vicentei
Philometra sp.
Family Mugilidae
Mugil curema Valenciennes Cucullanus sp.
Mugil liza Valenciennes Cucullanus (C.) grandistomis
Hysterothylacium sp.
Mugil platanus Günther Cucullanus pulcherrimus
Family Osphronemidae
Betta splendens Regan Camallanus cotti
Family Percophidae
Percophis brasiliensis Quoy & Gaimard Anisakis sp.
Contracaecum sp.
Hysterothylacium sp.
Procamallanus sp.
Raphidascaris sp.
Terranova sp.
Family Pinguipedidae
Pinguipes brasilianus Cuvier Cucullanus carioca
Hysterothylacium sp.
Paracapillaria (P.) argentinensis
Raphidascaris sp.
Pseudopercis numida Miranda-Ribeiro Anisakidae gen. sp.
Anisakis sp.
Capillaria sp.
Cucullanus carioca
Dichelyne sp.
Hysterothylacium sp.
Nematoda gen. sp.
Paracapillaria (P.) argentinensis
Procamallanus sp.
Raphidascaris sp.
Terranova sp.
Pseudopercis semifasciata (Cuvier) Capillaria sp.
Cucullanus carioca
Dichelyne sp.
Hysterothylacium sp.
Paracapillaria (P.) argentinensis
Procamallanus sp.
Raphidascaris sp.
Terranova sp.
Family Polynemidae
Polydactylus virginicus (Linnaeus) Procamallanus (S.) cruzi
Family Priacanthidae
Priacanthus arenatus Cuvier Contracaecum sp.
Nematoda gen. sp.
Oncophora melanocephala
Raphidascaris sp.
Terranova sp.
Family Rachycentridae
Rachycentron canadum (Linnaeus) Iheringascaris inquies
Family Sciaenidae
Cynoscion guatucupa (Cuvier) Cucullanus sp.
Hysterothylacium sp.
Terranova sp.
Cynoscion leiarchus (Cuvier) Nematoda gen. sp.
Macrodon ancylodon (Bloch & Schneider) Contracaecum sp.
Cucullanus sp.
Hysterothylacium sp.
Terranova trichiuri
Terranova sp.
Menticirrhus americanus (Linnaeus) Capillaria sp.
Contracaecum sp.
Hysterothylacium sp.
Nematoda gen. sp.
Procamallanus (S.) macaensis
Procamallanus (S.) pereirai
Procamallanus sp.
Micropogonias furnieri (Desmarest) Anisakidae gen. sp.
Anisakis sp.
Contracaecum sp.
Cucullanus cassinensis
Cucullanus pulcherrimus
Cucullanus sp.
Dichelyne (C.) amaruincai
Dichelyne (C.) elongatus
Dichelyne (D.) micropogonii
Dichelyne (C.) rodriguesi
Dichelyne (C.) sciaenidicola
Family Serranidae
Cephalopholis fulva (Linnaeus) Contracaecum sp.
Raphidascaris sp.
Epinephelus guttatus (Linnaeus) Hysterothylacium eurycheilum
Epinephelus sp. Nematoda gen. sp.
Mycteroperca sp. Nematoda gen. sp.
Family Sparidae
Archosargus rhomboidalis (Linnaeus) Hysterothylacium sp.
Diplodus argenteus argenteus (Valenciennes) Nematoda gen. sp.
Pagrus pagrus (Linnaeus) Anisakidae gen. sp.
Anisakis sp.
Contracaecum sp.
Family Sphyraenidae
Sphyraena guachancho Cuvier Anisakis sp.
Raphidascaris sp.
Family Stromateidae
Peprilus paru (Linnaeus) Contracaecum sp.
Hysterothylacium sp.
Raphidascaris sp.
Family Trichiuridae
Trichiurus lepturus Linnaeus Anisakis sp.
Contracaecum sp.
Hysterothylacium sp.
Procamallanus sp.
Pseudoterranova sp.
Raphidascaris sp.
Terranova sp.
Order Pleuronectiformes
Family Achiridae
Catathyridium jenynsii (Günther) Nematoda gen. sp.
Family Cynoglossidae
Symphurus tessellatus (Quoy & Gaimard) Procamallanus (S.) pereirai
Family Paralichthyidae
Citharichthys macrops Dresel Procamallanus (S.) halitrophus
Paralichthys isosceles Jordan Anisakis simplex
Anisakis sp.
Contracaecum sp.
Hysterothylacium sp.
Raphidascaris sp.
Terranova sp.
Syacium papillosum (Linnaeus) Procamallanus (S.) halitrophus
Family Dactylopteridae
Dactylopterus volitans (Linnaeus) Contracaecum sp.
Order Scorpaeniformes
Family Triglidae
Prionotus punctatus (Bloch) Anisakis sp.
Cucullanus sp.
Hysterothylacium sp.
Procamallanus sp.
Raphidascaris sp.
Terranova sp.
Order Siluriformes
“Bagre” Procamallanus (S.) rarus
Siluriformes fam. gen. sp. Ascarididae gen. sp.
Procamallanus sp.
Procamallanus (S.) solani
Family Ariidae
Bagre bagre (Linnaeus) Anisakis sp.
Terranova trichiuri
Cathorops agassizii (Eigenmann & Eigenmann) Falcaustra mirandafroesi
Philometra fariaslimai
Porrocaecum jardimfreire
Genidens barbus (Lacépède) Contracaecum sp.
Falcaustra mirandafroesi
Hysterothylacium sp.
Philometra fariaslimai
Porrocaecum jardimfreire
Genidens genidens (Cuvier) Falcaustra mirandafroesi
Philometra fariaslimai
Porrocaecum jardimfreire
Ariidae gen. sp. Cucullanus sp.
Family Auchenipteridae
Ageneiosus inermis (Linnaeus) Contracaecum sp.
Procamallanus (S.) inopinatus
Nematoda gen. sp.
Ageneiosus militaris Valenciennes Cucullanus (C.) pinnai pinnai
Goezia sp.
Ageneiosus ucayalensis Castelnau Cucullanus ageneiosus
Procamallanus (S.) belenensis
Procamallanus (S.) rarus
Pseudoproleptus sp.
Family Callichthyidae
Callichthys callichthys (Linnaeus) Eustrongylides sp.
Corydoras aeneus (Gill) Procamallanus (S.) pintoi
Corydoras paleatus (Jenyns) Procamallanus (S.) pintoi
Hoplosternum littorale (Hancock) Capillariinae gen. sp.
Goezia sp.
Lepthoplosternum pectorale (Boulenger) Eustrongylides sp.
Family Doradidae
Acanthodoras spinosissimus (Eigenmann & Rondonia rondoni
Eigenmann)
Franciscodoras marmoratus (Lütken) Porrocaecum sp.
Spinitectus rodolphiheringi
Spinitectus sp.
Oxydoras kneri Bleeker Rondonia rondonia
Spinoxyuris oxydoras
Oxydoras niger (Valenciennes) Cucullanus (C.) grandistomis
Platydoras costatus (Linnaeus) Brevimulticaecum sp.
Porrocaecum sp.
Pterodoras granulosus (Valenciennes) Cucullanus (C.) pinnai pterodorasi
Cucullanus sp.
Cucullanus sp. 2
Hysterothylacium sp.
Klossinemella iheringi
Neoparaseuratum travassosi
Paracamallanus amazonensis
Procamallanus (S.) inopinatus
Family Hemiodontidae
Hemiodus sp. Procamallanus (S.) sp.
Procamallanus sp.
Family Heptapteridae
Pimelodella gracilis (Valenciennes) Cucullanus (C.) pinnai pinnai
Pimelodella lateristriga (Lichtenstein) Brasilnema pimelodellae
Contracaecum sp.
Cucullanus (C.) pimelodellae
Procamallanus (S.) pimelodus
Procamallanus (S.) rarus
Rhabdochona acuminata
Spinitectus rodolphiheringi
Spinitectus yorkei
Rhamdia quelen (Quoy & Gaimard) Capillariinae gen. sp.
Contracaecum sp. Type 2
Contracaecum sp.
Cucullanus sp.
Cucullanus (C.) pinnai pinnai
Procamallanus (S.) hilarii
Family Loricariidae
Ancistrus cirrhosus (Valenciennes) Raphidascaris (S.) hypostomi
Raphidascaris sp.
Hypostomus affinis (Steindachner) Paracapillaria (Paracapillaria) piscicola
Hypostomus albopunctatus (Regan) Procamallanus (P.) annipetterae
Raphidascaris (S.) hypostomi
Raphidascaris sp.
Hypostomus cochliodon Kner Procamallanus (P.) annipetterae
Raphidascaris (S.) hypostomi
Hypostomus commersoni Valenciennes Raphidascaris sp.
Raphidascaris (S.) hypostomi
Family Pimelodidae
Bergiaria westermanni (Lütken) Procamallanus (S.) freitasi
Bergiaria sp. Contracaecum sp.
Brachyplatystoma sp. Ascarididae gen. sp.
Cucullanus sp.
Calophysus macropterus (Lichtenstein) Alinema amazonicum
Eustrongylides sp.
Procamallanus (S.) inopinatus
Procamallanus sp.
Conorynchus conirostris (Valenciennes) Anisakidae gen. sp.
Procamallanus (S.) sp.
Rhabdochona sp.
Hemisorubim platyrhynchos (Valenciennes) Brevimulticaecum sp.
Contracaecum sp. 1
Contracaecum sp. 2
Contracaecum sp. Type 1
Contracaecum sp. Type 2
Contracaecum sp.
Cucullanus (C.) zungaro
Eustrongylides ignotus
Goezia sp.
Nematoda gen. sp.
Hypophthalmus edentatus Spix & Agassiz Hysterothylacium sp.
Paracamallanus amazonensis
Paracamallanus sp.
Family Pseudopimelodidae
Pseudopimelodus mangurus (Valenciennes) Cucullanus (C.) pinnai pinnai
Lophiosilurus alexandri Steindachner Nematoda gen. sp.
Family Trichomycteridae
Pygidium brasiliensis Reinhardt Ascarididae gen. sp.
Procamallanus (S.) pexatus
Trichomycterus piurae (Eigenmann) Procamallanus (S.) hilarii
Order Synbranchiformes
Family Synbranchidae
Synbranchus marmoratus Bloch Eustrongylides ignotus
Eustrongylides sp.
Nematoda gen. sp.
Order Tetraodontiformes
Family Balistidae
Balistes capriscus Gmelin Contracaecum sp.
Dichelyne sp.
Raphidascaris sp.
Terranova sp.
Balistes vetula Linnaeus Anisakis sp.
Contracaecum sp.
Dichelyne travassosi
Dichelyne sp.
Hysterothylacium sp.
Family Tetraodontidae
Sphoeroides testudineus (Linnaeus) Capillaria (N.) carioca
Capillostrogyloides sentinosa
Cucullanus dodsworthi
Class Chondrichthyes
Order Carcharhiniformes
Family Carcharhinidae
Carcharhinus brachyurus (Günther) Contracaecum sp.
Carcharhinus signatus (Poey) Anisakis sp.
Contracaecum sp.
Galeocerdo cuvieri (Péron & Lesueur) Nematoda gen. sp.
Family Scyliorhinidae
Scyliorhinus haeckelii (Miranda Ribeiro) Contracaecum sp.
Raphidascaris sp.
Family Sphyrnidae
Sphyrna zygaena (Linnaeus) Contracaecum sp.
Parascarophis sphyrnae
Family Triakidae
Galeorhinus galeus (Linnaeus) Anisakis sp.
Pseudoterranova sp.
Contracaecum sp.
Pseudoterranova sp.
Raphidascaris sp.
Mustelus canis (Mitchill) Anisakis sp.
Contracaecum sp.
Pseudoterranova sp.
Mustelus schmitti Springer Contracaecum sp.
Pseudoterranova sp.
“Cação Panan” Terranova rochalimai
Order Hexanchiformes
Family Hexanchidae
Heptranchias perlo (Bonnaterre) Anisakis sp.
Contracaecum sp.
Order Rajiformes
Family Potamotrygonidae
Paratrygon sp. Eustrongylides ignotus
Eustrongylides sp.
Potamotrygon hystrix (Müller & Henle) Echinocephalus daileyi
Potamotrygon falkneri Castex & Maciel Brevimulticaecum sp.
Brevimulticaecum sp. of Moravec et al. (1997)
Cucullanus (C.) sp.
Cucullanus sp.
Echinocephalus sp.
Spinitectus sp.
Potamotrygon motoro (Müller & Henle) Ascarididae gen. sp.
Brevimulticaecum regoi
Brevimulticaecum sp.
Echinocephalus daileyi
Procamallanus (P.) peraccuratus
Procamallanus (S.) inopinatus
Nematoda gen. sp.
Family Rajidae
Atlantoraja castelnaui (Miranda Ribeiro) Procamallanus (S.) pereirai
Raphidascaris sp.
Dipturus trachydermus (Krefft & Stehmann) Anisakis sp.
Contracaecum sp.
Raphidascaris sp.
Psammobatis extent (Garman) Pseudanisakis sulamericana
Rioraja agassizii (Müller & Henle) Pseudanisakis sulamericana
Family Rhinobatidae
Order Squaliformes
Family Squalidae
Squalus megalops (Macleay) Anisakis sp.
Pseudoterranova sp.
Order Squatiniformes
Family Squatinidae
Squatina sp. Anisakis sp.
Contracaecum sp.
Squatina squatina (Linnaeus) Pulchrascaris caballeroi
Terranova trichiuri
In recent years, parasites have been recognized as an important component of global biodiversity. Given the inte-
gral roles played by parasites in natural ecosystems, identifying hotspots of high parasite diversity, as well as areas
of relatively low parasite diversity, is crucial for a complete understanding of the functioning of the biosphere
(Luque & Poulin 2007). According Luque & Poulin (2007), nematode species represent 13% and 27% of the total
number of host-parasite associations in Latin American and the Caribbean marine and freshwater fishes, respec-
tively. In Brazil, a ‘Megadiversity Country’ with the highest fish biodiversity of the planet, only 17.3% of the
known fish species have parasitological records. Consequently, we are in agreement with the statement of Moravec
(1998) that our current knowledge of nematodes parasitizing Neotropical fishes is too rudimentary to permit
answers even to basic questions concerning their biodiversity, biology and ecology, never mind other issues relat-
ing to veterinary medicine and public health. The nematode fauna, therefore, remains an underestimated compo-
nent of Neotropical fish parasite biodiversity.
However, there are some general and incipient patterns in the distribution of nematode species in Brazilian
fishes. For example, the number of nematode species parasitizing freshwater fishes is higher than occurs in marine
and brackish fish species, and some fish groups, such as the Perciformes and Siluriformes, have a greater number
of host-parasite associations. However, these differences may not reflect true biological patterns, as they may only
be an echo of study effort and local research priorities. As only a small portion of the potential hosts have been
examined for parasitic nematodes, further studies are needed to evaluate the possible implications of the faunal
composition of this group.
With regard to the distribution of nematode taxa, the greatest number of host-parasite associations involves
ascaridoids, with approximately 51% of the records in this study. The significant presence of anisakid nematodes is
of importance, because of possible consequences in terms of public health and fish marketing. Anisakids utilize
fish as definitive, intermediate or paratenic hosts. The third larval stage of some of those occurring as adults in
higher vertebrates can cause a disease referred to as anisakiasis, a typical food-borne parasitic zoonosis (Limbery &
Cheah 2007), in man when raw or poorly cooked fish are consumed. Although human anisakiasis in Brazil remains
unreported, the presence of anisakid larvae in several Brazilian commercial fish species at a high prevalence and
abundance, plus an increase in the habit of eating raw fish, are indicative of a potential health problem in this coun-
try (Tavares & Luque 2006).
Acknowledgements
The authors are very grateful to the Fundação Carlos Chagas Filho de Amparo à Pesquisa do Estado do Rio de
Janeiro (FAPERJ) and the Instituto Oswaldo Cruz for financial support. José L. Luque was supported by a research
fellowship from Conselho Nacional de Desenvolvimento Científico e Tecnológico do Brasil (CNPq); Fabiano M.
Vieira was supported by a Postdoctoral fellowship from CNPq (Conselho Nacional de Pesquisa e Desenvolvimento
Tecnológico, Brasil); and Julio C. Aguiar was supported by an MSc Fellowship from the Coordenação de Aperfei-
çoamento do Pessoal de Ensino Superior (CAPES), Brazil.
References
Abdallah, V.D., Azevedo, R.K. & Luque, J.L. (2004) Metazoários parasitos dos lambaris Astyanax bimaculatus (Linnaeus, 1758),
Astyanax parahybae Eigenmann, 1908 e Oligosarcus hepsetus (Cuvier, 1829) (Osteichthyes, Characidae), do Rio Guandu, Estado
do Rio de Janeiro, Brasil. Revista Brasileira de Parasitologia Veterinária, 13, 57–63.
Abdallah, V.D., Azevedo, R.K. & Luque, J.L. (2005) Ecologia da comunidade de metazoários parasitos do sairú Cyphocharax gilbert
(Quoy e Gaimard, 1824) (Characiformes, Curimatidae) do Rio Guandu, Estado do Rio de Janeiro, Brasil. Revista Brasileira de
Parasitologia Veterinária, 14, 154–159.
Abdallah, V.D., Azevedo, R.K. & Luque, J.L. (2006) Ecologia da comunidade de metazoários parasitos do tamboatá Hoplosternum lit-
torale (Hancock, 1828) (Siluriformes, Callichthyidae) do rio Guandu, Estado do Rio de Janeiro, Brasil. Acta Scientiarum, 28,
413–419.
Abdallah, V.D., Luque, J.L., Alves, D.R. & Paraguassú, A.R. (2002) Aspectos quantitativos das infrapopulações de metazoários parasi-
tos da cavalinha, Scomber japonicus (Osteichthyes, Scombridae), do litoral do Estado do Rio de Janeiro. Revista da Universidade
Rural, Série Ciências da Vida, 22, 103–107.
Moreira, L.H.A., Yamada, F.H., Ceschini, T.L., Takemoto, R.M. & Pavanelli, G.C. (2010) The influence of parasitism on the relative
condition factor (Kn) of Metynnis lippincottianus (Characidae) from two aquatic environments: the upper Parana river floodplain
and Corvo and Guairaca rivers, Brazil. Acta Scientiarum Biological Sciences, 32, 83–86.
Moreira, N.I.B., Oliveira, C.L. & Costa, H.M.A. (1991) New helminth parasite of fish: S. freitasi sp. n. (Nematoda, Camallanidae).
Memórias do Instituto Oswaldo Cruz, 86, 429–431.
Moreira, N.I.B., Oliveira, C.L. & Costa, H.M.A. (1994) Spirocamallanus inopinatus (Travassos, Artigas & Pereira, 1928) e Spiroca-
mallanus saofranciscensis sp. n. (Nematoda, Camallanidae) em peixes da represa de Três Marias. Arquivo Brasileiro de Medicina
Veterinaria e Zootecnia, 46, 485–500.
Moreira, N.I.B., Rocha, G.N. & Costa, H.M.A. (2000) A new nematode species (Seuratoidea, Cucullanidae) parasitizing
Parauchenipterus striatulus (Steindachner, 1876) (Pisces, Auchenipteridae) in Brazil. Memórias do Instituto Oswaldo Cruz, 95,
39–41.
Moreira, S.T., Ito, K.F., Takemoto, R.M. & Pavanelli, G.C. (2005) Ecological aspects of the Iheringichthys labrosus (Lütken, 1874)
(Siluriformes, Pimelodidae) in reservoirs of Paraná river basin and upper Paraná river floodplain, Brazil. Acta Scientiarum, Bio-
logical Sciences, 27, 317–322.
Motta, C.S. & Gomes, D.C. (1968) Sobre uma nova espécie do gênero Porrocaecum Railliet & Henry, 1912 (Nematoda, Anisakidae).
Atas da Sociedade de Biologia do Rio de Janeiro, 11, 163–164.
Muniz-Pereira, L.C., Vieira, F.M. & Luque, J.L. (2009). Checklist of helminth parasites of threatened vertebrate species from Brazil.
Zootaxa, 2123, 1–45.
Nelson, J.S. (2006) Fishes of the world. John Wiley and Sons, New York, 601 pp.
Padovani, R.E.S., Knoff, M., São-Clemente, S.C., Mesquita, E.F.M., Jesus, E.F.O. & Gomes, D.C. (2005) The effect of in vitro gamma
radiation on Anisakis sp. larvae collected from the pink cusk-eel, Genypterus brasiliensis Regan, 1903. Revista Brasileira de Ciên-
cia Veterinária, 12, 137–141.
Paraguassú, A.R, Alves, D.R. & Luque, J.L. (2005) Metazoários parasitos do acará Geophagus brasiliensis (Quoy & Gaimard, 1824)
(Osteichthyes, Cichlidae) do reservatório de Lajes, estado do Rio de Janeiro, Brasil. Revista Brasileira de Parasitologia Veter-
inária, 14, 35–39.
Paraguassú, A.R. & Luque, J.L. (2006) Ecologia das comunidades de metazoários parasitos de seis espécies de peixes do reservatório
de lajes, estado do Rio de Janeiro, Brasil. Revista Brasileira de Parasitologia Veterinária, 16, 121–128.
Paraguassú, A.R., Luque, J.L. & Alves, D.R. (2000) Aspectos quantitativos do parasitismo por larvas de anisakídeos (Nematoda,
Ascaridoidea, Anisakidae) no pargo, Pagrus pagrus (Linaneus, 1758) (Osteichthyes, Sparidae) do litoral do Estado do Rio de
Janeiro, Brasil. Contribuições Avulsas Sobre a Historia Natural do Brasil, 24, 1–8.
Paraguassú, A.R., Luque, J.L. & Alves, D.R. (2002) Community ecology of metazoan parasites of red porgy Pagrus pagrus (Osteich-
thyes, Sparidae) from the coastal zone of the state of Rio de Janeiro, Brazil. Acta Scientiarum, 24, 461–467.
Parra, J.E.P., Brandao, D.A. & Ceccarelli, P.S. (1997) Identificação e prevalência de nematóides do pacu Piaractus mesopotamicus
(Holmberg, 1887), da Estação de Piscicultura do CEPTA, Pirassununga, SP, Brasil. Ciência Rural, 27, 291–295.
Pavanelli, G.C., Machado, M.H. & Takemoto, R.M. (1997) Fauna helmintológica de peixes do rio Paraná, região de Porto Rico, Paraná.
In: Vazzoler, A.E.M., Agostinho, A.A. & Hahn, N.S. (Eds.), A planície de inundação do alto rio Paraná: Aspectos físicos, biológi-
cos e socioeconômicos. Editora da Universidade Estadual de Maringá, Maringá, pp. 307–329.
Pavanelli, G.C., Machado, M.H., Takemoto, R.M., Guidelli, G.M. & Lizama, M.A.P. (2004) Helminth fauna of fishes: diversity and
ecological aspects. In: Thomaz, S.M., Agostinho, A.A. & Hahn, N.S. (Eds.), The upper Paraná river and its floodplain: Physical
aspects, ecology and conservation. Backhuis Publishers, Leiden, pp. 309–329.
Pereira, C. (1935) Ascaridida e Spirurata parasitos de peixes do nordeste brasileiro. Archivos do Instituto Biológico São Paulo, 6, 53–
62.
Pereira, C. (1938) Travnema travnema n. gen. n. sp. (Nematoda, Oxyuridae) parasito de Curimata elegans (Pisces, Characinidae) no
nordeste brasileiro. In: Livro Jubilar do Professor Lauro Travassos. Tipografia do Instituto Oswaldo Cruz, Rio de Janeiro, Brasil,
pp. 318–385.
Pereira, C. & Vaz, Z. (1933) Nematóides parasitos de Cynolebias bellotti (Pisces, Cyprinodonta). Revista de Biologia e Hygiene, 4, 59–
61.