Checklist Of Acanthocephala Associated With The Fishes Of Brazil

Abstract

Twenty-three genera comprising thirty-four named and thirteen undetermined species of acanthocephalans are associated with one hundred and nineteen marine, brackish and freshwater fish species from Brazil. These are listed, with information on their hosts, habitat, distribution and records. A key to these parasites at the generic level is provided.

Full text

Accepted by S. Monks: 15 Aug. 2008; published: 21 Nov. 2008 1
ZOOTAXA
ISSN 1175-5326 (print edition)
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Copyright © 2008 · Magnolia Press
Zootaxa 1938: 1–22 (2008)
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Checklist of Acanthocephala associated with the fishes of Brazil
CLÁUDIA P. SANTOS1, DAVID I. GIBSON2, LUIZ E. R. TAVARES3 & JOSÉ L. LUQUE3
1Laboratório de Avaliação e Promoção da Saúde Ambiental, Instituto Oswaldo Cruz, Fundação Oswaldo Cruz, Av. Brasil 4365, Rio
de Janeiro, 21045–900, Brazil. E-mail: cpsantos@ioc.fiocruz.br
2Department of Zoology, Natural History Museum, Cromwell Road, London SW7 5BD, UK
3Departamento de Parasitologia Animal, Universidade Federal Rural do Rio de Janeiro, Caixa Postal 74508, Seropédica, RJ, 23851–
970, Brazil
Abstract
Twenty-three genera comprising thirty-four named and thirteen undetermined species of acanthocephalans are associated
with one hundred and nineteen marine, brackish and freshwater fish species from Brazil. These are listed, with informa-
tion on their hosts, habitat, distribution and records. A key to these parasites at the generic level is provided.
Key words: Acanthocephala, biodiversity, checklist, freshwater and marine fishes, key, Neotropics, Brazil
Introduction
Studies on the phylum Acanthocephala in Brazil began with the naturalist Johann Natterer, who arrived in
Brazil on the 5th of November of 1817. He was among the royal attendants accompanying the Austrian Arch-
duchess Leopoldina, who came to marry the future Emperor of Brazil, D. Pedro I (Straube, 2000). Natterer
participated in several expeditions, crossing the country from Rio de Janeiro to the Amazon, during a period
of 18 years (1817–1835). The material which he collected was brought to the Brazilian museum in Vienna
(actually the Naturhistorische Museum). Among this material were helminth parasites, including some
acanthocephalans which were later studied by Karl Moritz Diesing (1851, 1856), who described several spe-
cies, two of which were subsequently redescribed by Kritscher (1957, 1976) based on the original material.
In 1913 the Brazilian researcher Lauro Travassos started his helminthological studies with work on
acanthocephalans from birds, and in 1923 he described Echinorhynchus jucundus Travassos, 1923 from the
fish Piaractus brachypomus (Cuvier, 1818). During this period, Travassos (1917, 1926) carried out a taxo-
nomic review of the acanthocephalans from Brazil. Subsequently, our knowledge of the Brazilian acantho-
cephalan fauna was significantly increased by the work of Domingos Machado Filho, a student of Travassos,
who described numerous new genera and species (Machado Filho, 1941, 1948, 1951, 1954, 1959a, b). Among
the researchers who have contributed to this fauna in recent years, the works of Vernon Thatcher (1991, 2006)
are the most notable, as he listed the parasite species, including acanthocephalans, from fishes in the Amazon
River basin, with keys, descriptions and accounts of their biology.
Here we provide a checklist of the acanthocephalan species associated with marine and freshwater fishes
from Brazil. 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.

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Material and methods
The bibliographic review of the species of helminth parasites reported from fishes in Brazilian waters was
based on information collected from two sources. Firstly, searches of the Host-Parasite Catalogue of the Natu-
ral History Museum in London for 1920 to 1997 and, for the years 1988–2003, the Host-Parasite Database, a
partial implementation of which is now available on-line (Gibson et al., 2005). Secondly, searches of the
Zoological Record and CAB Abstract databases were made up to February 2008. In addition to these data-
bases various other on-line searches were undertaken. Other works checked included published reports and
papers based on original records. Original data from our own research are also included.
The checklist follows the parasite classification and systematic arrangement of Amin (1985, 1987, 2002).
The Acanthocephala are arranged according to the class, order and family, within which the species are pre-
sented in alphabetical order. The host species are given followed by their predominant habitat (marine—
MAR, freshwater—FW or brackish water—BW), localities and records (in chronological sequence). In
addition, the checklist includes acanthocephalans recorded only to generic, familial or class level (undeter-
mined species).
Acanthocephalan species names have been up-dated to agree with the recent literature, but inclusion in the
parasite 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) and then in
alphabetical order at the generic and species levels; the names used follow FishBase (Froese & Pauly 2008).
Results
In the present paper twenty-three genera comprising thirty-four named and thirteen undetermined species of
acanthocephalans parasitising Brazilian fishes are listed. These worms were recorded from one hundred and
nineteen marine, brackish and freshwater fish species.
Key to the genera of Acanthocephala from Brazilian fishes
1 Proboscis receptacle double-walled...............................................................Class Palaeacanthocephala (2)
- Proboscis receptacle absent or single-walled............................................................................................ (7)
2 Adult worms in intestine of fishes and amphibians........................................... Order Echinorhynchida (3)
- Larval worms encysted in tissues of fish; adults in birds and mammals........................................................
..........................................................................................Order Polymorphida; Family Polymorphidae (9)
3 Cement glands 2 ................................................................................................. Family Diplosentidae (12)
- Cement glands more than 2 ..................................................................................................................... (4)
4 Trunk unarmed or with minute spines...................................................................................................... (5)
- Trunk armed with strong spines or plaques.............................................................................................. (6)
5 Cement glands 4 ....................................................... Family Cavisomidae; Paracavisoma Kritscher, 1957
- Cement glands 6–8 ................................................................................................................................. (13)
6 Trunk spined in single undivided anterior region............. Family Illiosentidae; Dollfusentis Golvan, 1969
- Trunk spined differently ............................................................................ Family Rhadinorhynchidae (15)
7 Proboscis claviform, with numerous longitudinal rows of hooks; cement glands separate ..........................
Class Polyacanthocephala; Order Polyacanthorhynchida; Family Polyacanthorhynchidae; Polyacantho-
rhynchus Travassos, 1926

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- Proboscis usually small, with few radial rows of hooks; cement gland single..Class Eoacanthocephala (8)
8 Trunk armed............................................................ Order Gyracanthocephala; Family Quadrigyridae (17)
-Trunk unarmed.......................................... Order Neoechinorhynchida; Family Neoechinorhynchidae (19)
9 Anterior part of trunk bulbous................................................................................................................ (10)
- Anterior part of trunk not bulbous.......................................................................................................... (11)
10 Trunk spines end at same level................................................................................ Bolbosoma Porta, 1908
- Trunk spines extend more on ventral side............................................................. Corynosoma Lühe, 1904
11 Trunk spines arranged in narrow band.................................................. Hexaglandula Petrotschenko, 1950
- Trunk spines not arranged in narrow band ........................................................... Polymorphus Lühe, 1911
12 Proboscis short, globular; hooks vary in size and shape.................................................................................
.................................................................................... Amapacanthus Salgado-Maldonado & Santos, 2000
- Proboscis long, claviform; hooks similar ........................ Golvanorhynchus Noronha, Fabio & Pinto, 1978
13 Proboscis long, with gradual transition from basal to apical hooks............ Family Echinorhynchidae (14)
- Proboscis short, with abrupt transition from basal to apical hooks................................................................
....................................................................... Family Arhythmacanthidae; Heterosentis Van Cleave, 1931
14 Trunk rotund; testes spherical, parallel or diagonal........................................ Brasacanthus Thatcher, 2001
- Trunk elongate; testes tandem ........................................................ Echinorhynchus Zoega in Müller, 1776
15 Trunk with cuticular combs on ventral surface.............................................. Serrasentis Van Cleave, 1923
- Trunk without cuticular combs............................................................................................................... (16)
16 Proboscis long, frequently with basal hooks larger than others .................... Rhadinorhynchus Lühe, 1911
- Proboscis short, without large hooks at base.............................................. Gorgorhynchus Chandler, 1934
17 Trunk with 15 circular rows of spines different in size .............................. Machadosentis Noronha, 1992
- Trunk with 4–12 circular rows of similar spines.................................................................................... (18)
18 Proboscis hooks in 6–12 oblique rows ................................................. Palliolisentis Machado Filho, 1960
- Proboscis hooks in 3–4 circular rows.......................................................... Quadrigyrus Van Cleave, 1920
19 Proboscis hooks in 3 circular rows......................................................................................................... (20)
- Proboscis hooks in 8 diagonal rows of 8 hooks each ........................................... Floridosentis Ward, 1953
20 Proboscis hooks in 3 circular rows of 6 hooks each............................................................................... (21)
- Proboscis hooks in 3 circular rows of more than 6 hooks each.............................................................. (22)
21 Muscular sling surrounds proboscis receptacle ......................... Gorytocephalus Nickol & Thatcher, 1971
- Muscular sling around proboscis receptable absent................. Neoechinorhynchus Stiles & Hassall, 1905
22 Proboscis hooks in 3 circular rows of 12 hooks each.................................. Gracilisentis Van Cleave, 1919
- Proboscis hooks in 3 circular rows of 8 hooks each.................................. Octospiniferoides Bullock, 1957
Parasite-Host List
Class Palaeacanthocephala Meyer, 1931
Order Echinorhynchida Southwell & MacFie, 1925
Family Arhythmacanthidae Van Cleave, 1931
Heterosentis sp.—adult
Hosts and habitat: Pinguipes brasilianus (MAR), Pseudopercis numida (MAR)
Locality: Off Cabo Frio - Rio de Janeiro
References: Luque et al. (2008)

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Family Cavisomidae Meyer, 1932
Paracavisoma impudicum (Diesing, 1851)—adult
Host and habitat: Oxydoras kneri (FW), Oxydoras niger (FW)
Locality: Upper Parana River/Paraná, Mato Grosso
References: Diesing (1851), Travassos et al. (1928), Kritischer (1957), Golvan (1969), Eiras et al.
(1995), Rêgo and Vicente (1988), Thatcher (1991, 2006)
Family Diplosentidae Meyer, 1932
Amapacanthus amazonicus Salgado-Maldonado & Santos, 2000—adult
Hosts and habitat: Anableps microlepis (MAR), Sciades passany (MAR)
Locality: Off Maraca Island
References: Salgado-Maldonado and Santos (2000)
Golvanorhynchus golvani Noronha, Fabio & Magalhaes, 1978—adult
Host and habitat: Scomber colias (MAR)
Locality: Off Rio de Janeiro
References: Noronha et al. (1978)
Family Echinorhynchidae Cobbold, 1876
Brasacanthus sphoeroides Thatcher, 2001—adult
Host and habitat: Sphoeroides greeleyi (MAR)
Locality: Off Parana
References: Thatcher (2001)
Echinorhynchus briconi Machado Filho, 1959 (syn. Metechinorhynchus briconi)—adult
Host and habitat: Brycon hilarii (FW)
Locality: São Paulo
References: Machado Filho (1959c), Golvan (1969), Thatcher (1991, 2006)
Echinorhynchus gomesi Machado-Filho, 1948—adult
Host and habitat: Prochilodus nigricans (FW)
Locality: Mato Grosso
References: Machado Filho (1948)
Echinorhynchus gracilis Machado Filho, 1948—adult
Host and habitat: Brycon hilarii (FW)
Locality: São Paulo
References: Machado Filho (1948)
Echinorhynchus jucundus Travassos, 1923—adult
Hosts and habitat: Piaractus brachypomus (syn. Colossoma bidens) (FW), Piaractus mesopotamicus
(FW)
Locality: São João, Mato Grosso
References: Travassos (1923), Machado Filho (1948), Ferraz de Lima et al. (1990), Thatcher (1991,
2006)
Echinorhynchus paranensis Machado-Filho, 1959—adult
Host and habitat: Triportheus paranensis (FW)
Locality: Mato Grosso
References: Machado Filho (1959c), Thatcher (1991)

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Echinorhynchus salobrensis Machado Filho, 1948—adult
Host and habitat: Mylossoma duriventre (syn. M. paraguayensis) (FW)
Locality: Salobra, Mato Grosso
References: Machado Filho (1948), Thatcher (1991)
Echinorhynchus sp.—adult
Hosts and habitat: Brycon cephalus (FW), Schizodon borellii (FW)
Locality: Amazon River Basin, Upper Parana River/Paraná
References: Machado et al. (1994, 1995, 1996), Andrade et al. (2001), Pavanelli et al. (2004)
Echinorhynchidae gen. sp.—adult
Host and habitat: Kyphosus sp. (MAR)
Locality: Off Rio de Janeiro
References: Gomes et al. (1974)
Family Illiosentidae Golvan, 1960
Dollfusentis chandleri Golvan, 1969—adult
Hosts and habitat: Archosargus rhomboidalis (MAR), Eucinostomus argenteus (MAR), Haemulon sciu-
rus (MAR), Haemulon steindachneri (MAR), Micropogonias furnieri (MAR), Orthopristis ruber
(MAR), Umbrina coroides (MAR)
Locality: Off Rio de Janeiro
References: Kohn and Macedo (1984), Noronha et al. (1986), Vicente et al. (1989), Luque et al. (1996a,
b), Alves and Luque (1999, 2000, 2001a, b)
Family Rhadinorhynchidae Travassos, 1923
Gorgorhynchus trachinotus Noronha, Vicente, Pinto & Fábio, 1986—adult
Host and habitat: Trachinotus goodei (MAR)
Locality: Off Rio de Janeiro
Reference: Noronha et al. (1986)
Gorgorhynchus sp.—adult and larva
Hosts and habitat: Dactylopterus volitans (MAR—adult), Dipturus trachyderma (MAR—larva), Pin-
guipes brasilianus (MAR—adult) new record, Pseudopercis numida (MAR—adult), Sphyrna zyga-
ena (MAR—larva), Squatina sp. (MAR—larva)
Localities: Off Rio de Janeiro, off Rio Grande do Sul, off Paraná, off Santa Catarina
References: Knoff et al. (2001), Cordeiro and Luque (2005), Luque et al. (2008)
Rhadinorhynchus plagioscionis Thatcher, 1980—adult
Host and habitat: Plagioscion squamosissimus (FW)
Locality: Amazon River basin
Reference: Thatcher (1980, 1991)
Rhadinorhynchus pristis (Rudolphi, 1802)—adult
Hosts and habitat: Coryphaena hippurus (MAR), Balistes vetula (MAR), Euthynnus alletteratus
(MAR), Katsuwonus pelamis (MAR), Auxis thazard (MAR), Scomber japonicus (MAR)
Locality: Off Rio de Janeiro
References: Rudolphi (1819), Abdallah et al. (2002), Mogrovejo and Santos (2002), Alves et al. (2003,
2005), Alves and Luque (2006), Oliva et al. (2008)
Rhadinorhynchus sp.—adult
Hosts and habitat: Dactylopterus volitans (MAR), Merluccius hubbsi (MAR) (new record),

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Paralonchurus brasiliensis (MAR)
Locality: Off Rio de Janeiro
References: Ribeiro et al. (2002), Luque et al. (2003), Cordeiro and Luque (2005)
Serrasentis sp.—larva
Hosts and habitat: Balistes capriscus (MAR), Haemulon steindachneri (MAR), Oligoplites palometa
(MAR), Orthopristis ruber (MAR), Paralonchurus brasiliensis (MAR), Parona signata (new
record) (MAR)
Locality: Off Rio de Janeiro
References: Luque et al. (1995, 1996a, b, 2003), Takemoto et al. (1996), Luque and Poulin (2004), Alves
et al. (2005)
Order Polymorphida Petrochenko, 1956
Family Polymorphidae Meyer, 1931
Bolbosoma sp.—larva
Host and habitat: Scomber japonicus (MAR), Merluccius hubbsi (new host record) (MAR)
Locality: Off Rio de Janeiro
Reference: Rêgo and Santos (1983)
Corynosoma australe Johnston, 1937—larva
Hosts and habitat: Centropomus undecimalis (MAR), Cephalopholis fulva (MAR), Cynoscion guatu-
cupa (MAR), Diapterus rhombeus (MAR), Galeorhinus galeus (MAR), Genypterus brasiliensis
(MAR), Hexanchus griseus (MAR), Menticirrhus americanus (MAR), Micropogonias furnieri
(MAR), Mullus argentinae (MAR), Paralichthys isosceles (MAR), Pomatomus saltatrix (MAR),
Prionotus punctatus (MAR), Scomber japonicus (MAR), Squalus megalops (MAR), Squatina sp.
(MAR)
Localities: Off Rio de Janeiro, off Rio Grande do Sul, off Paraná
References: Pereira and Neves (1993), Luque and Chaves (1999), Alves and Luque (1999, 2000, 2001a,
b), Knoff et al. (2001), Abdallah et al. (2002), Luque et al. (2002), Alves et al. (2002a, b, 2003),
Sabas and Luque (2003), Tavares and Luque (2004b), Luque and Poulin (2004), Timi et al. (2005),
Bicudo et al. (2005), Oliva et al. (2008)
Corynosoma sp.—larva
Hosts and habitat: Euthynnus alletteratus (MAR), Galeorhinus galeus (MAR), Hexanchus griseus
(MAR), Merluccius hubbsi (MAR—new record), Micropogonias furnieri (MAR), Pagrus pagrus
(MAR), Pomatomus saltatrix (MAR), Sarda sarda (MAR), Scomber japonicus (MAR), Squalus
megalops (MAR), Squatina sp. (MAR), Umbrina canosai (MAR)
Localities: Off Rio de Janeiro, off Rio Grande do Sul, off Paraná, off Santa Catarina
References: Rêgo et al. (1983), Cezar and Amato (1998), Luque and Chaves (1999), Alves and Luque
(2000, 2001a, b, 2006), Knoff et al. (2001), Paraguassú et al. (2002), Luque and Poulin (2004),
Oliva et al. (2008)
Hexaglandula mutabilis (Rudolphi, 1819) (syn. Polymorphus mutabilis)—larva
Hosts and habitat: Atherina sp. (BW/MAR), Eucinostomus harengulus (BW/MAR), Sphoeroides
testudineus (BW/MAR), Strongylura marinus (BW/MAR)
Locality: Angra dos Reis, Rio de Janeiro
References: Travassos (1926)

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Polymorphus sp.—larva
Hosts and habitats: Anchoa tricolor (MAR), Astronotus ocellatus (FW), Genidens barbus (MAR),
Genypterus brasiliensis (MAR), Geophagus brasiliensis (FW), Oligosarcus hepsetus (FW), Pagrus
pagrus (MAR), Priacanthus arenatus (MAR), Trichiurus lepturus (MAR), Urophycis mystacea
(MAR)
Localities: Off Rio de Janeiro, Rio Guandú – Rio de Janeiro
References: Silva et al. (2000a, b), Tavares et al. (2001), Alves et al. (2002a, b, c), Paraguassú et al.
(2002), Luque and Poulin (2004), Abdallah et al. (2004), Tavares and Luque (2004a), Tavares et al.
(2004a, 2005), Azevedo et al. (2006, 2007)
Polymorphidae gen. sp.—larva
Host and habitat: Lophius gastrophysus (new record) (MAR)
Locality: Off Rio de Janeiro
Class Polyacanthocephala Amin, 1987
Order Polyacanthorhynchida Amin, 1987
Family Polyacanthorhynchidae Golvan, 1956
Polyacanthorhynchus macrorhynchus (Diesing, 1856)—adult
Host and habitat: Arapaima gigas (FW)
Locality: Amazon River basin
References: Diesing (1856), Travassos (1919), Travassos et al. (1928), Machado Filho (1947), Golvan
(1956), Travassos and Freitas (1964), Thatcher (1991, 2006)
Polyacanthorhynchus rhopalorhynchus (Diesing, 1851)—adult
Host and habitat: Arapaima gigas (FW)
Localities: Amazon River basin, Mato Grosso
References: Diesing (1851), Baylis (1927), Travassos (1919), Travassos et al. (1928), Machado Filho
(1947), Thatcher (1991, 2006)
Class Eoacanthocephala
Order Gyracanthocephala Van Cleave, 1936
Family Quadrigyridae Van Cleave, 1920
Machadosentis travassosi Noronha, 1992—adult
Host and habitat: Gymnothorax ocellatus (MAR)
Locality: Off Bahia
Reference: Noronha (1992)
Palliolisentis quinqueungulis Machado-Filho, 1960—adult
Hosts and habitat: Triportheus angulatus (FW), Triportheus paranensis (FW)
Locality: Salobra, Mato Grosso
References: Machado Filho (1960), Thatcher (1991)

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Quadrigyrus brasiliensis Machado Filho, 1941—adult
Hosts and habitat: Hoplerythrinus unitaeniatus (FW), Hoplias malabaricus (FW).
Locality: Salobra, Mato Grosso, Upper Parana River/Paraná
References: Machado Filho (1941), Thatcher (1991), Pavanelli et al. (2004)
Quadrigyrus machadoi Fabio, 1983—larva and adult
Hosts and habitat: Cichla monoculus (FW—larva), Hemisorubim platyrhynchos (FW—larva), Gymno-
tus sp. (FW—adult), Hoplias malabaricus (FW—adult)
Localities: Parana River/Paraná, São Paulo, Rio de Janeiro, Rio Baía, Mato Grosso do Sul
References: Fabio (1983), Weiblen and Brandão (1992), Pavanelli et al. ( 2001, 2002, 2004), Machado et
al. (2000), Guidelli et al. (2003), Isaac et al. (2004), Rosim et al. (2005)
Quadrigyrus torquatus Van Cleave, 1920— adult
Hosts and habitat: Acestrorhynchus lacustris (FW), Hoplias malabaricus (FW), Leporinus lacustris
(FW)
Localities: Upper Parana River/Paraná, Rio de Janeiro
References: Fabio (1983), Thatcher (1991) Pavanelli et al. (2004), Carvalho et al. (2003), Guidelli et al.
(2006)
Quadrigyrus sp.—adult
Hosts and habitat: Astyanax altiparanae (FW), Iheringichthys labrosus (FW), Pachyurus squamipennis
(FW), Prochilodus lineatus (FW)
Localities: Upper Parana River/Paraná São Francisco River, Mato Grosso, Paraná
References: Pavanelli et al. (2004), Lizama et al. (2005), Moreira et al. (2005),Takemoto et al. (2005)
Order Neoechinorhynchida Southwell & MacFie, 1925
Family Neoechinorhynchidae Ward, 1917
Floridosentis mugilis (Machado Filho, 1951) (syn. Atactorhynchus mugils Machado Filho, 1951)—adult
Host and habitat: Mugil platanus (MAR)
Locality: Off Rio de Janeiro
References: Knoff et al. (1997)
Gracilisentis variabilis (Diesing, 1851) (syns Echinorhynchus variabilis, Octospinifer variabilis)
Hosts and habitat: Achirus achirus (FW), Achirus lineatus (FW), Achirus sp. (FW), Glyptoperichthys lit-
uratus (FW), Hoplias malabaricus (FW), Hypostomus auroguttatus (FW), Hypostomus nem-
atopterus (FW), Hypostomus plecostomus (FW), Prochilodus lineatus (FW), Pleuronectes sp. (FW)
Localities: Pirassununga, São Paulo, Amazon River Basin
References: Diesing (1851, 1856), Travassos et al. (1928), Travassos & Kohn (1965), Kritscher (1976),
Thatcher (1991, 2006)
Gorytocephalus spectabilis (Machado Filho, 1959) (syn. Neoechinorhynchus spectabilis Machado Filho,
1959)—adult
Hosts and habitat: Cyphocharax gilbert (FW), Steindachnerina elegans (FW)
Locality: Emas, Pirassununga, São Paulo
References: Machado Filho (1959a), Travassos and Kohn (1965), Kohn and Fernandes (1987), Thatcher
(1991, 2006)
Gorytocephalus elongorchis Thatcher, 1979—adult
Host and habitat: Hypostomus carinatus (FW)
Locality: Amazon River Basin

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References: Thatcher (1979, 1991, 2006)
Neoechinorhynchus (Neoechinorhynchus) buttnerae Golvan, 1956—adult
Host and habitat: Colossoma macropomum (FW)
Locality: Amazon River Basin
References: Golvan (1956), Thatcher (1991), Ferraz de Lima et al. (1990), Malta et al. (2001), Fischer et
al. (2003)
Neoechinorhynchus (Neoechinorhynchus) curemai Noronha, 1973—adult
Hosts and habitats: Mugil curema (BW), Prochilodus lineatus (FW)
Localities: River Amazon, Pirassununga—São Paulo, Volta Grande reservoir, Minas Gerais, Mato
Grosso, Upper Parana River, Potengi River—Rio Grande do Norte
References: Noronha (1973, 1984), Kohn et al. (1985), Kohn and Fernandes (1987), Brasil-Sato and Pav-
anelli (1999), Ranzani-Paiva et al. (2000), Martins et al. (2000, 2001), Fortes et al. (2000), Santos et
al. (2003, 2005), Pavanelli et al. (2004), Lizama et al. (2005, 2006)
Neoechinorhynchus (Neoechinorhynchus) macronucleatus Machado Filho, 1954—adult
Hosts and habitat: Hoplias malabaricus (FW), Lycengraulis sp. (FW)
Localities: Juparanã lagoon, Espírito Santo, Campos, Rio de Janeiro
References: Machado Filho (1954), Fabio (1983)
Neoechinorhynchus (Neoechinorhynchus) paraguayensis Machado Filho, 1959—adult
Hosts and habitat: Cichlasoma monoculus (FW), Geophagus brasiliensis (FW), Hoplias malabaricus
(FW)
Locality: Guandu-Açú River—Rio de Janeiro
References: Machado Filho (1959b), Nickol and Padilha (1979), Azevedo et al. (2006), Thatcher (1991,
2006)
Neoechinorhynchus (Neoechinorhynchus) pimelodi Brasil-Sato & Pavanelli, 1998—adult
Hosts and habitat: Franciscodoras marmoratus (FW), Pimelodus maculatus (FW)
Locality: São Francisco River, Três Marias—Mato Grosso
References: Brasil-Sato and Pavanelli (1998, 1999), Brasil-Sato (2003), Santos and Brasil-Sato (2004)
Neoechinorhynchus (Neoechinorhynchus) pterodoridis Thatcher, 1981
Host and habitat: Pterodoras granulosus (FW)
Locality: Amazon River Basin
References: Thatcher (1981, 1991)
Neoechinorhynchus sp.—larva and adult
Hosts and habitat: Conorynchus conirostris (FW—larval), Corydoras julii (FW—adult), Pseudoplatys-
toma fasciatum (FW)
Locality: Aquidauana River, River São Francisco, aquarium in Germany
References: Moravec et al. (1999), Brasil-Sato and Santos (2005), Campos et al. (2008)
Octospiniferoides incognita Schmidt & Huggins, 1973—adult
Host and habitat: Leporinus lacustris (FW), Schizodon borellii (FW), Schizodon fasciatus (FW)
Locality: Upper Parana River/Paraná, Rondonia State
References: Machado et al. (1994, 1996), Thatcher (1998), Pavanelli et al. (2004), Guidelli et al. (2006)
Acanthocephala fam. gen. sp. (unidentified specimens)—larva and adult
Hosts and habitats: Ageneiosus ucayalensis (FW), Brycon hilarii (FW), Cichlasoma paranaense (FW),
Chloroscombrus chrysurus (MAR), Cichlasoma paranaense (FW), Cynoscion sp. (MAR), Hemi-
caranx amblyrhynchus (MAR), Hoplerythrinus unitaeniatus (FW), Hoplias malabaricus (FW), Lep-
orinus lacustris (FW), Menticirrus americanus (MAR), Mugil curema (MAR), Mugil platanus
(MAR), Paralonchurus brasiliensis (MAR), Piaractus mesopotamicus (FW), Pimelodus maculatus
(FW), Pseudoplatystoma corruscans (FW), Pterodoras granulosus (FW), Pygocentrus nattereri

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10 · Zootaxa 1938 © 2008 Magnolia Press
(FW), Rachycentron canadum (MAR), Rhaphiodon vulpinus (FW), Schizodon altoparanae (FW),
Schizodon nasutus (FW), Serrasalmus marginatus (FW), Serrasalmus spilopleura (FW)
Localities: Rivers Aquidauana, Miranda, Paraguay—Mato Grosso do Sul, Cuiabá and Manso Rivers—
Mato Grosso, Upper Paraná river-Paraná, off São Paulo, off Espírito Santo, Guandú River – Rio de
Janeiro and off Marambaia Island – Rio de Janeiro
References: Travassos and Freitas (1940), Travassos et al. (1963, 1964), Kohn and Fernandes (1987),
Pavanelli et al. (2004), Adriano et al. (2005), Santos et al. (2007).
Host-parasite list
Hosts Parasite species
Class Actinopterygii (ray-finned fishes)
Order Anguilliformes
Family Muraenidae
Gymnothorax ocellatus Agassiz Machadosentis travassosi
Order Atheriniformes
Family Atherinopsidae
Atherina sp. Hexaglandula mutabilis
Order Cyprinodontiformes
Family Anablepidae
Anableps microlepis Müller & Troschel Amapacanthus amazonicus
Order Beloniformes
Family Belonidae
Strongylura marina (Walbaum) Hexaglandula mutabilis
Order Clupeiformes
Family Engraulidae
Anchoa tricolour (Spix & Agassiz) Polymorphus sp.
Lycengraulis sp. Neoechinorhynchus macronucleatus
Order Characiformes
Family Acestrorhynchidae
Acestrorhynchus lacustris (Lütken) Quadrigyrus torquatus
Family Anostomidae
Leporinus lacustris Amaral Campos Octospiniferoides incognita
Quadrigyrus torquatus
Acanthocephala fam. gen. sp.
Schizodon altoparanae Garavello & Britski Acanthocephala fam. gen. sp.
Schizodon borellii (Boulenger) Echinorhynchus sp.
Octospiniferoides incognita
Schizodon fasciatus Spix & Agassiz Octospiniferoides incognita
Schizodon nasutus Kner Acanthocephala fam. gen. sp.
Family Characidae
Astyanax altiparanae Garutti & Britski Quadrigyrus sp.
Brycon cephalus (Günther) Echinorhynchus sp.
Brycon hilarii (Valenciennes) Echinorhynchus briconi
Echinorhynchus gracilis

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ACANTHOCEPHALA FROM FISHES OF BRAZIL
Acanthocephala fam. gen. sp.
Colossoma macropomum (Cuvier) Neoechinorhynchus buttnerae
Mylossoma duriventre (Cuvier) Echinorhynchus salobrensis
Oligosarcus hepsetus (Cuvier) Polymorphus sp.
Piaractus brachypomus (Cuvier) Echinorhynchus jucundus
Piaractus mesopotamicus (Holmberg) Echinorhynchus jucundus
Acanthocephala fam.gen. sp.
Pygocentrus nattereri Kner Acanthocephala fam. gen. sp.
Serrasalmus marginatus Valenciennes Acanthocephala fam. gen. sp.
Serrasalmus spilopleura Kner Acanthocephala fam. gen. sp.
Triportheus angulatus (Spix & Agassiz) Palliolisentis quinqueungulis
Triportheus paranensis (Günther) Echinorhynchus paranensis
Palliolisentis quinqueungulis
Family Cynodontidae
Rhaphiodon vulpinus Spix & Agassiz Acanthocephala fam. gen. sp.
Family Curimatidae
Cyphocharax gilbert (Quoy & Gaimard) Gorytocephalus spectabilis
Steindachnerina elegans (Steindachner) Gorytocephalus spectabilis
Family Erythrinidae
Hoplerythrinus unitaeniatus (Agassiz) Quadrigyrus brasiliensis
Acanthocephala fam. gen. sp.
Hoplias malabaricus (Bloch) Gracilisentis variabilis
Quadrigyrus brasiliensis
Quadrigyrus machadoi
Quadrigyrus torquatus
Neoechinorhynchus macronucleatus
Neoechinorhynchus paraguayensis
Acanthocephala fam. gen. sp.
Family Prochilodontidae
Prochilodus lineatus (Valenciennes) Gracilisentis variabilis
Neoechinorhynchus curemai
Quadrigyrus sp.
Prochilodus nigricans Agassiz Echinorhynchus gomesi
Order Gadiformes
Family Merlucciidae
Merluccius hubbsi Marini Bolbosoma sp.
Corynosoma sp.
Rhadinorhynchus sp.
Family Phycidae
Urophycis mystacea Ribeiro Polymorphus sp.
Order Gymnotiformes
Family Gymnotidae
Gymnotus sp. Quadrigyrus machadoi
Order Lophiiformes
Family Lophiidae
Lophius gastrophysus Ribeiro Polymorphidae gen. sp.

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12 · Zootaxa 1938 © 2008 Magnolia Press
Order Ophidiiformes
Family Ophidiidae
Genypterus brasiliensis Regan Corynosoma australe
Polymorphus sp.
Order Perciformes
Family Carangidae
Chloroscombrus chrysurus (Linnaeus) Acanthocephala fam. gen. sp.
Hemicaranx amblyrhynchus (Cuvier) Acanthocephala fam. gen. sp.
Oligoplites palometa (Cuvier) Serrasentis sp.
Parona signata (Jenyns) Serrasentis sp.
Trachinotus goodie Jordan & Evermann Gorgorhynchus trachinotus
Family Centropomidae
Centropomus undecimalis (Bloch) Corynosoma australe
Family Cichlidae
Astronotus ocellatus (Agassiz) Polymorphus sp.
Cichla monoculus Spix & Agassiz Quadrigyrus machadoi
Neoechinorhynchus paraguayensis
Cichlasoma paranaense Kullander Acanthocephala fam. gen. sp.
Geophagus brasiliensis (Quoy & Gaimard) Neoechinorhynchus paraguayensis
Polymorphus sp.
Family Coryphaenidae
Coryphaena hippurus Linnaeus Rhadinorhynchus pristis
Family Gerreidae
Diapterus rhombeus (Cuvier) Corynosoma australe
Eucinostomus argenteus Baird & Girard Dollfusentis chandleri
Eucinostomus harengulus Goode & Bean Hexaglandula mutabilis
Family Haemulidae
Haemulon sciurus (Shaw) Dollfusentis chandleri
Haemulon steindachneri (Jordan & Gilbert) Dollfusentis chandleri
Serrasentis sp.
Orthopristis ruber (Cuvier) Dollfusentis chandleri
Serrasentis sp.
Family Kyphosidae
Kyphosus sp. Echinorhynchidae gen. sp.
Family Mugilidae
Mugil platanus Günther Floridosentis mugilis
Acanthocephala fam. gen. sp.
Mugil curema Valenciennes Neoechinorhynchus curemai
Acanthocephala fam. gen. sp.
Family Mugiloididae
Pinguipes brasilianus Cuvier Gorgorhynchus sp.
Heterosentis sp.
Pseudopercis numida Ribeiro Gorgorhynchus sp.
Heterosentis sp.
Family Mullidae

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ACANTHOCEPHALA FROM FISHES OF BRAZIL
Mullus argentinae Hubbs & Marini Corynosoma australe
Family Pomatomidae
Pomatomus saltatrix (Linnaeus) Corynosoma australe
Corynosoma sp.
Family Priacanthidae
Priacanthus arenatus Cuvier Polymorphus sp.
Family Rachycentridae
Rachycentron canadum (Linnaeus) Acanthocephala fam. gen. sp.
Family Sciaenidae
Cynoscion guatucupa (Cuvier) Corynosoma australe
Cynoscion sp. Acanthocephala fam. gen. sp.
Menticirrhus americanus (Linnaeus) Corynosoma australe
Acanthocephala fam. gen. sp.
Micropogonias furnieri (Desmarest) Corynosoma australe
Corynosoma sp.
Dollfusentis chandleri
Pachyurus squamipennis Agassiz Quadrigyrus sp.
Paralonchurus brasiliensis (Steindachner) Rhadinorhynchus sp.
Serrasentis sp.
Acanthocephala fam. gen. sp.
Plagioscion squamosissimus (Heckel) Rhadinorhynchus plagioscionis
Umbrina canosai Berg Corynosoma sp.
Umbrina coroides (Cuvier) Dollfusentis chandleri
Family Scombridae
Auxis thazard (Lacepède) Rhadinorhynchus pristis
Euthynnus alletteratus (Rafinesque) Corynosoma sp.
Rhadinorhynchus pristis
Katsuwonus pelamis (Linnaeus) Rhadinorhynchus pristis
Sarda sarda (Bloch) Corynosoma sp.
Scomber colias Gmelin Golvanorhynchus golvani
Scomber japonicus Houttuyn Bolbosoma sp.
Corynosoma australe
Corynosoma sp.
Golvanorhynchus golvani
Rhadinorhynchus pristis
Family Serranidae
Cephalopholis fulva (Linnaeus) Corynosoma australe
Family Sparidae
Archosargus rhomboidalis (Linnaeus) Dollfusentis chandleri
Pagrus pagrus (Linnaeus) Corynosoma sp.
Polymorphus sp.
Family Trichiuridae
Trichiurus lepturus Linnaeus Polymorphus sp.
Order Pleuronectiformes
Family Achiridae
Achirus achirus (Linnaeus) Gracilisentis variabilis

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14 · Zootaxa 1938 © 2008 Magnolia Press
Achirus lineatus (Linnaeus) Gracilisentis variabilis
Achirus sp. Gracilisentis variabilis
Family Paralichthyidae
Paralichthys isosceles Jordan Corynosoma australe
Family Pleuronectidae
Pleuronectes sp. Gracilisentis variabilis
Order Scorpaeniformes
Family Dactylopteridae
Dactylopterus volitans (Linnaeus) Gorgorhynchus sp.
Rhadinorhynchus sp.
Family Triglidae
Prionotus punctatus (Bloch) Corynosoma australe
Order Siluriformes
Family Ariidae
Sciades passany Valenciennes Amapacanthus amazonicus
Genidens barbus (Lacepède) Polymorphus sp.
Family Auchenipteridae
Ageneiosus ucayalensis Castelnau Acanthocephala fam. gen. sp.
Family Callichthyidae
Corydoras julii Steindachner Neoechinorhynchus sp.
Family Doradidae
Franciscodoras marmoratus (Reinhardt) Neoechinorhynchus pimelodi
Oxydoras kneri Bleeker Paracavisoma impudica
Oxydoras niger (Valenciennes) Paracavisoma impudica
Pterodoras granulosus (valenciennes) Acanthocephala fam. gen. sp.
Neoechinorhynchus pterodoridis
Family Loricariidae
Glyptoperichthys lituratus (Kner) Gracilisentis variabilis
Hypostomus auroguttatus Kner Gracilisentis variabilis
Hypostomus carinatus (Steindachner) Gorytocephalus elongorchis
Hypostomus nematopterus Isbrücker & Nijssen Gracilisentis variabilis
Hypostomus plecostomus (Linnaeus) Gracilisentis variabilis
Family Pimelodidae
Hemisorubim platyrhynchos (Valenciennes) Quadrigyrus machadoi
Iheringichthys labrosus (Lütken) Quadrigyrus sp.
Pimelodus maculatus (Lacepède) Acanthocephala fam. gen. sp.
Neoechinorhynchus pimelodi
Pseudoplatystoma corruscans (Spix & Agassiz) Acanthocephala fam. gen. sp
Pseudoplatystoma fasciatum (Linnaeus, 1766) Neoechinorhynchus sp.
Order Siluriformes (species incertae sedis)
Conorynchus conirostris (Valenciennes) Neoechinorhynchus sp.
Order Osteoglossiformes
Family Arapaimatidae
Arapaima gigas (Schinz) Polyacanthorhynchus macrorhynchus
Polyacanthorhynchus rhopalorhynchus

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ACANTHOCEPHALA FROM FISHES OF BRAZIL
Discussion
Acanthocephalan parasites of fishes come in two forms, adults from the intestine and larvae (post-cysta-
canths) from the tissues. The latter are usually members of the family Polymorphidae and utilise fishes
(mainly marine) as a second intermediate host in the life-cycle. Brazilian records belong to species of Coryno-
soma, Bolbosoma, Hexaglandula and Polymorphus, the adults of which are found in aquatic mammals and
birds. The majority of Brazilian records from fishes are of adult specimens, although ingested cystacanths and
juvenile worms are also found in the intestine. All acanthocephalans utilise arthropods as intermediate hosts;
in the case of species from fishes these are normally crustaceans. The host-specificity of the worms to their
fish hosts varies considerably; in some cases they are found in a single host and in others in a group of unre-
lated hosts, which perhaps reflects the host’s diet.
The phylum Acanthocephala is a relatively poorly known helminth group in Neotropical fishes. Accord-
ing to Luque & Poulin (2007), they represent fewer than 5% of the host-parasites associations recorded in
Order Tetraodontiformes
Family Balistidae
Balistes capriscus Gmelin Serrasentis sp.
Balistes vetula Linnaeus Rhadinorhynchus pristis
Family Tetraodontidae
Sphoeroides greeleyi Gilbert Brasacanthus sphoeroides
Sphoeroides testudineus (Linnaeus) Hexaglandula mutabilis
Class Chondrichthyes (cartilaginous fishes)
Order Carcharhiniformes
Family Sphyrnidae
Sphyrna zygaena (Linnaeus) Gorgorhynchus sp.
Family Triakidae
Galeorhinus galeus (Linnaeus) Corynosoma australe
Corynosoma sp.
Order Hexanchiformes
Family Hexanchidae
Hexanchus griseus (Bonnaterre) Corynosoma australe
Corynosoma sp.
Order Rajiformes
Family Rajidae
Dipturus trachyderma Krefft & Stehmann Gorgorhynchus sp.
Order Squaliformes
Family Squalidae
Squalus megalops (Macleay) Corynosoma australe
Corynosoma sp.
Order Squatiniformes
Family Squatinidae
Squatina sp. Corynosoma australe
Corynosoma sp.
Gorgorhynchus sp.

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16 · Zootaxa 1938 © 2008 Magnolia Press
freshwater and marine fishes from this region. This checklist shows that from 123 fish hosts species reported,
116 are ray-finned fishes and seven are cartilaginous, and the most parasitized fish family is the Characidae
with 15 acanthocephalan species, followed by the Sciaenidae with nine.
The composition of the known acanthocephalan fauna shows that the Neoechinorhynchidae is the most
represented family, comprising some 12 named species parasitic in freshwater, brackish water and marine
fishes. Some genera are clearly specific to South American fishes, whereas others are cosmopolitan. The host
with greatest number of reported acanthocephalan species is the marine fish Haemulon sciurus, and the acan-
thocephalan recorded fom the greatest number of hosts is Corynosoma australe, a larval form.
Some apparent Brazilian records are not included in the lists above. Although previously listed (Thatcher,
1991, 2006) as occurring in the region, Palliolisentis ornatus Machado-Filho, 1960, fromTriportheus paran-
ensis (FW), was originally described from the Paraguay River, Paraguay. We can find no definite Brazilian
record. Similarly, Megapriapus ungriai (Gracia-Rodrigo, 1960) from Potamotrygon hystrix, and Quadrigyrus
torquatus Van Cleave, 1920 from Astyanax bimaculatus (FW), Crenicichla geayi (FW), Gephyrocharax
valencia (FW) and Symbranchus marmoratus (FW), have apparently been reported only from Venezuela and
are consequently omitted from our list.
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).
Luiz E. R. Tavares was supported by a Junior Posdoctoral Fellowship from CNPq.
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