Amazon Fish Parasites Vol I PDF

ABLA Vol.
1 – Thatcher: Amazon Fish Parasites 1
Aquatic Biodiversity in Latin America
Biodiversidad Acuática en América Latina
Volume 1
Amazon Fish Parasites
(Second edition)
Volumen 1
Parásitos de Peces Amazónicos
(Segunda edición)
2 ABLA Vol. 1 – Thatcher: Parásitos de Peces Amazónicos
BIODIVERSIDAD ACUÁTICA EN AMÉRICA LATINA
Volumen 1
PARÁSITOS DE PECES
AMAZÓNICOS
(Segunda edición)
por
Vernon E. Thatcher
Lector Científico a los Editores:

František Moravec
Editores de la Serie:
Joachim Adis, Jorge R. Arias,
Guillermo Rueda-Delgado & Karl Matthias Wantzen
Sofia–Moscow
2006
ABLA Vol. 1 – Thatcher: Amazon Fish Parasites 3
AQUATIC BIODIVERSITY IN LATIN AMERICA
Volume 1
AMAZON FISH
PARASITES
(Second edition)
by
Vernon E. Thatcher
Scientific Reader to the Editors:

František Moravec
Series Editors:
Joachim Adis, Jorge R. Arias,
Sofia–Moscow
2006

Series Editors/Editores de la Serie:

Joachim Adis, Jorge R. Arias, Guillermo Rueda-Delgado & Karl Matthias Wantzen
VOLUME 1. VOLUMEN 1.
AMAZON FISH PARASITES PARÁSITOS DE PECES AMAZÓNICOS
(Second edition) (Segunda edición)
by por
Vernon E. Thatcher Vernon E. Thatcher
Scientific Reader to the Editors: Lector Científico a los Editores:
František Moravec František Moravec
Front cover:
Nerocila armata (Cymothoidae, Isopoda) and Amazon River near Manaus, Brazil
(photos: V.E. Thatcher, W.J. Junk; design: Zheko Aleksiev & Elke Bustorf)
Aquatic Biodiversity in Latin America Vol. 1

ISSN 1312-7276
First published 2006

ISBN-10: 954-642-258-4
ISBN-13: 978-954-642-258-3
© PENSOFT Publishers
All rights reserved. No part of this publication may be reproduced, stored in a retrieval system or
transmitted in any form by any means, electronic, mechanical, photocopying, recording or
otherwise, without the prior written permission of the copyright owner.
Pensoft Publishers
Geo Milev Str. 13a, Sofia 1111, Bulgaria
pensoft@mbox.infotel.bg
www.pensoft.net
Printed in Bulgaria, March 2006

Series Editors: Joachim Adis, Jorge R. Arias,

Biodiversity is a key word in science and global management schemes, however very few
people are able to identify the species and their ecology that make up “biodiversity”. For
many years, researchers and students from numerous countries complain about the lack of
“tools” to identify aquatic invertebrates from Latin America. Keys found in accepted
entomological textbooks are mostly highly limited, superficial and rarely cover Neotropical
biota in sufficient detail. On the other hand, specialized information on taxonomy or ecology
is scattered throughout the literature in many single publications.
An international team of editors have combined their efforts with Pensoft Publishers
to launch a new major series on the Aquatic Biodiversity of Latin America (ABLA). Their
goal was to find experts who combine the current state of knowledge in taxonomy and
ecology, in order to produce a concise and affordable handbook for each group. About 15
separate monographs, written by reference scientists from various countries will offer a
new, unrivalled view on the aquatic fauna of South America. Information on the ecology
and status of the taxa (written in English) is combined with illustrated identification keys to
families and genera, in both English and Spanish. The series is aimed at zoologists, ecologists,
hydrobiologists, biogeographers, conservationists and students interested in aquatic
biodiversity. The series will be an essential tool for any biological library.
Volume 1: Amazon Fish Parasites (Second edition) by Vernon E. Thatcher.

Pensoft Publishers, Sofia-Moscow, ISBN-10: 954-642-258-4, ISBN-13: 978-954-642-
258-3, 165x240 mm, 508 pp., including 194 plates of figures in line drawings and
photos (16 plates in color); publication date: March 2006.
This book covers the following Phyla and Classes: Protozoa (Myxozoa, Sporozoa),
Plathelminthes (Trematoda, Monogenoidea, Cestoda), Nematoda, Acanthocephala,
Crustacea (Copepoda, Branchiura, Isopoda), Hirudinea and Pentastomida that are known
to parasitize Amazonian fishes. In addition to the keys and checklists, each chapter gives
information on the morphology, life cycles, pathology, prevention, treatment, collection
and methodology, as well as general reference data on each taxon. An alphabetic table of
host fishes with their respective parasites is provided. The book will prove to be useful for
parasitologists, ichthyologists, aquaculturists, pisciculturists and all those interested in
Neotropical fish parasites.
ADDRESSES
Author:
PROF. DR. VERNON E. THATCHER DR. KARL MATTHIAS WANTZEN
Universidade Federal do Paraná Institute of Limnology
Faculdade de Ciências Biológicas University of Konstanz
Departamento de Zoologia Postfach M 659
Caixa Postal 19020 78457 Konstanz, Germany
81531-990 Curitiba, Paraná, Brazil e-mail: matthias.wantzen@uni-konstanz.de
e-mail: thatcher@uol.com.br,
thatcher@ufpr.br Scientific Reader to the Editors:
DR. FRANTIŠEC MORAVEC
Series Editors: Institute of Parasitology
PROF. DR. JOACHIM ADIS Academy of Sciences of the Czech Republic
Max-Planck-Institute for Limnology Branišovská 31
Tropical Ecology Working Group 37005 České Budějovice, Czech Republic
Postfach 165 e-mail: moravec@paru.cas.cz
24302 Plön, Germany
e-mail: adis@mpil-ploen.mpg.de
DR. JORGE R. ARIAS

5870 Colfax Avenue
Alexandria, Virginia 22311, USA
e-mail: jaria2@fairfaxcounty.gov
M.Sc. GUILLERMO RUEDA-DELGADO

Grupo de Investigación en Cuencas y
Humedales Tropicales GICHT
UNIMAG-INTROPIC
Universidad Jorge Tadeo Lozano
Laboratorio de Limnología
Carrera 4 No. 22-61
Bogotá, D.C.
Colombia
e-mail: guillermo.rueda@utadeo.edu.co
FOREWORD
Amazon Fish Parasites by Vernon E. Thatcher
It is an honor to provide this introductory statement to the 2nd edition of “Amazon Fish
Parasites” compiled by my dear friend, Dr. Vernon E. Thatcher. Vernon’s contributions to
parasitology span a period of more than five decades, during which time he has published
more than 160 scientific papers, most dealing with the parasites of Neotropical fishes. His
overall contributions include coverage of a wide variety of parasitic taxa, including the
Kinetoplasta (Protista), Digenea, Monogenoidea, Nematoda, Cestoda, Acanthocephala,
Copepoda, Isopoda and Diptera. On one occasion, I remember Vernon commenting that
a “true” parasitologist is one that was knowledgeable, competent, and sufficiently versed
concerning most (if not all) parasitic groups such that he/she could conduct significant
research on any one of them. Vernon has certainly lived up to this definition of a
parasitologist, and it is fitting that he be the person to compile the available information
about the parasites of Amazonian fishes.
Knowledge of the diversity of the parasites of Neotropical fishes, and those of the
Amazon in particular, has significantly increased since publication of the 1st edition of
“Amazon Fish Parasites” in 1991. Much of this new information is due to a large extent
to the more than 60 publications originating from Thatcher’s laboratory since 1991, as
well as works published by other investigators, who undoubtedly were encouraged to
conduct new investigations as a result of the publication of the 1st edition. The new
edition includes coverage of all parasites representing the major parasitic groups infesting
Amazonian fishes listed in the first edition, as well as that of species subsequently reported
in the literature. For each of the major parasitic groups, the 2nd edition includes sections
on Definition and Morphology, Life Cycle and Transmission, Pathology, Prevention and Treatment, as
well as Collection and Methods. These introductory sections are then followed with
identification, keys in English and Spanish, a checklist of species and numerous figures
and references pertaining to the respective groups. The checklist of species not only lists
all species of the taxon reported from the region, but includes diagnoses of families,
subfamilies and genera represented by the species in the list. Finally, the book ends with
a detailed list of hosts and their respective parasites.
Although I am not totally aware of the extensive literature of all parasite groups
represented in the Amazon, coverage of the Monogenoidea (my specialty) suggests that
records presented in the 2nd edition are complete and up-to-date. The organization of
the book and that of each chapter dealing with the individual parasite groups is extremely
conducive for use by other investigators. There is little doubt that the 2nd edition of
“Amazon Fish Parasites” will greatly assist future investigators with identification of
Amazonian fish parasites and provide a convenient means for surveying the literature for
a variety of studies including taxonomic, phylogenetic and ecologic investigations of the

parasites found in the region. Copies of the 2nd edition of “Amazon Fish Parasites” will
certainly find their way to the book shelves of many libraries and most workers dealing
with the parasites of fishes in the region.
DELANE C. KRITSKY, Ph.D.

Idaho State University
Pocatello, USA
PREFACE TO THE SECOND EDITION
The first edition of the book, Amazon Fish Parasites, was well received especially in
Europe and tropical American countries. It has proven to be useful to aquarists and
pisciculturists. The book went to press in 1990 and since that time, advances have been
made in several areas of fish parasite research. New genera and species of parasites have
been discovered and described. New finds have been reported in Monogenoidea,
Trematoda, Nematoda, Copepoda and Isopoda and these have been incorporated into the
present edition. Keys in both English and Spanish are provided to aid in the identification
of the parasites. The author hopes that the present edition will continue to be helpful to
those interested in tropical fishes and their parasites and stimulate further studies on these
fascinating animals.
VERNON E. THATCHER
Departamento de Zoologia
Universidade Federal do Paraná
Curitiba, Paraná, Brazil, August 2005
PREFACE TO THE FIRST EDITION
My first encounter with fish parasites occurred some forty years ago when I took a course
in the subject from the late Dr. Ivan Pratt at Oregon State University. Although I received
only one credit for the course (and had to work very hard for it), I gained a field of
interest that has lasted through the years. Fish parasites are so numerous and varied, and
have adapted to the parasitic way of life in so many ways, that they never cease to
fascinate.
I have had the opportunity to live and work in the American tropics for nearly thirty
years. My studies have frequently involved human and mammalian parasites, but I have
always found time to examine a few fish as well. I have collected and studied fish parasites
from the Grijalva River basin in Tabasco, Mexico, the Chagres River in Panama, the Cauca
River system in Colombia and the Guandu-Aço River in Rio de Janeiro State, Brazil. For
the past twelve years, I have been able to study fish parasites and the pathology they
provoke at the Brazilian National Institute of Amazonian Research (INPA) in Manaus,
Amazonas, Brazil. It has been possible to examine wild caught fish from various parts of
the Amazon region as well as captive fish held in the INPA pisciculture station.
The present book is an attempt to compile all available information on Amazon fish
parasites. For the sake of comparison, freshwater fish parasites from other parts of the
Neotropical Region have been included. Marine species have been omitted, however.
Every effort has been made to produce a work that is as complete as possible. Inevitably,
some references will have been overlooked. Even so, this book should serve as a basic
source of information and stimulate further research in the field.
VERNON E. THATCHER
Manaus, Amazonas, Brazil, February, 1990
ACKNOWLEDGMENTS
The author is grateful for the continued support of Profs. WOLFGANG J. JUNK and
JOACHIM ADIS of the Tropical Ecology Working Group, Max-Planck-Institute for
Limnology, Plön, Germany. Thanks also to Dr. DELANE C. KRITSKY, Idaho State
University, Pocatello, Idaho (Monogenoidea); Dr. OTTO SEY, University of Agricultural
Science, Keszthely, Hungary and Dr. FRANTIŠEK MORAVEC, Institute of Parasitology,
České Budějovice, Czech Republic, who not only aided in the publication of new species
but also granted permission to reproduce some of the figures used. The author is also
grateful for the collaboration of Dr. WALTER A. BOEGER and R. N. VIANNA, Department
of Zoology, Universidade Federal do Paraná, on the chapter concerning Monogenoidea,
and to Dr. JORGE R. ARIAS, Director of the West Nile Virus Program at the Fairfax
County Health Department, for checking the Spanish translations. Special thanks are due
to Dr. BEDSY DUTARY THATCHER for converting the text of the first edition of the book
to digital format.
VERNON E. THATCHER
Curitiba, Paraná, Brazil, August 2005
Table of Contents
1I NTRODUCTION
Cited and general references
19
22
2 P I. Definition andCmorphology
ROTOZOA AND ILIOPHORA
23
II. Life-cycle and transmission 24
III. Pathology 25
IV. Prevention and treatment 26
V. Collection and study methods 26
VI. Identification and keys 27
Key to the genera of Neotropical Myxosporida 28
Clave para los géneros de Myxosporida Neotropicales 28
Key to Neotropical fish ciliophorans 28
Clave para los cilioforos de peces Neotropicales 29
VII. Checklist of Myxosporida from Neotropical
freshwater fishes 29
Checklist of Ciliophora from Neotropical freshwater fishes 31
VIII. Plates of Protozoa and Ciliophora (Figs. 2–1 to 2–44) 33
IX. Cited and general references 40
3 MI. Definition and morphology

ONOGENOIDEA
42
II. Life cycle and transmission 44
III. Pathology 44
Key to higher taxa of freshwater Amazonian Monogenoidea 50
Key to genera of freshwater Amazonian Gyrodactylidae 51

Key to genera of freshwater Amazonian Dactylogyrinea 51
Clave para las taxas superiores de Monogenoidea
Neotropicales de agua dulce 55
Clave para los géneros de Gyrodactylidae
Clave para los Dactylogyrinea Neotropicales de agua dulce 56
VII. Checklist of Monogenoidea from Amazonian
freshwater fishes and diagnoses 60
VIII. Plates of Monogenoidea (Figs. 3-1 to 3-64) 92
4 T I. Definition (Dand morphology

REMATODA IGENEA )
117
III. Pathology 119
Key to the Neotropical freshwater fish trematodes 123
Clave para identificación de Tremátodos, parásitos de
peces Neotropicales 127
VII. Checklist of Trematoda (Digenea) from
Neotropical freshwater fishes 131
VIII. Plates of Trematoda (Digenea) (Figs. 4-1 to 4-99) 154
5 C I. Definition and morphology

ESTODARIA AND C ESTODA
206
III. Pathology 207
Key to Cestodaria and Cestoda of Neotropical

Clave para Cestodaria y Cestoda de peces
VII. Checklist of Cotyloda (= Cestodaria) and Eucestoda
(= Cestoda) from Neotropical freshwater fishes 213
VIII. Plates of Cestoda and Cestodaria (Figs. 5-1 to 5-44) 222
6 NI. Definition and morphology

EMATODA
234
III. Pathology 235
Key to the superfamilies of Nematoda from Amazonian
Key to the Ascaridoidea of Amazonian freshwater fishes 238
Key to the Oxyuroidea (Pharyngodonidae) of Amazonian
Key to the Camallanoidea (Camallanidae) of Amazonian
Key to the Cosmocercoidea of Amazonian freshwater fishes 240
Key to the Dracunculoidea of Amazonian freshwater fishes 240
Key to the Seuratoidea of Amazonian freshwater fishes 241
Key to the Trichinelloidea (Capillariidae) of Amazonian
Key to the Gnathostomatoidea (Gnathostomatidae) of
Amazonian freshwater fishes 242
Key to the Thelazioidea (Rhabdochonidae) of Amazonian
Key to the Physalopteroidea (Physalopteridae) of Amazonian
Key to the Habronematoidea of Amazonian freshwater fishes 242

Clave para superfamílias de Nematoda de peces
Amazonicos de agua dulce 242
Clave para las Ascaridoidea de peces Amazonicos de
agua dulce 243
Clave para las Oxyuroidea (Pharyngodonidae) de peces
Clave para las Camallanoidea (Camallanidae) de
peces Amazonicos de agua dulce 244
Clave para las Cosmocercoidea de peces Amazonicos de
agua dulce 245
Clave para las Dracunculoidea de peces Amazonicos de
agua dulce 245
Clave para las Seuratoidea de peces Amazonicos de
agua dulce 246
Clave para las Trichinelloidea (Capillariidae) de
Clave para las Gnathostomatoidea (Gnathostomatidae) de
Clave para las Thelazioidea (Rhabdochonidae) de peces
Clave para las Physalopteroidea (Physalopteridae) de peces
Clave para las Habronematoidea de peces Amazonicos de
agua dulce 247
VII. Checklist of Nematoda from Amazonian fishes 248
VIII. Plates of Nematoda (Figs. 6-1 to 6-98) 265
7 A I. Definition and morphology

CANTHOCEPHALA
299
III. Pathology 301

VI. Identification and key 302
Key to Acanthocephala of Neotropical freshwater fishes 302
VI. Identificación y clave 303
Clave de identificación para Acanthocephala de peces
VII. Checklist of Acanthocephala from Neotropical freshwater fishes 305
VIII. Plates of Acanthocephala (Figs. 7-1 to 7-32) 311
8 C I. Definition and morphology

OPEPODA
326
Ergasilidae 326
Vaigamidae 328
Lernaeidae 329
Therodamasidae 329
III. Pathology 331
Key to Amazonian freshwater ergasiloid females 333
Key to the South American genera and species of
postmetamorphic female Lernaeidae 334
Clave para hembras Ergasiloides dulceacuícolas Amazonicas 335
Claves para los géneros y especies de Lernaeidae
hembras post-metamórficas de América del Sur 336
VII. Checklist of Copepoda from Amazonian freshwater fishes 337
VIII. Plates of Copepoda (Figs. 8-1 to 8-62) 346
9 B I. Definition and morphology

RANCHIURA
390
III. Pathology 391

Key to the Neotropical genera of Branchiura 392
Key to the Neotropical species of Argulus 392
Key to the Neotropical species of Dolops 393
Clave para los géneros Neotropicales de Branchiura 394
Clave para las especies Neotropicales de Argulus 394
Clave para las especies Neotropicales de Dolops 395
VII. Checklist of Branchiura from Neotropical freshwater fishes 397
VIII. Plates of Branchiura (Figs. 9-1 to 9-31) 401
10 II. Definition and morphology

SOPODA
416
III. Pathology 417
Key to the Genera of Cymothoidae from Neotropical
Clave para los géneros de Cymothoidae de peces
VII. Checklist of Isopods from Neotropical freshwater fishes 422
VIII. Plates of Isopoda (Figs. 10-1 to 10-62) 427
11 H H IRUDINEA,
IRUDINEA
PENTASTOMIDA AND OTHER PATHOGENS
454
I. Definition and morphology 454
III. Pathology 455

VI. Identification and key 456
Key to leeches from Neotropical freshwater fishes 456
Clave para Sanguijuelas de peces Neotropicales de agua dulce 456
VII. Checklist of Hirudinea from Neotropical fishes 456
PENTASTOMIDA 458
I. Definition and morphology 458
III. Pathology 458
VI. Identification 459
VII. Checklist for Pentastomida 459
Other pathogens 460
VIII. Plates of Hirudinea and miscellaneous
pathological conditions (Figs. 11-1 to 11-7) 461
12 A DDENDUM: HOST–PARASITE TABLE 465
13 S UBJECT INDEX 497

1
INTRODUCTION
Fish parasitology and pathology are fields of growing importance in view of a worldwide
expansion of pisciculture efforts. It would seem to be inevitable that world populations
will come to depend more and more on artificially cultured fish as naturally occurring fish
populations become depleted. Fish parasitology and pathology are not really separate
areas of study since most fish parasites cause mild to severe pathological alterations in
their hosts. Although it would be possible to study fish parasites and disregard the
pathology they produce, it would not be feasible to delve into fish pathology (ichthyopa-
thology) while ignoring the parasites that cause most of the damage.
Although no universally applicable definition of parasitism has ever been devised,
for our purposes we can consider a parasite as an animal that lives on or in an animal of a
different species (host) at the expense of the latter. The host pays the bill by providing
sustenance to the parasite. During the evolution of such a relationship over long time
periods, the parasite gradually loses its ability to elaborate certain molecules that are
essential to its own metabolic well being. The true parasite therefore has a physiological
dependence on a particular host species and cannot survive without the association.
Ultimately then, parasitism would have to be defined in terms of the biochemical
dependency of the parasite on the host. Even though details of these associations at the
biochemical level are incomplete or lacking, we can surmise a great deal from differences
in degrees of host specificity. As it turns out, most parasites are host specific and some
can develop only on a single host species. Others are limited to host species of a single
genus while some can grow on host genera of only one family group. Even those parasites
that occur on a variety of hosts are usually found to have preferences. Parasites on the
preferred host generally are found in greater numbers, have individuals of larger size and
produce more eggs or larvae per parasite.
Pathology is literally the study of “pathos”. Pathos means suffering, especially that
brought on by disease, physical injury or nutritional deficiencies. Pathology is also the
resultant damage produced in an organism by these conditions. Ichthyopathology differs
considerably from human or mammalian pathology for several important reasons. For
one thing, we cannot ask a fish where it hurts so we have to depend on visible indications
such as alterations in cells and tissue (histopathology) and the presence and identification
of invading disease organisms. Also, the aquatic environment is very different from the
terrestrial one in terms of disease transmission. The human pathologist spends much time
diagnosing and characterizing diseases caused by microorganisms (viruses, bacteria and
fungi), those of a hereditary nature and those associated with ageing. He is little
concerned with parasites, at least in Northern temperate climes. The ichthyopathologist,

on the other hand, is more involved with parasitic and nutritional disorders. Microbial and
hereditary diseases have been little studied in Neotropical fish and sick animals are usually
removed from the environment by predators before they grow very old.
The following major groups of animal parasites invade Neotropical freshwater
fishes: Protozoa, Myxozoa (= Myxosporida), Ciliophora, Platyhelminthes (Monogenoidea,
Trematoda (Digenea), Cestodaria, Cestoda), Nematoda, Acanthocephala, Arthropoda
(Copepoda, Branchiura, Isopoda), Annelida (Hirudinea) and Pentastomida. The groups
listed here represent 14 Classes or Orders of eight or nine animal Phyla. Of the 14
groups, only five infect man and only seven invade other mammals.
All of these parasitic forms evolved from free-living, frequently predatory, ancestors.
With the exception of Myxosporida, Acanthocephala and Pentastomida, the free-living
relatives are still around and can be found in nearly any body of water. All of these
invertebrate animal groups had already evolved before the first vertebrates appeared on
the scene. First to show up were the fishes and they offered numerous new niches for the
invertebrate groups to exploit. Fish therefore have a greater quantity and variety of
parasites than any other class of vertebrates because they have lived for a longer time
period in close association with the greatest variety of invertebrate forms. Parasitism, as
such, did not arise as a result of the appearance of the first vertebrates, however. Small
invertebrates had long been invading larger ones and many of these were therefore pre-
adapted to parasitize vertebrates when the opportunity arose.
Parasites are usually divided into ecto and endoparasites depending on whether they
live on the surface or within their hosts. In the case of fish ectoparasites, they may live on
the external surface, within the mouth or on the gills. All other habitats within the fish are
considered to be interior. To these general categories a third should be added, namely,
blood and tissue parasites. For a parasite to penetrate the circulatory system or tissues of
a host represents an evolutionary advance and these sites must be considered more
“interior” than the lumen of the digestive tract and associated organs.
One of the first requisites of the successful parasite is its ability to remain in position
on or within its host and not be swept away by the current. This is particularly evident in the
case of ectoparasites living on the slippery skin of fast swimming fish. As every fisherman
knows, one good way to catch a fish is with a fishhook. Parasitic crustaceans (Copepoda,
Branchiura and Isopoda) have evolved claws that closely resemble fishhooks and some even
have barbed points. Other attachment devices found among fish parasites include entangling
threads, suckers, spines, clamps, tentacles and clasping or latching appendages.
Much of the histopathology observed in fish is a direct result of the physical damage
done to the tissues by the attachment devices of the parasites. Such damage includes
localized hemorrhaging, cellular enlargement (hypertrophy), increased cellular growth
(hyperplasia) and the transformation of epithelial cells to mucous cells (metaplasia).
Parasites can also damage host tissues by their physical presence (causing pressure
atrophy), by chemical secretions or excretions and by their mode of feeding. For example,
some monogenean worms individually cause little harm to the gills of their hosts, but
sometimes they are present in such large numbers that they impede the breathing function of
the lamellar surfaces. Other monogenean species actively feed on epithelial and blood cells, and
can destroy the gill lamellae and filaments (Fig. 3-63). Still others provoke tumoroid growths
near the site of attachment presumably by chemical excretions (Fig. 3–64).
Sick or parasitized fish can frequently be recognized as such by the observant
pisciculturist or aquarist because their behavior alters. Signs to watch for include: color
changes, failure to keep up with others in a school, erratic or circular swimming patterns,
alterations in swimming posture (i.e. head, tail or belly frequently uppermost) and surface
gasping. The latter often indicates a low level of dissolved oxygen in the water, but it can
also signal the presence of excessive numbers of gill parasites.
Treatment of parasitized fish can sometimes be accomplished but is not without its
inherent difficulties. Chemical medications can be mixed with pellet-type ration and
offered to the fish, but if they are already sick, they may not be eating. This method works
better as a control to prevent the build-up of excessive numbers of parasites in relatively
healthy fish. The most common technique for treating fish is the chemical “dip” where
the fish are removed from their tanks and placed in chemically altered water for a short
time, and then returned to their previous environment. This method can be quite
successful, especially against ectoparasites. It should be remembered, however, that
catching and transferring the fish introduces the element of stress which can be fatal to
animals that are already sick. Anti-parasite chemicals can also be introduced directly into
the stomach by tube, or injected into the muscles or circulatory systems by hypodermic
syringe. These methods work admirably at times, but again the factor of stress must be
taken into consideration. If, for example, we took a chicken and held it under water for
several minutes while administering some painful treatment, the stress would be
comparable to that of a fish taken from water.
Since the treatment of fish is admittedly difficult, it would be far better if we could
keep them from getting infected in the first place. In order to be able to do so, we should
have detailed information concerning parasite life-cycles, especially as regards their
transmission from one host to another. Unfortunately, few studies have been made on
parasite life-cycles in the Neotropics. In many cases, however, we can deduce the mode of
transmission from studies done on similar parasites elsewhere. Some parasitic organisms
invade fish by direct means. That is, as active larvae or adults they swim up to a fish and
attach or penetrate. Ciliophora, Copepoda, Branchiura, Isopoda and Hirudinea use this
approach. Others have larval forms that are eaten by the fish, sometimes within
intermediate hosts or attached to aquatic vegetation. Trematoda, Cestoda, Nematoda and
Acanthocephala gain entry into fish in this way. Specific suggestions for ways to break the
transmission cycles are to be found in the chapters that follow.
Original papers on Neotropical fish parasites are scattered through the scientific
journals of six countries and were published in five different languages. Until the first
edition of this book, no general reference work about these parasites had been available.
The existing books on fish parasites and pathology, such as: HOFFMAN (1967, 1999); KINNE
(1984); REICHENBACH-KLINKE & LANDOLT (1973); RIBELIN & MIGAKI (1975); and ROBERTS
(1978), provide useful information based on North American and European forms, but
they are of little help in identifying Neotropical species.
Each of the following chapters of this work deals with a major group of Neotropical
fish parasites. It will be noted that Chapter 3, Monogenoidea and Chapter 8, Copepoda are
longer than the others. This reflects a reality in that these parasites are more numerous and
varied on these fish than any of the others. They are also the forms that cause the most
problems for pisciculturists and aquarists. Each chapter contains the following topics:
I. Definition and morphology; II. Life-cycle and transmission; III. Pathology; IV.
Prevention and treatment; V. Collection and study methods; VI. Identification and keys
(English and Spanish); VII. Checklist of species; VIII. Figures and IX. References. The
keys facilitate identification to genus. After the genus of a parasite is determined, the
checklist should be consulted to find out what species is represented. The checklists are
arranged alphabetically and after each specific name, the host, country where found,
figure citation and measurements are given. If the user wishes to find out what other
parasites have been reported from the same host, he may consult the addendum, which is
a host-parasite table.
Cited and general references
HERWIG, N. (1979): Handbook of drugs and chemicals used in the treatment of fish diseases. -
Charles C. Thomas Publ., Springfield: 272 pp.
HOFFMAN, G. L. (1967): Parasites of North American freshwater fishes. - Univ. Calif. Press,
Berkeley: 486 pp.
HOFFMAN, G. L. (1999): Parasites of North American freshwater fishes. - 2nd. ed., Cornell Univ.
Press, Ithaca: 539 pp.
KINNE, O. (ed.) (1984): Disease of marine animals. Vol. 4, Part 1. Introduction, Pisces. - Biologische
Anstalt Helgoland, Hamburg: i–xv, 541 pp.
OLSEN, O. W. (1974): Animal parasites, their life cycles and ecology. - Univ. Park Press, Baltimore:
562 pp.
REICHENBACH-KLINKE, H. H. & M. LANDOLT (1973): Fish pathology. - TFH Press, Neptune City:
512 pp.
RIBELIN, W. E. & G. MIGAKI (eds.) (1975): The pathology of fishes. - Univ. Wisconsin Press,
Madison: 1004 pp.
ROBERTS, R.J. (ed.) (1978): Fish pathology. - Bailliere Tindall, London: 318 pp.
THATCHER, V. E. & J. BRITES NETO (1991): Diagnóstico, prevenção e tratamento das enfermidades
de peixes Neotropicais de água doce. - In: VAL, A. L., FLIGLIUOLO, R. & E. FELDBERG (eds.):
Bases científicas para estratégias de preservação e desenvolvimento da Amazônia: 339–371.
Fatos e perspectivas. Impressa CNPq INPA/UA, Manaus: 440 pp.
2
PROTOZOA AND CILIOPHORA
I. Definition and morphology
The Phylum Protozoa formerly included all small unicellular animals, but in recent
years there has been a tendency to regard some of the component groups as separate
phyla. After all, a Paramecium and a malaria parasite have little in common in terms of
either morphology or life-cycle. Representatives of the protozoan groups Mastigo-
phora, Myxosporida, Coccidida, and some genera of the Phylum Ciliophora (formerly
Class Ciliata) have been reported from Neotropical freshwater fishes and these forms
will be treated here.
Trypanosoma spp. are the only mastigophorans known from Neotropical freshwater
fishes. Species of this genus have a single flagellum arising from a posterior blepharoplast
which is fused along the length of the body to form an undulating membrane. Beyond the
front end, the same flagellum is free and whip-like. The body tapers towards both
extremities and the nucleus is usually near the middle or in the anterior half (Fig. 2–35).
Trypanosomes are found in the blood of their hosts.
The myxosporidans, especially species of the genera Myxobolus and Henneguya, are
among the most common and conspicuous of fish parasites. These invaders form spore
filled cysts in the host tissues that vary in size from less than one millimeter to more than
one centimeter. Such cysts are found in and on the gills, under the skin and within internal
organs, such as muscles, liver, spleen and intestinal wall. Subcutaneous cysts sometimes
cause such deformations in fish that they can be observed in nature (Figs. 2-40 & 41).
Small cysts, when numerous, give an organ like the liver a granular or white spotted
appearance. Identification of myxosporidan cysts is made by opening them with dissecting
needles and examining the contents under a microscope at 400x magnification. Myxospori-
dan cysts contain large numbers of vegetative (non-motile) spores composed of two shell-
like valves. Within each spore, from one to four cylindrical polar capsules may be seen and
each of these contains a spiral filament. From one to five nuclei may also be visible.
Coccidia are usually found in the intestinal wall, or in associated organs such as the
liver. Identification of Calyptospora sp. (Eimeriidae), the only coccidian presently known to
occur in Neotropical fishes, can be made under 400x magnification by finding oocysts
containing four spores, each of which has two sporozoites (Figs. 2-28 & 2-44 A-B).
Fish ciliophorans are usually found on the skin or gills, or within the intestinal
contents of their hosts. These animals are easily recognized at 100x magnification by the
presence of motile cilia which cover the external surfaces (Figs. 2-29 to 2-34). In living
specimens, it is often possible to see the large macronucleus. In stained preparations, a
smaller micronucleus near to, or attached to, the macronucleus may be discerned.
II. Life-cycle and transmission
Trypanosomes have been described from several siluriform fishes and a few characins from
Brazil, but their life-cycles have not been studied. We can presume, however, that they must
have a leech (Hirudinea) as intermediate host and vector as do the North American species.
Transmission to an uninfected fish is therefore by way of the bite of an infected leech.
Transmission of myxosporidans from one host to another is by means of the
bivalved spores. When these spores are ingested by the specific host fish, the polar
capsules discharge the filaments to entangle in the cells of the gut wall and this serves to
anchor the spore. After anchoring, the valves open and a single ameboid parasite (sporont)
leaves the spore and penetrates the intestinal wall. The sporont is carried throughout the
body in the circulatory system until the proper organ is reached. Myxosporidan species are
apparently quite specific both in host and site selection. After settling at the site, the
sporont (trophozoite) begins to produce numerous new spores internally by asexual
nuclear divisions (for details on sporozoan life-cycles; see HOFFMAN, 1999).
Although myxosporidan life-cycles would appear to involve only one host, there is
some evidence to suggest that at least some species need two animals. According to WOLF
(1982), the spores of Myxosoma (or Myxobolus) cerebralis must “age” for three months or so
before they become infective. The same author found that oligochaete worms, especially
those of the genus Tubifex, ingested the spores and became infective to experimental trout.
Some myxosporidans alter the behavior of their host fishes to the extent that they
fall easy prey to predators. This suggests that piscivorous animals may be important in
disseminating the spores and it may be that passage through the digestive system of
another animal changes the capsules and makes the spores infective. In the Cauca River
valley of Colombia, entire populations of Astyanax fasciatus periodically become
infected with a species of Henneguya. The parasites form large cysts under the skin, in
the musculature and their swimming ability is impaired. Heavily infected fish swim near
the surface in a slow and disoriented manner. Such fish are easy for piscivorous birds to
catch and it is evident that the spores could be rapidly dispersed to other ponds in the
bird feces. Piscivorous fish, reptiles and mammals could also spread the spores, but to a
more limited extent. Personal field observations indicate that this species of Henneguya
acts as a control on populations of A. fasciatus. The parasite spores can only escape from
the fish after its death so by incapacitating its host, the parasite is aiding in the
completion of its cycle and in its own dispersal. This is an exception to the rule that a
parasite does not harm its host. TAYLOR (1978) showed experimentally that whirling
disease in trout, caused by Myxosoma cerebralis, a myxosporidan which invades the
nervous system, could be transmitted in bird feces.
In our studies of Amazonian fishes, we have frequently encountered a coccidian
(Eimeriidae) in the livers of Triportheus elongatus. Although life-cycle studies of this species
have not been done, it seems probable that an intermediate host would be required. In this
case, the species would belong to the genus Calyptospora OVERSTREET, HAWKINS & FOURNIE,
1984. It may be that freshwater shrimp, Macrobrachium spp., are intermediate hosts for this
parasite and transmission occurs when fish eat infected shrimp.
Intestinal ciliophorans enter the fish by way of food and detritus eaten by the
host. Most of these small invaders are probably harmless commensals and they are
common in fish that ingest bottom detritus, such as the loricariid catfish. The case of
Ichthyophthirius multifiliis is quite different since these animals actively seek out a fish
host and invade the epithelium of its skin and gills. This parasite is not limited to the
Neotropics, but has become a pest of aquarium fishes the world over. There is a body
of evidence to suggest that “Ich” is more of a symptom than a disease. It may be that
this ciliophoran only invades fish that are already stressed by overcrowding, inade-
quate nutrition or poor water quality. In our study of wild-caught Amazon River
fishes, we have never been able to detect the presence of I. multifiliis. Cauca River fish,
in Colombia, only became infected with “Ich” after being maintained in aquaria for
several months.
While on a fish host, I. multifiliis grows from a size of less than 50 microns to nearly
one millimeter. After this, it drops off of the fish, settles to the bottom and encloses itself
in a protective capsule (cyst). Within the capsule, divisions occur which result in the
production of several hundreds of oval, ciliated tomites, each about 2-30 microns long.
The capsule eventually ruptures freeing the tomites to seek new hosts.
III. Pathology
Trypanosomes in man and other mammals cause a severe disease and they can invade and
weaken the heart musculature. We do not know if trypanosomes in Neotropical fishes
cause similar problems since detailed studies of these parasites within the tissues of the
hosts have not been made.
As noted above, Myxosporida and Coccidida cause the deformation of fish and
alter their behavior. The invasion of these parasites does not provoke an inflammato-
ry reaction on the part of the normal host so the cysts are able to grow continuously,
taking up the space of host cells, until the death of the fish becomes inevitable. An
infection known as “whirling disease” is a serious problem in Northern trout
hatcheries. This disease gets its name from the circular swimming behavior of
infected fish, which is caused by the invasion of the central nervous system by a
myxosporidan (Myxosoma cerebralis). The only cerebral myxosporidan known in
Neotropical fishes, is Myxobolus inaequus KENT & HOFFMAN, 1984, but it is not known
if this parasite modifies host behavior.
Intestinal ciliophorans, such as Balantidum and Nyctotherus, do not produce any
notable pathology. Ichthyophthirius multifiliis, on the other hand, causes skin lesions that are
easy to recognize. Infected areas are of a grey or whitish appearance and they have
irregular margins. Examination with a dissecting microscope, or a hand lens, will reveal the
presence of spherical trophozoites, typically in slow rotation.
IV. Prevention and treatment
No treatment is required for intestinal ciliophorans and the blood inhabiting trypano-
somes since these organisms have not been shown to be pathogenic. Eliminating leeches
from the aquaria breaks the transmission cycle of trypanosomes and prevents the
contamination of uninfected fish.
Prevention of Myxosporida and Coccidida is made more difficult by the fact
that the life-cycles are little understood. Infected fish showing obvious tumors
should be removed from the aquarium, or pond, and incinerated to limit further
dissemination of the organisms. Surviving fish should be placed in a clean aquarium
which is free of small invertebrate animals, such as Tubifex. As a treatment for
Coccidida, Monesin (Eli Lilly Co.) is routinely used in chickens and has been
reported to give good results against a fish coccidian (SOLANGI & O VERSTREET
1980). The drug may be injected or mixed with food, however, dosages would have
to be determined for particular host-parasite systems. This drug would probably be
useful against myxosporidan infections as well.
Prevention of Ichthyophthirius multifiliis and other skin invaders, involves care in
maintaining good water quality and good nutrition. At the first sign of the disease,
the fish should be removed to clean aquaria, separating the infected fish from the
uninfected ones. Sick fish should be treated by chemical dip, such as Malachite
green at 1:15,000 parts for 10-30 seconds or Formalin at 1:4,000 for 10-30 minutes.
Either of these treatments should be repeated on three consecutive days and then
the fish should be kept in a clean aquarium for one week of observation to see if
additional treatment is needed.
V. Collection and study methods
For trypanosomes, fresh blood smears are made on slides, allowed to air dry, fixed in
absolute methanol and stained in either Giemsa or Wright’s stain. The parasites should be
studied under oil immersion at 1,000x magnification.
Myxosporidan cysts in fish tissues are white and can frequently be seen with the
naked eye. The cysts can be excised, fixed in AFA (85 parts 85 % alcohol, 10 parts of
commercial formaldehyde and five parts glacial acetic acid), or in 10 % formalin solution,
and stored in glycerinated alcohol (90 parts of 80 % alcohol to 10 parts of glycerin).
Spores may be removed from the cysts with dissecting needles, partially dehydrated in 95
% alcohol and cleared in pure phenol for identification and study.
Ciliophorans are best studied alive, and vital stains, such as neutral red, can be used
to better visualize the nuclei. If permanent preparations are desired, the animals should be
killed by immersion in hot AFA, and after fixation in this solution for at least 5 minutes,
the slides may be made by the following procedure:
1) Place the material containing ciliophorans in a tapered centrifuge tube and spin at
low speed for 30 seconds.
2) Decant the AFA, replace with water, stir and centrifuge for 30 seconds.
3) Decant the water, replace with water containing one drop of Mayer’s carmalum
stain and stir.
4) Leave material in this stain solution for 5 minutes, then centrifuge for 30 seconds.
5) Decant the stain solution, replace with 50 % alcohol, stir and spin for 30 seconds.
6) Dehydrate by sequentially passing material in same tube through 70, 85, 95, 100 %
alcohol, xylol and half xylol-half balsam (spin for 30 seconds between each solution).
7) Sample of material can be removed from the xylol-balsam with a thin pipette and
mounted in balsam on microscope slides.
VI. Identification and keys
Blood inhabiting mastigophorans are easily identified as the genus Trypanosoma (Fig. 2-35).
Characters used to separate species are: body shape and size; position and size of the
blepharoplast and nucleus; length of the free part of the flagellum; and host identity.
Unfortunately, these parasites are extremely variable. There may be more than one
morphological type within a single species, and it is also possible for more than one
species to occur in the same host. Existing names for trypanosome species from
Neotropical fish are of doubtful validity since they are mainly based on host identity. Only
after these parasites have been studied by electron microscopy, artificial medium culturing
and physiological techniques will we know how many species are involved and which
names are valid. In Brazil, nominal species have been proposed as follows:
TRYPANOSOMA HOST FISH

albopunctatus Hypostomus albopunctatus
chaetostomi * Chaetostomus sp.
chagasi Hypostomus punctatus
dorbignyi Rhinodoras dorbigny
ferreirae * Characinidae
francirochai * Otocinclus francirochai
guaibensis Hypostomus commersoni
hypostomi Hypostomus aurogutatus
iheringi Franciscodoras marmoratus
immanis Loricariichthys anus
larai *Prochilodus sp.
loricariae Loricaria sp.
macrodontis * Hoplias malabaricus
margaririferi Hypostomus margaritifer
paivae * Characinidae
piracicabae Loricaria piracicabae
plecostomi Hypostomus sp.
regani Hypostomus regani
rhamdiae Rhamdia quelen and R. sapo
splendorei Rhamdia sapo
strigaticeps Hypostomus strigaticeps

zungaro Pseudopimelodus zungaro
(Hosts marked with an asterisk are characins and all others are catfish, Siluriformes).
Important considerations in the identifications of Myxosporida are: size and form of
the spore; number, position and size of the polar capsules; host identity; and site within
the host. The following key is offered as an aid in determining genera. Known species may
be identified by recourse to the figures and to the measurements in the checklist. All
measurements are given in micrometers (µm).
Key to the genera of Neotropical Myxosporida
I. Polar capsules located in one extremity of the spore only.

A. Spore capsule provided with 1-3 tail-like extensions.
1. Four polar capsules present ....................................................... Agarella (Fig. 2–16)
2. Two polar capsules present .................................... Henneguya (Figs. 2–17 to 2–24)
B. Spore capsule without tail-like extensions .................. Myxobolus (Figs. 2–1 to 2–15)
II. Polar capsules located in opposite extremities ...................... Myxidium (Figs. 2–25 to 2–27)
Clave para los géneros de Myxosporida Neotropicales
I. Cápsulas polares localizadas en solo una de las extremidades de la espora.

A. Cápsula de la espora provista de 1-3 extensiones en forma de cola.
1. Cuatro cápsulas polares presentes ............................................ Agarella (Fig. 2–16)
2. Dos cápsulas polares presentes ................................ Henneguya (Figs. 2–17 a 2-24)
B. Cápsula de la espora sin extensiones en forma de cola ................................................
................................................................................................ Myxobolus (Figs. 2–1 a 2–15)
II. Cápsulas polares localizadas en extremos opuestos ........ Myxidium (Figs. 2–25 a 2–27)
Key to Neotropical fish ciliophorans
Fish ciliophorans can be identified by: size and form of the body; shape of the
macronucleus; and nature of the cytostome.
I. Macronucleus horseshoe-shaped.
A. Body spherical, striate .............................................. Ichthyophthirius multifilis (Fig. 2–31)
B. Body bell-shaped, with oral denticles ............... Trichodina fariai (Figs. 2–32 & 2–34)
II. Macronucleus bean-shaped.
A. Body rounded ...................................................................Balantidium piscicola (Fig. 2–29)
B. Body elongate ......................................................... Rhynchodinium paradoxum (Fig. 2–33)
III. Macronucleus triangular ........................................................ Nyctotherus piscicola (Fig. 2–30)
Clave para los cilioforos de peces Neotropicales
Los cilioforos de peces pueden ser identificados por: tamaño y forma del cuerpo; forma
del macronúcleo; y naturaleza del citostoma.
I. Macronúcleo en forma herradura.

A. Cuerpo esférico, estriado ........................................ Ichthyophthirius multifilis (Fig. 2–31)
B. Cuerpo en forma de campana con dentículos orales ....................................................
................................................................................... Trichodina fariai (Figs. 2–32 & 2–34)
II. Macronúcleo en forma de frijol.
A. Cuerpo arredondeado .....................................................Balantidium piscicola (Fig. 2–29)
B. Cuerpo alargado ..................................................... Rhynchodinium paradoxum (Fig. 2–33)
III. Macronúcleo triangular ......................................................... Nyctotherus piscicola (Fig. 2–30)
VII. Checklist of Myxosporida from Neotropical freshwater fishes
Measurements are given in micrometers (µm).

Myxosporida
Myxozoa. Unicellular parasites. Trophozoite amoeboid, but without pseudopodia;
resistant infective spores form within trophozoite. Tissue parasites of fishes, rarely
amphibians.
Agarella DUNKERLY, 1915.
Spore elongate; of two valves having tail-like extensions; four polar capsules
present, two of which are smaller. Tissue parasites of freshwater fish.
A. gracilis DUNKERLY, 1915: testes of Lepidosiren paradoxa: Brazil. (Fig. 2–16).
Spore = 16 x 4; tail = 14.
Henneguya THÉLOHAN, 1892. Spore elongate; of two valves having tail-like
extensions; two polar capsules present. Tissue parasites of freshwater fish.
H. iheringi PINTO, 1928: gill filaments of Serrasalmus spilopleurus: Brazil. (Fig. 2–
17). Spore = 22 x 6; polar capsules = 3.4 x 2.
H. leporini NEMECZEK, 1926: urinary bladder of Leporinus mormyrops: Brazil.
(Fig. 2–18). Spores = 13–15 x 5; tail = 15–18; polar capsule = 5–8.
H. linearis (GURLEY, 1893) LABBÉ, 1899: peritoneum of Rhamdia sebae and Pseudo-
platystoma fasciatum: Brazil. (Fig. 2–21). Spore = 3 x 4 times longer than wide.
H. lutzi CUNHA & FONSECA, 1918: gall bladder of Pseudopimelodus zungaro: Brazil.
(Fig. 2–19). Spore = 11 x 7; tail = 22.
H. malabarica AZEVEDO & MATOS, 1996: gill filaments Hoplias malabaricus: Pará
State, Brazil. Spore = 26.6-29.8.
H. oculta NEMECZEK, 1916: gills of Loricaria sp.: Brazil. (Fig. 2–22). (Measure-
ments unavailable).
H. schizodon EIRAS, MALTA, VARELLA & PAVANELLI, 2004: kidney of Schizodon
fasciatus: Amazon River, Manaus, Brazil. Spore = 12–13 x 3–4; caudal
process = 15–17.
H. theca KENT & HOFFMAN, 1984: brain of Eigenmannia virescens: Brazil. Spore
= 40.6–52.6 x 3.0–4.1.
H. wenyoni PINTO, 1928: gills of Astyanax fasciatus: Brazil. (Fig. 2–20). Spore =
11–12 X 4.5–6.0; tail = 8–12; polar capsule = 1.5.
Myxidium BÜTSCHLI, 1882. Spores elongate, with one polar capsule at each
extremity. Tissues of fishes, rarely of amphibian and aquatic reptiles.
M.cruzi PENIDO, 1927: bile duct of Chalcinus nematurus: Brazil. (Fig. 2-25).
Spores = 17-18 x 5-7; polar capsules = 5-6 x 2-3.
M.fonsecai PENIDO, 1927: gallbladder of Carapus fasciatus: Brazil. (Fig. 2-26).
Spores = 7-9 x 2.5-3; polar capsules = 2-3.
M.gurgeli PINTO, 1928: bile duct Acestrorhamphus sp.: Brazil. (Fig. 2-27). Spores
= 14.6 x 8.5-3; polar capsules = 3.4 x 3.
Myxobolus BÜTSCHLI, 1882. Spores oval or pyriform, with two polar capsules in
one extremity; valves without tail-like extensions. Tissues of fishes and
rarely amphibians.
M.associatus NEMECZEK, 1926: kidney of Leporinus mormyrops: Brazil. (Fig. 2-1).
Spore = 15 x 10; polar capsule = 7.
M.braziliensis CASAL, MATOS & AZEVEDO, 1996: gills of Bunocephalus coracoideus:
Amazon River, Pará State, Brazil. Spore = 9.5-11 x 4.7-6.
M.cunhai PENIDO, 1927: intestine of Serrasalmus piraya and Pimelodus clarias:
Brazil. (Fig. 2-2). Spore = 9-11 x 4-6.
M.galaxii SZIDAT, 1953: gills of Galaxias maculatus: Argentina. (Fig. 2-4). Spore
= 14-15 x 9-10.
M.inaequalis GURLEY, 1893: skin of Pimelodus clarias: Brazil. (Fig. 2-3). Spore =
5.2-3.3.
M.inaequus KENT & HOFFMAN, 1984: brain of Eigenmannia virescens: Brazil.
Spore = 15.6-22 x 7.8-9.3.
M.kudoi GUIMARÂES & BERGAMIN, 1938: skin of unidentified fish: Brazil. (Fig.
2-5). Spore = 8.8-8.9 x 6.5-7.3; capsules = 3.5-4.2 x 1.3-2.0.
M. lutzi ARAGÂO, 1919: testis of Poecilia vivipara: Brazil. (Fig. 2-6). Spore = 10 x 7.
M. magellanicus SZIDAT, 1953: gills of Galaxius maculatus: Argentina. (Fig. 2-
7). Spore = 10-13 x 8-9; polar capsules = 3.
M.noguchii PINTO, 1928: gills of Serrasalmus spilopleurus: Brazil. (Fig. 2-8). Spore
= 13.6 x 8.5; polar capsules = 6.8-2.2.
M.pygocentris PENIDO, 1927: intestine of Serrasalmus piraya: Brazil. (Fig. 2-9).
Spore = 15-16 x 9-11; polar capsules = 9-11 x 3-4.
M.serrasalmi WALLIKER, 1969: spleen, kidney & liver of Serrasalmus rhombeus:
Brazil. (Fig. 2-10 A-B). Macrospore = 12-18 x 7-10; polar capsules = 6-9 x
2-4; microspore = 7.0-9.5 x 3.5-5; polar capsules = 5-7 x 1.2.
M. stokesi PINTO, 1928: skin of Pimelodus sp.: Brazil. (Fig. 2-11). Spore = 8.5 x 5.3.
Myxobolus sp. SZIDAT, 1953: skin of Pimelodus albicans: Argentina. (Fig. 2-12).
Spore = 15 x 8.
Myxobolus sp. WALLIKER, 1969: spleen and liver of Colossoma bidens: Brazil. (Fig.
2-14). Spore = 8-10 x 4-7; polar capsules = 3.5-5.0 x 1.0-2.5.
Myxobolus sp. WALLIKER, 1969: kidney of Serrasalmus sp.: Brazil. Spore = 9 x 11

x 5-6.5; polar capsules = 5-6 x 1.5-2.0.
Checklist of Ciliophora from Neotropical freshwater fishes
Measurements are given in micrometers (µm).

Phylum Ciliophora.
Unicellular animals possessing cilia or structures composed of cilia in
some stage of the life-cycle. Basal granules for coordinating ciliary motion
also present. Cystostome (mouth), macronucleus and micronucleus usually
present. Reproduction by binary fission and conjugation. Free-living and
parasitic.
Balantidium CLAPARÈDE & LACHMAN, 1858. Trophozoite oval or elliptical. Cytostome
near anterior extremity; cytopharynx not well developed. Body ciliation
uniform. Cytopyge and contractile vacuole terminal. Intestine of inverte-
brates and vertebrates.
B. piscicola GÈZA, 1913: gut of Piaractus brachypoma and Pimelodus clarias: Brazil.
(Fig. 2-29). Body = 36 x 28; macronucleus = 12 x 6-7; cilia = 3-4.
Nyctotherus LEIDY, 1877. Body oval, compressed; cytostome in midbody; cytopharynx
long, with undulating membrane. Macronucleus large, triangular, anterior.
Cytopyge and contractile vacuole terminal. Gut of invertebrates, fish and
mammals.
N. dilleri EARL & JIMÉNEZ, 1969: Cichlasoma fenestratum: México. Body = 154-
280 x 80-140; macronucleus = 6-10 x 12-30.
N. piscicola DADAY, 1905: Colossoma brachipomum, Pimelodus clarias and Acestrorham-
pus sp.: Brazil. (Fig. 2-30). Body = 112-153; macronucleus = 12-30 x 30-90.
Rhynchodinium CUNHA & PENIDO, 1927. Body elongate, cylindrical, tapering anteriorly;
cilia long, lacking posteriorly. Macronucleus ovoid, anterior in position.
Cytopyge and contractile vacuole terminal. Intestine of fish.
R. paradoxum CUNHA & PENIDO, 1927: Pterodoras granulosus: Brazil. (Fig. 2-33).
Body = 20-30 x 5-10.
Trichodina EHRENBERG, 1834. Body cup or bell-shaped, with three circular rows of
cilia; oral surface provided with chitinized ring and circle of denticles.
Macronucleus U-shaped. On skin of marine and freshwater crusta-
ceans and fish.
T. fariai CUNHA & PINTO, 1928. Sphaeroides testudineus: Brazil. (Figs. 2-32 & 2-
34). Body = 32 x 20-22; with 24-28 denticles.
Sporozoa
Calyptospora OVERSTREET, HAWKINS & FOURNIE, 1984. Sporocyst covered with a thin
veil supported by one or more sporopodia; lacking Stieda body; with
membrane covered oblong opening. Suture extending a short distance
distally on opposite sides then converting to low ridges that continue to
posterior end. Sporozoite developing in invertebrate host. Parasites of

marine and freshwater fishes.
C. tucunarensis BÉKÉSI & MOLNÁR, 1991: liver of Cichla ocellaris: Ceará State,
Brazil. (Figs. 2-44 A-B). Oöcysts = 23-26 in diameter. Sporocysts = 7.2-9.1
x 3.5-5.0.
C. spinosa AZEVEDO, P. MATOS & E. MATOS, 1993: liver of Crenicichla lepidota:
Pará State, Brazil (measurements unavailable).
VIII. Plates of Protozoa and Ciliophora (Figs. 2–1 to 2–44)
2-1 2-2 2-3 2-4
2-5 2-6 2-7 2-8

10 µm
2-10
2-9 A B 2-11
2-12 2-13 2-14 2-15

Myxoxporida spores: 2-1. Myxobolus associates; 2-2. M. cunhai; 2-3. M. inaequalis; 2-4. M. galaxii; 2-5.
M. kudoi; 2-6. M. lutzi; 2-7. M. magellanicus; 2-8. M. noguchii; 2-9. M. pygocentris; 2-10a. M. serrasalmi
(macrospore); 2-10b. M. serrasalmi (microspore); 2-11. M. stokesi; 2-12. Myxobolus sp. from Pimelodus
albicans; 2-13. Myxobolus sp. from Serrasalmus sp.; 2-14. Myxobolus sp. from Colossoma bidens; 2-15.
Myxobolus sp. from Curimata elegans (2-15. orig., others redrawn from PINTO (1928c, d) and
WALLIKER (1969)).
10 µm
2-16 2-17 2-18
2-19 2-20
2-21 2-22 2-23 2-24
2-16. Agarella gracilis (spore); 2-17 to 2-24. Henneguya spp. (spores): 2-17. H. iheringi; 2-18. H.leporinus;
2-19. H. lutzi; 2-20. H. wenyoni; 2-21. H. linearis; 2-22. H. oculta; 2-23. Henneguya sp. from Brycon
melanopterus; 2-24. Henneguya sp. from Mylossoma duriventris (2-23. & 2-24. orig.; 2-16. to 2-22. redrawn
from PINTO (1928 b-d).
10 µm
2-25 2-26 2-27
25 µm
2-28 2-29
25 µm
250 µm
2-30 2-31
2-25. Myxidium cruzi; 2-26. M. fonsecai; 2-27. M. gurgeli; 2-28. Calyptospora sp. (Eimeridae) (2-25. to 2-
28. to same scale); 2-29. Balantidium piscicola; 2-30. Nyctotherus piscicola; 2-31. Ichthyophthirius multifilis
(2-28. orig., others redrawn from PINTO (1928 a, d)).
10 µm
2-32
25 µm
2-33
10 µm
2-34
2-35
2-32. Trichodina fariai (lateral view); 2-33. Rhynchodinium paradoxum; 2-34. Trichodina fariai (oral
surface); 2-35. Trypanosoma sp. (2-32. to 2-34. redrawn from PINTO (1928d); 2-35. from FRÖES et al.
(1978)).
CyanMagentaYellowBlack page 37
2-36 2-37
2-38 2-39
2-36. Henneguya sp. (cysts in gill filament of Mylossoma duriventris); 2-37. Henneguya sp. (spores from gill
filament of Semaprochilodus insignis); 2-38. Henneguya sp. (cysts in gill filament of Brycon melanoptera);
2-39. Myxobolus sp. (spores from liver of Semaprochilodus insignis). (Original photographs).
2-40
2-41
2-40. Astyanax fasciatus from Cauca River, Colombia, with cysts of Henneguya sp.; 2-41. Curimata
elegans from Amazon River, near Belém, Pará, Brazil with cysts of Myxobolus sp. (Original
photographs).
2-42 2-43
2-44
A B
2-42. Myxobolus braziliensis mature spore (redrawn from CASAL et al. 1996); 2-43. Myxobolus inaequus
(redrawn from KENT & HOFFMAN 1984); 2-44. Calyptospora tucunarensis: A. mature sporocyst; B.
sporulated oöcyst (redrawn from BÉKÉSI & MOLNÁR 1991).
IX. Cited and general references
BARASSA, B., CORDEIRO, N.S. & S. ARANA (2003): A new species of Henneguya, a gill parasite of
Astyanax altiparane (Pisces: Characidade) from Brazil, with comments and histopathology and
seasonality. - Mem. Inst. Oswaldo Cruz 98(6): 761-765.
BÉKÉSI, L. & K. MOLNÁR (1991): Calyptospora tucunarensis n. sp. (Apicomplexa : Sporozoa) from the
liver of tucunaré Cichla ocellaris in Brazil. - Systematic Parasitology 18: 127-132.
BOTELHO, C., JR. (1907): Sur deux nouveaux trypanosomes des poissons. - C.R. Soc. Biol. Paris
59(2): 128-129.
CASAL, G., MATOS, E. & C. AZEVEDO (1996): Ultrastructural data on the life cycle stages of
Myxobolus braziliensis n. sp., parasite of an Amazonian fish. - Europ. J. Protistol 32: 123-127.
CHEUNG, P.J., NIGRELLI, R.F. & G.D. RUGGIERI (1983): Coccidian parasite from the liver of the black
piranha, Serrasalmus niger SCHOMBURGK (Abstr.). - Eastern Fish Health Workshop, 21-23 June,
1983. Nat. Fish Health Res. Lab., Kearneysville, West Virginia: 3 pp.
CUNHA, A.M. & J.C.N. PENIDO (1926): Nouveau protozoaire parasite des poissons: Zelleriella piscicola
n. sp. - C.R. Soc. Biol. Paris 94: 1003.
CUNHA, A.M. & J.C.N. PENIDO (1927): Rynchodinium paradoxum n. gen., n. sp. protozoaire parasite
d’um poisson d’eau douce. - C.R. Soc. Biol. Paris 97(36): 1793-1794.
DADAY, E. (1905): Nyctotherus piscicola n. sp., ein neuer Fischparasit aus Südamerika. - Zool. Anz.
28: 233-238.
EARL, P.R. & G. JEMÉNEZ (1969): Nyctotherus dilleri n. sp. from the fish Cichlasoma fenestratum in
Veracruz. - Trans. Am. Mic. Soc. 88(2): 287-292.
EIRAS, J.C., MALTA, J.C., VARELA, A. & G.C. PAVANELLI (2004): Henneguya schizodon n. sp. (Myxosoa,
Myxobolidae), a parasite of the Amazonian teleost fish Schizodon fasciatus (Characiformes,
Anostomidae). - Parasite 11: 169-173.
FONSECA, F. (1935): Trypanosomas de peixes brasileiros. Decrição de uma nova espécie. - Mem.
Inst. Butantan 9: 151-184.
FONSECA, F. & Z. VAZ (1928a): Novos trypanosomas de peixes brasileiros. - Ann. Fac. Med. Univ.
São Paulo 3: 69-94.
FONSECA, F. & Z. VAZ (1928b): Trypanosoma francirochai n. sp. parasito de Otocinclus francirochai
HERRING, 1928 (peixe de água doce do Estado de São Paulo). - Bol. Biol. 11: 4-5.
FONSECA, F. & Z. VAZ (1929): Novas espécies de trypanosomas de peixes brasileiros de água doce.
- Bol. Biol. 13: 36-41.
FRÖES, O,M., FORTES, E., LIMA, D.F. & V.R.V. LEITE (1978): Três espécies novas de tripanosomas de
peixes de água doce do Brasil (Protozoa, Kinetoplastida). - Rev. Brasil. Biol. 38(2): 461-468.
HOFFMAN, G.L. (1999): Parasites of North American freshwater fishes. Second Edition. – Cornell
Univ. Press: 539 pp.
HORTA, P.P. (1910): Trypanosoma chagasi n. sp. parasito de Plecostomus punctatus. - Brasil Médico
24(28): 273.
HORTA, P.P. & A. MACHADO (1911): Estudos citológicos sôbre o “Trypanosoma chagasi” n. sp.
encontrado em peixes do gênero Plecostomus. - Mem. Inst. Oswaldo Cruz 3(2): 344-366.
KENT M.L. & G.L. HOFFMAN (1984): Two new species of Myxozoa, Myxobolus inaequus sp. n. and
Henneguya theca sp. n. from the brain of a South American knife fish, Eigemannia virescens (V.). -
J. Protozool. 31(1): 91-94.
MARTINS, M.L., SOUZA, V.N., MORAES, J.R. & F.R. MORAES (1999): Gill infection of Leporinus
macrocephalus GARAVELLO & BRITSKI, 1988 (Osteichthyes: Anostomidae) by Henneguya leporinico-
la n. sp. (Myxozoa: Myxobolidae) description, histopathology and treatment. - Rev. Brasil.
Biol. 59(3): 537-534.
NEMECZEK, A. (1926): Beiträge zur Kenntnis der Myxosporidenfauna Brasiliens. - Arch. Protistenk.
54(1): 137-149.
OVERSTREET, R.M., HAWKINS, W.E. & J.W. FOURNIE (1984): The coccidian genus Calyptospora n. gen.
and family Calyptosporidae n. fam. (Apicomplexa), with members infecting primarily fishes. -
J. Protozool. 31(2): 332-339.
PENIDO, J.C.N. (1927): Quelques nouvelles Myxosporidies parasites des poisons d’eau douce du
Brésil. - C.R. Soc. Biol. Paris: 850-852.
PINTO, C. (1928a): Myxidium gurgeli nova espécie. Myxosporídeo parasito da vesícula biliar de peixe
de água doce do Brasil. - Sci. Med. 6(2): 86-87.
PINTO, C. (1928b): Henneguya wenyoni n. sp. myxosporidie parasite des branchies de poisson d’eau
douce du Brésil. - C.R. Soc. Biol. París 98(17): 1580.
PINTO, C. (1928c): Myxobolus noguchii, M. stokesi e Henneguya iheringi, espécies novas de myxosporídeos
de peixes de água doce do Brasil. - Bol. Biol. 12: 41-43.
PINTO, C. (1928d): Myxosporídeos e outros protozoários intestinais de peixes observados na
América do Sul. - Arch. Inst. Biol. 1: 101-136, 42 figs.
SOLANGI, M.A. & R.M. OVERSTREET (1980): Biology and pathogenesis of the coccidium Eimeria
funduli infecting killifishes. - J. Parasitol. 66(3): 513-526.
SPLENDORE, A. (1910): Trypanosomes de poisons brésiliens. - Bull. Soc. Path. Exot. 3: 521-524.
TAYLOR, R.L. (1978): Transmission of salmonid whirling disease by birds fed trout infected with
Myxosoma cerebralis. - J. Protozool. 25(1): 105-106.
WALLIKER, D. (1969): Myxosporidea of some Brazilian freshwater fishes. - J. Parasitol. 55(5):
942 - 948.
WOLF, K. (1982): Tubifex worms identified as essential hosts for salmonid whirling disease. - Res.
Infor. Bull. U. S. Fish & Wildlife Service: 2 pp.
3
MONOGENOIDEA
In collaboration with W. A. BOEGER and R. T. VIANNA
The Class Monogenoidea (also known as Monogenea or monogenetic trematodes)

consists of hermaphroditic mostly ectoparasitic platyhelminths with direct life-cycles. In
fishes, they inhabit the gills, skin, nasal fossae, ureters and a few are found in the intestinal
ducts. In amphibians and aquatic reptiles they may be found in the mouth, cloaca, or
urinary bladder. Exceptionally, species of monogenoids are reported from mammals
(hippopotamus) and invertebrates (ex. squid, copepods).
The most useful recognition feature of monogenoids is the posterior attachment
organ (haptor). This structure is usually flattened and frequently disk-like. It may be
armed with sclerotized structures such as anchors, bars (to support the anchors), and
hooks (Fig. 3-4D). In some genera, the haptor may have suckers, clamps or loculi along
with or in place of these sclerites. The anterior end of monogenoids can also attach to
host tissue by means of adhesive secretions from the cephalic lobes that are stored on the
head organs. Two pairs of eyes are often present on the cephalic area, though in some
species these have been lost.
The digestive system of monogenoids consists of a mouth, a pharynx, an esophagus
and, usually, two intestinal ceca. These may be simple, have diverticula or be posteriorly
confluent. The female reproductive system contains a germarium (= ovary), oviduct,
ootype, seminal receptacle (sometimes absent), uterus, and one or two vaginae (these may
be absent as well). The male system includes a testis (or several), a vas deferens, a seminal
vesicle (saccate or as an expansion of the vas deferens) and a copulatory complex. The
copulatory complex is usually composed of a male copulatory organ (MCO) and an
accessory piece, that may be absent in some taxa. The MCO may be either muscular or
sclerotized and its morphology is important in the classification of the group. In some
genera, other sclerites may be associated with the reproductive system (e.g., a gonadal bar
in Gonocleithrum and a hook-like vaginal sclerite in Urocleidoides).
Monogenoidea is perhaps the least known taxon of fish parasites in the Amazon
region, which is also true for the entire Neotropical realm. Until the 1970’s, very few
species were known from Amazonian and Neotropical continental waters. The earliest
references of the taxon in Amazonian fish species are limited to fish kept in aquaria
abroad, mainly in the United States (see references of MIZELLE and coauthors). At that
time, very few local scientists contributed to the recovery of the richness of the group.
Undoubtedly, the collaboration between two fish parasitologists, VERNON E. THATCHER (at
that time in Cali, Colombia) and DELANE C. KRITSKY (Idaho State University, USA) and
subsequent efforts and collaborations represent the most intense and consistent effort
towards the understanding of the diversity of this group, especially in Brazilian waters.
Recently other groups have awakened to the abundance and richness of the Neotropical
Monogenoidea of freshwater fish species (ex. TAKEMOTO, LUQUE, VIOZZI and others).
While there are evidences of monogenoids with a broad host species spectrum
(KRITSKY et al. 1996 a, b & 1997 a, b), most show relatively high host specificity,
occurring on a single species or on closely related fish. The richness per host species
appears, however, to be highly variable. The red-breasted piranha (Pygocentrus nattereri)
hosts about 30 different species while many species of Siluriformes, for example, bear
a single or a few species. Most medium-sized and large fish harbor Monogenoidea. A
few such hosts have been reported to be negative for these parasites but it may be that
this merely reflects inadequate sampling methods. On the other hand, small fish
species have few monogenoids. We have searched intensively among the small
siluriforms (eg. whale catfishes and others) without turning up a single representative
of this group of worms. Characiform fish, especially Serrasalmidae, show the greatest
abundance of species of monogenoids.
Despite the efforts of taxonomists, the diversity of the Monogenoidea in the
Amazon and in the Neotropics is still largely unknown. Around 308 species are described
in approximately 70 genera, from 144 species of fish or an average of 2.14 species of
monogenoids per fish species. The family Dactylogyridae is overwhelmingly the most
abundant taxon in continental waters of South America. However, species of Gyrodactyl-
idae are being systematically described while only a few Diplectanidae, Monocotylidae and
Hexabothriidae appear to have representatives in these waters.
Assuming a conservative absence of species of monogenoid from 10 % of fish
species, a rough estimate of species of this clade of parasites suggest that fewer than 3
% of freshwater fish Monogenoidea are known. These estimates consider the most
recent evaluations of freshwater Neotropical fish species. Ichthyologists expect that
more than 6,025 species of freshwater fishes exist in this realm (MALABARBA & ROSA
2003). This estimated number of species of Monogenoidea may be greater, however,
since the composition of the monogenoid fauna of a single fish species may vary
greatly according to the geographic distribution (see Boeger & Kritsky 1988) and site
specificity of the host (even species known to harbor monogenoids have had few of the
possible microhabitats screened).
Although little is known about the association of the orders and families of hosts
and the genera of monogenoids, some indications are appearing. This is especially true of
the Dactylogyridae For example, species of Gussevia and Sciadicleithrum are reported solely
on Cichlidae; Anacanthorus spp are known only from Characiformes; Vancleaveus spp.,
Demidospermus spp. and Aphanoblastella spp are recorded from Pimelodidae Siluriformes;
while Amphithecium spp, Mymarothecium spp, Nothozothecium spp., and others, are unique to
Serrasalmidae hosts. There are, obviously, many other examples. In general one can guess
the parasite group if one knows the host species.
II. Life cycle and transmission
The life cycle of most monogenoids is direct in that all stages are completed on a single host.
These worms are also highly host specific and will develop on only one host species or on
closely related species of a single genus. Monogenoids can be either viviparous (some
Gyrodactylidae) or oviparous (all other families). In most cases, the larval stage (oncomira-
cidium) is ciliated and already has posterior sclerites (e.g., hooks and sometimes even
anchors) with which to attach to the same or a different host after hatching takes place. The
bands of cilia on the larva permit it to swim to a nearby fish of the same species. As the larva
settles down and begins to grow, the cilia are lost. The eggs of oviparous forms frequently
have one or more polar filaments that may serve to anchor them in the gill mucous until a
young adult hatches, with the haptoral sclerites already well formed. The eggs of oviparous
Gyrodactylidae have a drop of cement on the tip of a short filament that is used to “glue”
the egg on the external surface of their hosts. Viviparous gyrodactylids give birth to
complete worms, which already carry an offspring in the uterus when born.
According to HARRIS (1983), Oogyrodactylus farlowellae can complete its cycle from egg to
adult in 11 to 13 days at 27 °C. Other species probably have similarly short lifecycles.
III. Pathology
Most monogenoids attach superficially to the gill epithelium. They can move about at will
and generally produce little damage to the fish. However, even the relatively non-pathogenic
forms can produce extensive damage when large numbers are present on the gills. Direct
transmission, as observed in Monogenoidea, is greatly favored by proximity of hosts of the
same species and large populations of parasites develop rapidly. In the crowded conditions
of intensive pisciculture, monogenoids can rapidly become a problem. There are unreported
cases, especially on cultured Colossoma macropomum, in which specimens of apparently non-
pathogenic species of monogenoids were capable of consuming a significant proportion of
the gill tissue, leaving only cartilage in certain areas of the gill filaments.
Even if they do not cause excessive tissue damage, some monogenoids may provoke an
excessive production of mucus by the gill filaments. Apparently, such species feed on mucus
and its presence appears to stimulate the abundant production of this substance. Whatever the
cause, gill filaments coated with mucus have a reduced respiratory capacity. Thus, fish that
seem to tolerate heavy infestations may die suddenly when the oxygen content of the water
falls slightly. We should therefore consider all monogenoids as potentially harmful and try to
eliminate them from culture tanks, or at least control their numbers.
Most Neotropical monogenes belong to the family Dactylogyridae, which is
generally believed to include highly pathogenic species. Gyrodactylidae, with the excep-
tion of G. salaris, a notorious pathogen of salmon in European waters, are apparently less
pathogenic. In Europe, the species of Linguadactyla are known to provoke extensive
epithelial hyperplasia and the epithelium overgrows the haptor, which fixes the worm in
place (BYCHOWSKY 1957). These parasites feed on epithelial cells. A similar species,
Linguadactyloides brinkmanni THATCHER & KRITSKY, 1983, has been found on a Brazilian
Amazon food fish, Colossoma macropomum. This species appears more pathogenic since it
actively penetrates the tissues of the gill filament. This action is aided by secretions from
penetration glands that L. brinkmanni has located in the haptoral peduncle. When the
haptor has penetrated to the cartilaginous supporting rod of the filament, the anchors
directly engage this structure. As a host response, the cartilaginous tissue overgrows the
anchor points, effectively fixing the worm to a permanent location (Fig. 3-61).
The initial host response to the penetration of L. brinkmanni is basically an
inflammatory reaction. Both erythrocytes and leucocytes are numerous around the site of
the lesion. A funnel-like tube from the surface of the filament to the cartilage support is
maintained open by the constant movements of the worm. Apparently, this lesion
hemorrhages, at least initially. Epithelial hyperplasia occurs for some length along the
filament, and becomes more pronounced at the penetration site (Fig. 3-62). Epithelial
overgrowth at this site is probably a continuous process, which is partially checked by the
feeding activity of the worm. These parasites ingest blood cells as well as epithelium and
are usually red in color. The prolonged activities of this parasite can lead to the almost
complete destruction of the gill filaments (Fig. 3-63).
An as yet undescribed monogenoidean has been found on a doradid catfish that may
be even more pathogenic than L. brinkmanni. This species also penetrates to the cartilage,
but it does so from the other (lamellar) side of the filament (Fig. 3-64). The presence of
the haptor in the center of the filament provokes intensive cellular concentration and a
tumoroid growth that swells the filament and alters blood circulation.
Newly obtained fish, from whatever source, should be assumed to be infested and should be
treated before being introduced into a tank, pond, or aquarium containing other specimens
of the same or related species. Dip treatment can be used effectively against monogenoids
since they are relatively sensitive to chemicals and are in direct contact with the water.
Although many different chemical dips will dislodge monogenoids from living fish, the
easiest, most efficient and cheapest is a 1:4,000 solution of commercial formaldehyde (1 cm3
of formaldehyde for each 4 liters of water) used for 10-30 minutes. The time for immersion
in these dips is variable because different species of fish have different degrees of tolerance
for the chemicals. The sensitivity of the fish to the chemicals can also vary with water
temperature and the physiological condition of the individuals. One or two of the infested
fish should be treated first to test sensitivity before large scale treatment is attempted. At the
first sign of distress, the fish should be removed from the dip.
Based on the limited knowledge of the richness of the group, it is obvious that studies of
the alpha taxonomy of Monogenoidea are still essential in the Amazon region and in the
entire Neotropics. Taxonomic studies on the group, however, require basic scientific skills
and special care with the descriptions and proposal of new taxa in order to produce
robust knowledge to be applied in several areas of biology, including systematics, ecology,
and historical biogeography.
Freshwater monogenoids are usually very small. Some are as small as 200-300
micrometers long and, thus, are easily lost if the person collecting these animals is
untrained or the collection method is inadequate. Our experience has demonstrated that
the warm-water method (described below) is not only the best collection method
available, but also produces excellent specimens for morphological studies. This technique
allows rapid and adequate processing of specimens in the field, leaving the time-
consuming work of picking the worms from the gills and other organs to the laboratory,
where more adequate equipment and time are readily available.
Specimen preparation is also fundamental, for obvious reasons. In general, descrip-
tions of species involve cleared specimens, for the study of hard, sclerotized, structures,
and stained worms, which are necessary for the study of the morphology of internal
organs and structures and to complement the study of the sclerotized parts. The choice
of stain is fundamental and frequently makes the difference between an extensive and a
limited morphological characterization.
The use of phase contrast or, sometimes differential contrast microscopy is
necessary for precise illustrations and descriptions. Often, bad descriptions are caused by
the exclusive use of light microscoscopy which does not provide the resolution required
to see the fine details. Many taxonomic accounts fail to present detailed and adequate
morphological interpretations of the sclerotized parts of haptor, vagina, male copulatory
organ. These limitations repeatedly result in morphologic, taxonomic, and systematic
errors that greatly hinder the evolution of the knowledge in the group.
Illustrations, especially those of the hard sclerites, need to be very precise.
Frequently, species can be differentiated by small, frequently overlooked, inconspicuous
features of these structures. The most difficult structure to be illustrated is, by far, the
hook. Curiously, although the morphology of these structures is fundamental for the
systematic studies in the group, they receive the least attention from many investigators.
The structure of the male copulatory organ of many species, on the other hand requires
the most detailed morphological study and interpretation.
An important point to consider in taxonomic studies of Amazonian and Neotropical
Monogenoidea is the establishment and proposal of supra-specific taxa. While some genera
appear to represent clear specific groups, easily separated from other genera and groupings
by large morphological gaps (e.g., Curvianchoratus), many generic (or other supra-specific)
groupings are characterized by faint, often inconspicuous, morphological differences. In the
absence of an adequate phylogeny, the decision of proposing new supra-specific taxa
requires a “fish-eye-lens” study. Groupings of species are often recognized only after
extensive morphological study of many related species (in general, species of closely related
fish species) that allow the comprehension of shared characters, hopefully reflecting
common ancestry. Certification of the taxonomic status of these groups will certainly
depend on future phylogenetic reconstructions. Thus, a conservative approach is strongly
recommended and investigators working on this group of platyhelminths are urged to
exercise extreme caution in the proposal of supra-specific (and specific) taxa.
Mistakes of the past are being repeated in the present so a word of caution is
required. Except, perhaps, for lineages that invaded the Neotropical freshwater
environments recently, apparently from and with marine host lineages (e.g., the
freshwater dactylogyrids and diplectanids from Sciaenidae), most monogenoid parasites
of primary fish groups are unique to South and sometimes Central America. In the past,
Neotropical species were described as members of North American or European
genera, reflecting the lack of knowledge of the diversity of the group in the region. For
example, species of Urocleidus and Cleidodiscus, described from piranhas by MIZELLE and
co-authors (e.g., MIZELLE & PRICE 1965) are presently allocated to exclusively Neotropi-
cal genera (e.g., Amphithecium, Calpidothecium, Urocleidoides) (check KRITSKY et al. 1983,
and BOEGER & KRITSKY 1988 for many examples). Although the study of the group in
South America accumulated much information on species diversity there are still
unjustified attempts to include new species in genera with species from other continents
(e.g., Acolpenteron australe VIOZZI & BRUGNI, 2003 described from a Percichtyidae from
Argentina).
Most of the fauna of freshwater Monogenoidea of South America is closely related
to the African fauna (e.g., KRITSKY & BOEGER 1995). These faunas, however, are composed
of apparent sister but easily distinguishable generic groupings. Examples of genera with
species in the Neotropical region and elsewhere includes Euryhaliotrema, Diplectanum
(although there are strong indications that this genus does not represent a monophyletic
group) and Gyrodactylus. The latter, however, probably also represent an exceptional case
of rapid dispersion to other regions and continents of the globe, as suggested by BOEGER
et al. (2003). Summarizing, if a new species collected does not fit the diagnosis of
Neotropical genera, the investigator should seriously consider the possibility that it
represents the member of an undescribed genus.
Although common in the initial stages of the taxonomic studies of Amazonian and
Neotropical Monogenoidea, the use of specimens collected from aquarium fishes should
be accompanied by careful consideration of the type host species. Exchange of parasites
between hosts kept in captivity is very common, even between species of distantly related
higher categories of hosts (e.g., orders).
Collection of fish monogenoids is easily accomplished in the field and does not
require much more than buckets, a portable stove, formalin, ethanol (if DNA samples are
required), vials, labels, a notebook, and a pencil. It is, however, absolutely necessary that
animals are killed relaxed in order to allow adequate study of internal anatomy. The
method described below can be modified according to the local conditions, size of the
animal, and requirements of each specific study.
If monogenoids of the fish body surface are the object of the study, special care
should be given to the method of capture and handling of hosts. In general, gill nets and
trawl nets are not good fishing methods to meet this objective. Gill nets, however, may be
used if the fish is not left in the net and in the water for too long. Hook and line are great
for this purpose, but this method is not very efficient for catching fish of a variety of
species. Dip nets are fine and minnow traps are excellent.
Once collected, the fish should be killed by pithing, or other method, placed in a
container and washed with hot water (about 60-70 °C). The container should be
vigorously shaken in order to detach the parasites from the fish tissue. The “vigorously”
should be stressed as anything less than this may result in poor sampling of the parasite
population of a fish. The fish can be removed from the container, the liquid can be
decanted and fixed according to the objective of the study: in ethanol, formalin or both.
An adequate method, depending on the size of the fish, is to fix half of the decanted
liquid and sediments in formalin, for morphological studies, and the other half in ethanol,
if the investigator is interested in DNA studies. In this case, and in the procedures
described below, the final concentration for both methods of fixation would be about 4-
5 % of formalin and 70-80 % of ethanol, respectively.
Nasal cavities can be washed with hot water (about 60-70 °C) and scraped with a
probe into another vial. Fixation can be achieved in the same way described above. Gills
can be excised, the gill arches separated, placed in a vial, flooded with hot water (about 70-
75 °C). Once again, the vial should be vigorously shaken to detach parasites from the gill
tissue. Fixation, as in the other cases, should be accomplished with ethanol and formalin,
in the same concentrations, but the gill arches should accompany the sediment and liquid
in the formalin vial - if added to the ethanol vial, the ratio ethanol : tissue may result in
poor fixation and hinder later processing of the DNA.
Collection of internal Monogenoidea requires necropsy with the use of stereomicro-
scope and should be carried out in an adequate place, such a laboratory. In this case, fresh
or living host specimens should be used. Fixation can be performed with the same
methods mentioned above.
Slide making
A few comments are in order before we start. Monogenoids, especially the small
ones, are easily hydrated or dehydrated with ethanol, not requiring extensive time nor
long, stepwise, ethanol series. Hence, the methods of slide preparation in the group are
usually much simpler than for any other group of Platyhelminthes. Additionally, an
adequate morphological study of Monogenoidea requires specimens prepared for the
study of sclerotized (“hard”) structures and internal (“soft”) morphology.
Sclerotized structures may be studied through the clearing of the specimens in one
of two media: Hoyer’s Mounting Medium1 or Gray and Wess Mounting Medium2. Hoyer’s
clears faster, is more reliable but has two major problems: it may overly flatten the
specimen and they may not last as long as specimens mounted in Gray and Wess. Gray
and Wess, on the other hand, is not as reliable in producing nicely cleared specimens and
requires longer “incubation” in a oven or on a hot plate (at about 56 ºC) to produce good
1
Formula for Hoyer’s Mounting Medium-Chloral hydrate: 200 g; Crystalline gum Arabic: 30 g; Glycerol: 20 ml;
Distilled water: 50 ml. Dissolve gum arabic in distilled water and let it sit over night. Add the chloral hydrate and
glycerol and either filter the mixture through clean glass wool, or more efficiently, centrifuge the mixture. Store
Hoyer’s in a brown bottle to prevent degradation by light.
2
Formula for Gray and Wess Medium-PVA (polyvinyl alcohol) 71-24 powder: 2.0 g; 70 % acetone: 7.0 ml;
Glycerin: 5.0 ml; lactic acid: 5.0 ml; distilled water: 10.0 ml. Make a paste of the dry alcohol with acetone. Mix
half of the water with glycerin and lactic acid, stir into paste. Add remaining of water drop by drop while
stirring. Solution will be cloudy but becomes transparent if warmed in water bath for 10 min.
specimens. This medium, however, is necessary if one applies the method indicated below
to observe and study the shield on the superficial bar of Gyrodactylidae.
In both cases, specimens that are kept in aqueous liquid (such as 4-5 % formalin) can
be directly transferred into a drop of either media on a glass slide. After orientation of the
specimen (preferably, with the ventral side up), a coverslip is placed on the drop of the
mounting medium and, especially in the case of Hoyer’s mounting media, it should be
sealed with transparent nail polish. Study of specimens prepared this way is necessarily
done with a phase-contrast or DIC microscope, as sclerites (such as anchors, bars and
hooks) are not readily visible with normal light microscopy.
Specimens to be used for the study of internal morphology should be stained with
Gomori’s trichrome3 as follows:
1. Place a drop of Gomori’s trichome in a small Petri dish and transfer 1-5 animals
from the formalin into this drop;
2. Stain the specimens for 1-5 minutes (according to the size of the specimens-it
would be wise to test the time with few specimens first);
3. Flood the dish with absolute ethanol;
4. Concentrate the specimens in the center of the dish by rotation of this container;
5. Pouring drops of water on the specimens should remove the stain from the overly
stained specimens (differentiation). This is the most important step in the process.
The organs should stain dark while the mesenchyma should be lightly stained
(usually light green).
6. Once the specimens are differentiated (as described above), the dish with ethanol
and the specimens should be lightly shaken in order to mix the remaining water
with the ethanol.
7. If only a few water drops were used, the specimens can, then, be cleared in
Beachwood (or synthetic) Creosote4, otherwise, it is recommended that the
specimens are further dehydrated in absolute ethanol prior to clearing.
8. With small specimens, it is better to place a drop of Creosote on a glass slide,
place the specimen in this liquid and, after it is cleared, carefully remove the
excess Creosote with a paper tissue.
9. Once cleared, the specimens can be mounted in Canada’s Balsam between slide
and coverslip.
The specimens prepared in this manner should be examined under normal
illumination but, often, the use of phase contrast and DIC helps in revealing some hidden
detail of sclerotized organs and parts.
Another method has been developed to allow visualization and study of sclerites
with very delicate and thins areas, which are extremely hard to differentiate using
specimens cleared in Hoyer’s or Gray and Wess.
3
Formula for Gomori’s Trichrome - Chromotrope 2R (CI 16570): 0.6 g; aniline blue: 0.6 g; phosphomolybdic
acid: 1.0 g; dissolve in distilled water: 100.0 ml; add hydrochloric acid: 1.0 ml. Allow to stand 24 hours in
refrigerator before use. Store in refrigerator and use cold.
4
Be especially careful passing the specimens from absolute ethanol into Creosote. If transfer is slow, the
ethanol dries rapidly and the specimen may dry.
This method was proposed by KRITSKY et al. (1978) and is extremely simple:
1. place one drop of Gomori’s trichrome on the center of a glass slide;
2. place the specimen you want to study in this drop;
3. Allow the specimen to stain for 30 s to 1 min;
4. Remove the excess of stain with a tissue paper (make a tip by twisting the paper
between your fingers and be careful not to remove the specimen);
5. Rapidly place a drop of Gray and Wess (not Hoyer’s) on the specimen;
6. Cover with a coverslip and put the slide in a warmer or oven (about 57 °C) for
1-2 days.
This slide can be studied under light microscope. As Gray and Wess is a water-base
stain, it will remove the excess stain. Sclerites stain blue, red or a mixture of both colors,
making small and very thin and delicate parts visible. This technique is fundamental for
the study of the morphology of the shield of the superficial bar of viviparous
Gyrodactylidae.
The keys that follow are intended solely as a support for the initial process of
determination of the taxa of specimens under study. Generic determination should be
checked using the diagnoses presented in the next part. There are no keys for species,
however. Species determination depends on the study of original literature based on the
list of species available for each genus and the list of references of the chapter. The
numbering of hooks used herein is that proposed by MIZELLE (1936). In this system,
numbering of hook pairs proceeds in an anterior-posterior direction ventrally and
posterior-anterior direction dorsally (when dorsal hooks are present) (examples in Figs. 3-
4A; 3-13A; 3-14A). Keys were prepared with the software DELTA Editor (DALLWITZ,
1980) and Key (DALLWITZ, 1974).
Key to higher taxa of freshwater Amazonian Monogenoidea
1. Haptor armed with a combination of hooks, anchors, bars, spines, squamodisc

................................................................................................................. (Polyonchoinea)..2
– Haptor armed with haptoral suckers or clamps ............................................................
................................................................................. (Heteronchoinea, Oligonchoinea)..4
2(1). 16 hooks, hinged, all marginal in a palmate, sucker-like haptor; viviparous or
oviparous worms. ...................................................................................... Gyrodactylidae
– 14 hooks, not hinged, on a haptor of variable morphology; oviparous worms .. 3
3(2). Haptor sucker-like, crossed by septa; 14 hooks marginal; a single pair of ventral
anchors; bars absent; parasites of potamotrygonid rays .............................................
.................................................................................... Monocotylidae (Potamotrygonocotyle)
– Haptor sucker-like or otherwise, never with septa; 14 hooks; anchors, bars, and
other haptoral sclerites present or absent; parasites of Teleostei ...... Dactylogyrinea
4(1). Haptor armed with multiple clamps; parasites of Teleostei .......................................

.................................................................................................. Microcotylidae (Paranaella)
– Haptor with 6 haptoral suckers, each with haptoral sclerite; parasites of
potamotrygonid rays ............................................ Hexabothriidae (Paraheteronchocotyle)
Key to genera of freshwater Amazonian Gyrodactylidae
1. Oviparous worms ............................................................................................................... 2

– Viviparous worms .............................................................................................................. 5
2(1). Male copulatory organ (MCO) without accessory piece ........................................... 3
– Copulatory organ composed of MCO and accessory piece (Fig. 3-8B) ..................
..................................................................................................................... Nothogyrodactylus
3(2). Eggs in uterus more than 2, usually up to 10 or more ...................... Phanerothecium
– Eggs in uterus never more than one ............................................................................. 4
4(3). Muscular MCO without spines .................................................................. Oogyrodactylus
– Muscular MCO with spines ........................................................................... Hyperopletes
5(1). Ventral anchors, bars absent .....................................................................Anacanthocotyle
– Ventral anchor, bars (superficial and deep) present ................................................... 6
6(5). Cylindrical haptoral sclerite absent ................................................................................. 7
– Cylindrical haptoral sclerite present (Fig. 3-4B) .......................................... Accessorius
7(6). Superficial (anterior) bar with shield (Fig. 3-6E) or lacking posterior projections
.............................................................................................................................. Gyrodactylus
– Superficial (anterior) bar with double posterior projections similar to ribbons (Fig.
3-11B) .................................................................................................................. Scleroductus
Key to genera of freshwater Amazonian Dactylogyrinea
1. Dorsal bar absent ............................................................................................................... 2

– Dorsal bar present, double ............................................................................................... 9
– Dorsal bar present, single ............................................................................................... 10
2(1). Hooks 14 on posterior margin of haptor ..................................................................... 3
– Hooks 14, 12 ventral, on posterior margin of haptor, 2 more central .................. 5
– Hooks 14 marginal ..................................................................................... Anacanthoroides
– Hooks 14, with anacanthorine distribution (6 dorsal, 8 ventral) ......... Anacanthorus
– Hooks 14, with ancyrocephaline distribution (4 dorsal, 10 ventral) ....................... 6
3(2). Haptor well set off from trunk ...........................................................................Kritskyia
– Haptor not differentiated from trunk ........................................................................... 4
4(3). Accessory piece one, non articulated to MCO ........................................... Pavanelliella
– Accessory piece articulated directly to MCO by copulatory ligament ..... Telethecium
5(2). Ventral bar with smooth surface on anterior margins; gonads tandem; germarium
anterior to testis; accessory piece articulated to MCO by copulatory ligament;
hook shank simple .......................................................................................... Cacatuocotyle
– Ventral bar with two submedian elongate processes; gonads overlapping;

accessory piece articulated directly to MCO; hook shank divided in two portions,
clearly defined .................................................................................................... Rhinonastes
6(2). Vagina ventral ...................................................................................................................... 7
– Vagina dextral, marginal ................................................................................................... 8
– Vagina sinistral, marginal .................................................................................. Rhinoxenus
7(6). Accessory piece one, not articulated to MCO; hook shank simple; ventral anchor
rod shaped ......................................................................................................... Trinidactylus
– Accessory piece articulated directly to MCO; hook shank divided in two portions,
clearly defined; ventral anchor with point, shaft, roots well defined ........................
.......................................................................................................................... Monocleithrium
– Accessory piece articulated to MCO by copulatory ligament; hook with entire
shank inflated, conferring a robust morphology to hook; ventral anchor with
point, shaft and roots not defined ..................................................................... Unibarra
8(6). Gonads tandem; testes multiple, anterior to germarium; ventral anchor robust,
with deep root longer than superficial ................................................ Linguadactyloides
– Gonads overlapping; testis single; ventral anchor with point, shaft and roots not
well defined ........................................................................................................... Trinigyrus
– Gonads tandem; germarium anterior to single testis; ventral anchor with point,
shaft, roots well defined ............................................................................ Euryhaliotrema
9(1). Vagina dextral, marginal; dorsal anchor modified, base apparently greatly
elongate, distorted, sometimes hook-like; ventral bar strongly V-shaped ................
........................................................................................................................ Curvianchoratus
– Vagina sinistral, marginal; dorsal anchor with shaft, point and base not clearly
separated in roots; ventral bar lip-shaped ................................................... Diplectanum
– Vagina dextral, ventral; dorsal anchor with shaft, point and base composed of two
roots; ventral bar slightly V-shaped ............................................................. Trinibaculum
10(1). Ventral bar with smooth surface on anterior margins ............................................. 11
– Ventral bar with short sclerotized muscle-attachment ........................ Euryhaliotrema
– Ventral bar with slightly projected margin, with or without medial cleft (Fig. 3-2B)
.............................................................................................................................................. 34
– Ventral bar with anterior transversal groove .............................................................. 35
– Ventral bar with antero-median projection ................................................................ 36
– Ventral bar with double umbeliform membranes on anterior margin .....................
.......................................................................................................................... Sciadicleithrum
11(10). Ventral anchor with point, shaft and roots not well defined .................................. 12
– Ventral anchor with point, shaft, base very elongated with roots not well defined
......................................................................................................................... Protorhinoxenus
– Ventral anchor with point, shaft, roots well defined ................................................ 15
– Ventral anchor robust, with elongate superficial and deep roots ..................... Jainus
12(11). Gonads overlapping ......................................................................................................... 13
– Gonads tandem; germarium anterior to testis ........................................................... 14
13(12). Eyes four; MCO coiled; hook pair 5 significantly distinct from other pairs; ventral
anchor filament robust, very conspicuous ........................................................ Gussevia
– Eyes absent; MCO somewhat straight or slightly curved; hook pair 5 similar to
other pairs; ventral anchor filament delicate, often inconspicuous ......... Heterotylus
14(12). Hooks 14, 8 dorsal, 6 ventral; anchors two pairs, dorsal ............................... Unilatus
– Hooks 14, with ancyrocephaline distribution (4 dorsal, 10 ventral); anchors ventral
pair, dorsal pair ............................................................................................ Demidospermus
15(11). MCO J-shaped .................................................................................................................. 16
– MCO sinuous .............................................................................................. Aphanoblastella
– MCO somewhat straight or slightly curved ............................................................... 18
– MCO coiled ....................................................................................................................... 27
16(15). Gonads overlapping; seminal vesicle C-shaped ......................................... Notothecium
– Gonads tandem; germarium anterior to testis; seminal vesicle fusiform ............ 17
17(16). Posterior projections on dorsal bar one ................................................. Demidospermus
– Posterior projections on dorsal bar two, ribbon-like ......................... Cosmetocleithrum
18(15). Seminal vesicle C-shaped ................................................................................................ 19
– Seminal vesicle fusiform ................................................................................................. 20
19(18). Vagina dextral, dorsal; dorsal bar strongly V-shaped ................................ Notothecium
– Vagina sinistral, dorsal; dorsal bar slightly U-shaped ............................. Enallothecium
20(18). MCO single ....................................................................................................................... 21
– MCO double .................................................................................................................... 25
21(20). Anterior projections of dorsal bar absent .................................................................. 22
– Anterior projections of dorsal bar with small subterminal knob at each extremity
............................................................................................................................Philocorydoras
– Anterior projections of dorsal bar with single median projection .........................
......................................................................................................................... Mymarothecium
22(21). Vagina double ................................................................................................ Amphithecium
– Vagina single ...................................................................................................................... 23
23(22). Vagina middorsal, looping left caecum ..................................................... Notothecioides
– Vagina sinistral marginal ................................................................................................. 24
– Vagina dextral, dorsal ................................................................................. Mymarothecium
24(23). Gonads overlapping; dorsal bar slightly V-shaped; posterior projections on dorsal
bar absent ....................................................................................................... Calpidothecium
– Gonads tandem; germarium anterior to testis; dorsal bar strongly V-shaped; single
posterior projection on dorsal bar ........................................................... Demidospermus
25(20). Vagina double ................................................................................................ Amphithecium
– Vagina single ...................................................................................................................... 26
26(25). Vagina sclerotized, dextral, marginal; ventral bar slightly U-shaped; tegument scaled
........................................................................................................................ Pithanothecium
– Vagina muscular, sinistral, dorsal; ventral bar rod-shaped; tegument smooth ........
............................................................................................................................ Heterothecium
27(15). Posterior projection on dorsal bar absent ................................................................... 28
– Posterior projection on dorsal bar one ........................................................................ 33
– Posterior projections on dorsal bar two, ribbon-like ......................... Cosmetocleithrum
28(27). Ventral bar without posterior median projection ...................................................... 29
– Ventral bar with posterior median projection ............................................................ 32
29(28). Accessory vaginal sclerite absent .................................................................................. 30

– Accessory vaginal sclerite present ................................................................Urocleidoides
30(29). Accessory sclerite associated with base of ventral anchor absent ........................ 31
– Accessory sclerite associated with base of ventral anchor present ......................
........................................................................................................................... Tereancistrium
31(30). Hook pair 5 similar to other pairs; ventral anchor filament delicate, often
inconspicuous; gonadal bar present .......................................................... Gonocleithrum
– Hook pair 5 significantly distinct from other, reduced; ventral anchor filament
robust, very conspicuous; gonadal bar absent ................................................. Gussevia
32(28). Eyes four; accessory piece one, non articulated to MCO; vagina sinistral, marginal;
dorsal bar slightly V-shaped ..................................................................... Aphanoblastella
– Eyes absent; accessory piece articulated to MCO by copulatory ligament; vagina
sinistral, ventral; dorsal bar slightly U-shaped ................................... Pseudovancleaveus
33(27). Gonads overlapping; ventral bar rod-shaped; accessory sclerite associated with
base of ventral anchor present ................................................................... Tereancistrium
– Gonads tandem; germarium anterior to testis; ventral bar strongly V-shaped;
accessory sclerite associated with base of ventral anchor absent ...........................
......................................................................................................................... Demidospermus
34(10). Eyes four; vagina sclerotized, single; dorsal bar slightly U-shaped ......... Odothecium
– Eyes two; vagina muscular, double; dorsal bar slightly V-shaped ..............................
....................................................................................................................... Calpidothecioides
35(10). Prostatic reservoirs unknown; hook pair 5 similar to other pairs; ventral anchor
filament delicate, often inconspicuous .................................................... Ancistrohaptor
– Prostatic reservoirs round to short, ovate; hook pair 5 significantly distinct from
other, reduced; ventral anchor filament robust, very conspicuous ............. Gussevia
36(10). Accessory piece one, non articulated to MCO .......................................................... 37
– Accessory piece articulated directly to MCO ............................................................ 39
– Accessory piece articulated to MCO by copulatory ligament ................................ 41
37(36). Eyes four; prostatic reservoirs round to ovate; annulations on body absent;
seminal vesicle with thin walls, of variable length .................................................... 38
– Eyes absent; prostatic reservoirs very long, often looping posteriorly; annulations
on body present; seminal vesicle with thick walls, very long, extending from level
of base of MCO to initial portion of germarium ..................................... Vancleaveus
38(37). Ventral anchor with point, shaft, roots well defined; ventral bar slightly V-shaped,
without any posterior median projection ............................................... Diaphorocleidus
– Ventral anchor robust, with elongate superficial and deep roots; ventral bar rod-
shaped, with posterior median projection ............................................................. Jainus
39(36). MCO J-shaped ............................................................................................... Notozothecium
– MCO somewhat straight or slightly curved .......................................... Mymarothecium
– MCO coiled ....................................................................................................................... 40
40(39). Seminal vesicle two, fusiform; ventral bar rod-shaped; gonadal bar absent ...........
...............................................................................................................................Dawestrema
– Seminal vesicle fusiform; ventral bar slightly U-shaped; gonadal bar present .......
........................................................................................................................... Gonocleithrum
41(36). MCO J-shaped .................................................................................................................. 42

– MCO somewhat straight or slightly curved .......................................... Mymarothecium
– MCO coiled .................................................................................................... Ameloblastella
42(41). Vagina sinistral marginal; dorsal anchor with shaft, point and base composed of
two roots ...................................................................................................Annulotrematoides
– Vagina dextral, dorsal; dorsal anchor robust with deep root longer than superficial
root ................................................................................................................... Notozothecium
Clave para las taxas superiores de Monogenoidea Neotropicales de agua dulce
1. Haptor armado con una combinación de ganchos, áncoras, barras, espinas,

escuamodiscos ..................................................................................... (Polyonchoinea)..2
– Haptor armado con ventosas haptoriales o tenazas .....................................................
................................................................................. (Heteronchoinea, Oligonchoinea)..4
2(1). 16 ganchos, articulados, todos marginales en un haptor palmeado y con forma de
ventosa; helmintos vivíparos u oviparos .............................................. Gyrodactylidae
– 14 ganchos, sin articulación; haptor de morfología variable; helmintos ovíparos
................................................................................................................................................ 3
3(2). Haptor como ventosa, cruzado por un septo; 14 ganchos marginales; un único par
de áncoras ventrales; barras ausentes; parásitos de rayas potamotrygonideas ........
.................................................................................... Monocotylidae (Potamotrygonocotyle)
– Haptor como ventosa o no, nunca con septos; 14 ganchos; áncoras, barras y otras
escleritas presentes o ausentes en el raptor; parásitos de Teleostei .........................
....................................................................................................................... Dactylogyrinea
4(1). Haptor armado con múltiples tenazass; parásitos de Teleostei .................................
.................................................................................................. Microcotylidae (Paranaella)
– Haptor con 6 ventosas en el haptor, cada una con escleritas haptorial; parásito de
rayas potamotrygonideas .................................... Hexabothriidae (Paraheteronchocotyle)
Clave para los géneros de Gyrodactylidae Neotropicales de agua dulce
1. Helmintos ovíparos ............................................................................................................ 2

– Helmintos vivíparos ........................................................................................................... 5
2(1). Organo copulador masculino (OCM) simple sin piezas accesorias. ....................... 3
– Organo copulador compuesto por OCM y pieza accesoria (Fig. 3-8B) ..................
..................................................................................................................... Nothogyrodactylus
3(2). Más de 2 huevos dentro del útero, generalmente hasta 10 ................ Phanerothecium
– Nunca más de 1 huevos dentro del útero..................................................................... 4
4(3). OCM muscular sin espinas ......................................................................... Oogyrodactylus
– OCM muscular con espinas ........................................................................... Hyperopletes
5(1). Ancoras ventrales presentes, barras ausentes ........................................ Anacanthocotyle
– Ancoras ventrales presentes, barras (superficiales y profundas) presentes ............ 6
6(5). Esclerita haptorial cilíndrica ausente .............................................................................. 7

– Esclerita haptorial cilíndrica presente (Fig. 3-4B) ....................................... Accessorius
7(6). Barra ventral (anterior) con escudo (Fig. 3-6E) o sin proyecciones posteriores ....
.............................................................................................................................. Gyrodactylus
– Barra ventral (anterior) con proyecciones posteriores dobles similares a cintas
(Fig. 3-11B) ......................................................................................................... Scleroductus
Clave para los Dactylogyrinea Neotropicales de agua dulce
1. Barra dorsal ausente .......................................................................................................... 2

– Barra dorsal presente, doble ............................................................................................ 9
– Barra dorsal presente, única ........................................................................................... 10
2(1). 14 ganchos en el margen posterior del haptor ............................................................ 3
– 14 ganchos, 12 ventrales, en el margen posterior del haptor, 2 más centrales ..... 5
– 14 ganchos marginales .............................................................................. Anacanthoroides
– 14 ganchos, con distribución anacantorina (6 dorsales, 8 ventrales) ...... Anacanthorus
– 14 ganchos, con distribución ancirocefalina (4 dorsales, 10 ventrales) .................. 6
3(2). Haptor bien diferenciado del tronco .................................................................Kritskyia
– Haptor no diferenciado del tronco ................................................................................ 4
4(3). Pieza accesoria único, no-articulada al OCM .............................................. Pavanelliella
– Pieza accesoria directamente articulada al órgano copulador masculino (OCM)
por el ligamento copulador .............................................................................. Telethecium
5(2). Barra ventral con superficie lisa en el margen anterior; gónadas en hilera;
germarium anterior a los testículos; pieza accesoria articulada al OCM por el
ligamento copulatorio; cuerpo del gancho simple. .................................. Cacatuocotyle
– Barra ventral con dos procesos submedianos alargados; gónadas sobrepuestas; la
pieza accesoria articulada directamente al OCM; parte final del gancho dividido
en dos porciones claramente definidas ........................................................ Rhinonastes
6(2). Vagina ventral ...................................................................................................................... 7
– Vagina derecha, marginal .................................................................................................. 8
– Vagina izquierda, marginal ............................................................................... Rhinoxenus
7(6). Pieza accesoria única, no-articulada al OCM; cuerpo del gancho es simple; áncora
ventral en forma de varilla ............................................................................. Trinidactylus
– Pieza accesoria articulada directamente a la OCM; cuerpo del gancho dividido en
dos porciones claramente definidas; áncora ventral con punta, cuerpo y base bien
definidas ......................................................................................................... Monocleithrium
– Pieza accesoria articulada a OCM por el ligamento copulador; gancho con todo
cuerpo inflado, confiriéndole una morfología robusta al gancho; áncora ventral
con punta, cuerpo y base no definida ............................................................... Unibarra
8(6). Gónadas en hilera; testículos múltiples, anteriores al germarium; áncora ventral
robusta con raíz profunda más larga que superficial ....................... Linguadactyloides
– Gónadas sobrepuestas; testículo único; áncora ventral en punta, cuerpo y raíces
base indefinidos .................................................................................................... Trinigyrus
– Gónadas en hilera; germario anterior al único testículo; áncora ventral con punta,
cuerpo y raíces bien definidos .................................................................. Euryhaliotrema
9(1). Vagina dextral, marginal; áncora dorsal modificada, base aparentemente muy
alargada, distorsionada, algunas veces en forma de gancho; barra ventral en
forma de V .................................................................................................. Curvianchoratus
– Vagina sinistral, marginal; áncora dorsal con cuerpo, punta y base sin separación
clara con las raíces; barra ventral en forma de labio ................................ Diplectanum
– Vagina dextral, ventral; áncora dorsal con cuerpo, punta y base compuesto de dos
raíces; barra ventral ligeramente en forma de V ...................................... Trinibaculum
10(1). Barra ventral con superficie lisa en los márgenes anteriores .................................. 11
– Barra ventral con corta proyección muscular esclerotizada .............. Euryhaliotrema
– Barra ventral con margen ligeramente proyectado, con o sin hendidura media (Fig.
3-2B) ................................................................................................................................... 34
– Barra ventral con estría antero-transversal ................................................................. 35
– Barra ventral con proyecciones antero-mediana ....................................................... 36
– Barra ventral con doble membrana umbeliforme en el margen anterior ................
.......................................................................................................................... Sciadicleithrum
11(10). Ancora ventral con punta, cuerpo y raíces sin definición ....................................... 12
– Ancora ventral con punta, cuerpo, base muy alargada con raíces no definidas .....
......................................................................................................................... Protorhinoxenus
– Ancora ventral con punta, cuerpo, raíces bien definidas ......................................... 15
– Ancora ventral robusta, con raíces superficiales y profundas alargadas ......... Jainus
12(11). Gónadas sobrepuestas ..................................................................................................... 13
– Gónadas en hileras; germario anterior al testículo ................................................... 14
13(12). Cuatro ojos; OCM enrollado; el quinto par de ganchos significativamente distinto de
los otros pares; filamento del áncora ventral robusto, muy conspicuo ......... Gussevia
– Ojos ausentes; OCM casi recto o ligeramente curvado; quinto par de ganchos
similar a los otros pares; filamento del áncora ventral delicado, con frecuencia
insconspicuo ........................................................................................................ Heterotylus
14(12). 14 ganchos, 8 dorsales, 6 ventrales; 2 pares de áncoras dorsales ................. Unilatus
– 14 ganchos, con distribución ancirocefalina (4 dorsales, 10 ventrales); par de
áncoras ventrales; par de áncoras dorsales ............................................ Demidospermus
15(11). OCM en forma de J ........................................................................................................ 16
– OCM sinuoso .............................................................................................. Aphanoblastella
– OCM casi recto o ligeramente curvado ...................................................................... 18
– OCM enrollado ................................................................................................................. 27
16(15). Gónadas sobrepuestas; vesícula seminal en forma de C ......................... Notothecium
– Gónadas en hilera; germario anterior al testículo; vesícula seminal fusiforme ......
.............................................................................................................................................. 17
17(16). Proyecciones posteriores en la primera barra dorsal ........................... Demidospermus
– Proyecciones posteriores en la segunda barra dorsal, en forma de cinta ................
....................................................................................................................... Cosmetocleithrum
18(15). Vesícula seminal en forma de C .................................................................................... 19
– Vesícula seminal fusiforme ............................................................................................ 20
19(18). Vagina dextral, dorsal; barra dorsal em forma de V ................................ Notothecium

– Vagina sinestral, dorsal; barra dorsal ligeramente en forma de U ....... Enallothecium
20(18). OCM único ........................................................................................................................ 21
– OCM doble ........................................................................................................................ 25
21(20). Proyecciones anteriores de la barra dorsal ausentes ................................................. 22
–Proyecciones anteriores de la barra dorsal con pequeños nudos subterminales en
cada extremidad ..............................................................................................Philocorydoras
– Proyecciones anteriores de la barra dorsal con proyección media única .................
......................................................................................................................... Mymarothecium
22(21). Vagina doble ................................................................................................... Amphithecium
– Vagina única ...................................................................................................................... 23
23(22). Vagina dorsal media, circundando el intestino izquierdo ...................... Notothecioides
– Vagina sinestral marginal ................................................................................................ 24
– Vagina dextral, dorsal ................................................................................. Mymarothecium
24(23). Gónadas sobrepuestas; barra dorsal ligeramente en forma de V; proyecciones
posteriores ausentes en la barra dorsal .................................................... Calpidothecium
– Gónadas en hilera; germario anterior al testículo; barra dorsal en forma de V;
proyección única posterior en la barra dorsal ....................................... Demidospermus
25(20). Vagina doble ................................................................................................... Amphithecium
– Vagina única ...................................................................................................................... 26
26(25). Vagina esclerotizada, destral, marginal; barra ventral ligeramente en forma de U;
tegumento con escamas .............................................................................. Pithanothecium
– Vagina muscular, sinestral, dorsal; barra ventral en forma de varilla; tegumento liso
............................................................................................................................ Heterothecium
27(15). Proyección posterior ausente en la barra dorsal ........................................................ 28
– Proyección posterior en la primera barra dorsal ....................................................... 33
– Proyecciones posteriores en la segunda barra dorsal, en forma de cinta ................
....................................................................................................................... Cosmetocleithrum
28(27). Barra ventral sin proyección mediana posterior ........................................................ 29
– Barra ventral con proyección mediana posterior ...................................................... 32
29(28). Esclerita accesoria vaginal ausente ............................................................................... 30
– Esclerita accesoria vaginal presente .............................................................Urocleidoides
30(29). Esclerita accesoria asociada a la base del áncora ventral ausente .......................... 31
– Esclerita accesoria asociada a la base del áncora ventral presente ......................
........................................................................................................................... Tereancistrium
31(30). Par de ganchos 5 similar a los otros pares; filamento del áncora ventral delicado,
con frecuencia incospicuo; barra gonadal presente ............................... Gonocleithrum
– Par de ganchos 5 significativamente distinto de los otros, reducido; filamento del
áncora ventral robusto, muy conspicuo; barra gonadal ausente .................. Gussevia
32(28). Cuatro ojos; primera pieza accesoria no articulada al OCM; vagina sinestral,
marginal; barra dorsal ligeramente en forma de V ............................. Aphanoblastella
– Ojos ausentes; pieza accesoria articulada al OCM por ligamento copulatorio; vagina
sinistral, ventral; barra dorsal ligeramente en forma de U ................ Pseudovancleaveus
33(27). Gónadas sobrepuestas; barra ventral en forma de varilla; esclerita accesoria

asociada a la base del áncora ventral presente ........................................ Tereancistrium
– Gónadas en hilera; germarium anterior al testículo; barra ventral en forma de
V; esclerita accesoria asociada a la base del áncora ventral ausente ...................
......................................................................................................................... Demidospermus
34(10). Cuatro ojos; vagina esclerotizada, única; barra dorsal ligeramente en forma de U
................................................................................................................................ Odothecium
– Dos ojos; vagina muscular, doble; barra dorsal ligeramente en forma de V ..........
....................................................................................................................... Calpidothecioides
35(10). Reservorios prostáticos desconocidos; par de ganchos 5 similar a los otros
pares; filamento del áncora ventral delicado, frecuentemente incospicuo ...........
.......................................................................................................................... Ancistrohaptor
– Reservorios prostáticos cortos, redondos u ovalados; par de ganchos 5
significativamente diferentes de los otros, reducidos; filamento del áncora ventral
robusto, muy conspicuo ........................................................................................ Gussevia
36(10). Pieza accesoria unica no articulada al OCM .............................................................. 37
– Pieza accesoria articulada directamente al OCM ...................................................... 39
– Pieza accesoria articulada al OCM por el ligamento copulatorio .......................... 41
37(36). Cuatro ojos; reservorios prostáticos redondos a ovalados; cuerpo sin anillos;
vesícula seminal con paredes delgadas de longitud variable ................................... 38
– Ojos ausentes; reservorios prostáticos muy largos, frecuentemente dando la vuelta
posteriormente; anillos presentes en el cuerpo; vesícula seminal con paredes
gruesas, muy larga, extendiendose desde el nível de la base del OCM hasta la parte
inicial del germarium ........................................................................................ Vancleaveus
38(37). Ancora ventral con punta, cuerpo, raíces bien definidas; barra ventral ligeramente
en forma de V, sin ninguna proyección posterior mediana ............... Diaphorocleidus
– Ancora ventral robusta, con raíces superficiales y profundas alargadas; barra
ventral en forma de varilla, con proyección posterior mediana ........................ Jainus
39(36). OCM en forma de J ..................................................................................... Notozothecium
– OCM casi recto o ligeramente curvado ................................................. Mymarothecium
– OCM enrollado ................................................................................................................. 40
40(39). Dos vesículas seminales, fusiformes; barra ventral em forma de varilla; barra
gonadal ausente ................................................................................................. Dawestrema
– Vesícula seminal fusiforme; barra ventral ligeramente en forma de U; barra
gonadal presente ............................................................................................ Gonocleithrum
41(36). OCM en forma de J ........................................................................................................ 42
– OCM casi recto o ligeramente curvo ..................................................... Mymarothecium
– COM enrollado .............................................................................................. Ameloblastella
42(41). Vagina sinestral marginal; áncora dorsal con cuerpo, punta y base compuesta de
dos raíces ...................................................................................................Annulotrematoides
– Vagina destral, dorsal; áncora dorsal robusta con raíz profunda más larga que la
raíz superficial ................................................................................................ Notozothecium
VII. Checklist of Monogenoidea from Amazonian freshwater fishes and diagnoses
The diagnoses and list of species below refer solely to those species of Mono-
genoidea from the Amazon Region or from fish groups (species, genera) that are
known to occur in this region, even though some of the species are originally
reported from other areas. A list of species of genera of uncertain taxonomic validity
is presented at the end of this section. The descriptions were prepared with the
software DELTA Editor (DALLWITZ, 1980).
Subclass Heteronchoinea BOEGER & KRITSKY, 2002

Infrasubclass Oligonchoinea BYCHOWSKY, 1937
Hexabothriidae PRICE, 1942
Body elongate, fusiform. Oviparous. Spike sensilla absent. Eyes, eye
granules absent. Mouth terminal; oral sucker present, pharynx bulbous.
Ceca non-confluent, with diverticula. Copulatory organ (MCO) muscular,
armed with spines or not. Testes multiple, post-germarian. Germarium
elongate, U-shaped; genito-intestinal canal present. Vagina two, bilateral,
ventral; ductus vaginalis. Haptor round to ovate, armed with three pairs of
haptoral suckers with hook-like sclerites; haptoral appendix present with
pair of suckers, hooks, pair of anchors; hooks present or absent in adult;
anchors present or absent.
Paraheteronchocotyle MAYES, BROOKS & THORSON, 1981
Body elongate. MCO muscular, composed of two portions; distal portion
elongate, unarmed; proximal portion elongate with thick walls; prostatic
region absent. Proximal germarium lobate, descending branch sinuous,
ascending branch absent; oviduct dilated. Ootype smooth, lacking longitu-
dinal rows of cells. Haptor asymmetrical; haptoral sucker distributed
linearly, except a pair adjacent to haptoral appendix. Haptoral appendix
marginal, anchors absent. Eggs with one long filament. Parasites of species
of Potamotrygonidae rays.
Type and only species: Paraheteronchocotyle amazonensis MAYES, BROOKS &
THORSON, 1981 from Potamotrygon circularis. (Figs. 3-1 A-C).
Microcotylidae TASCHENBERG, 1879
Body elongate, fusiform. Oviparous. Spike sensilla absent. Eyes, eye granules
absent. Mouth terminal; buccal organs present, pharynx bulbous. Ceca non-
confluent with diverticula. Copulatory organ (MCO) present or absent,
genital atrium muscular, usually armed with spines. Testes multiple, post-
germarian. Germarium elongate, double-inverted-U-shaped; genito-intesti-
nal canal present. Vagina dorsal; ductus vaginalis. Haptor elongate, armed
with multiple microcotylid clamps (n>8); hooks, anchors, bars absent.
Paranaella KOHN, BAPTISTA-FARIAS & COHEN, 2000
Body lanceolate. Haptor subsymmetrical with two subequal rows of
clamps, similar in shape. Buccal organs with septum. MCO non-differenti-
ated. Genital atrium armed with two concentric rows of spines around
muscular aperture. Eggs with one filament each. Vagina mid-dorsal,

unarmed. Parasites of gills of species of Loricariidae (Siluriformes).
Type and only species: Paranaella luquei KOHN, BAPTISTA-FARIAS & COHEN,
2000 from Hypostomus sp., Rhinelepis aspera, Hypostomus regani. (Figs. 3-2 A-B).
Subclass Polyonchoinea
Monocotylidae TASCHENBERG, 1879
Body robust to fusiform. Oviparous. Spike sensilla absent. Eyes present or
absent. Haptor sucker-shaped, usually divided by septa. Oral sucker present.
Pharynx single, bulbous. Ceca non-confluent. Copulatory organ (MCO)
sclerotized, lacking accessory piece. Germarium loops right intestinal ceca.
Vagina ususally present. Testis single, post-germarian. Hooks dactylogyrid,
14 marginal. Ventral anchor present or absent. Dorsal anchor, bars absent.
Potamotrygonocotyle MAYES, BROOKS & THORSON, 1981
Eyes absent; eye granules present or absent. Tegument smooth. Vagina
sinistro-marginal, muscular.. Hooks with simple shank. Ventral anchor pair
present. Haptor divided ventrally into eight external loculi, one central; septa
lacking sinuous ridges or sclerites; dorsal surface of haptor with six muscular
papillae. Parasites of the gills of Potamotrygonidae species (Rajiformes).
Type and only species: P. tsalickisi MAYES, BROOKS & THORSON, 1981 from
Potamotrygon circularis. (Figs. 3-3 A-B).
Gyrodactylidae VAN BENEDEN & HESSE, 1863
Oviparous or viviparous. Spike sensilla present. Eyes granules absent.
Haptor sucker-shaped. Pharynx two bulbous subunits, distal frequently with
finger-like projections. Ceca non-confluent. Copulatory organ (MCO) mus-
cular or sclerotized; when muscular it may be elongate, form a cirrus sac, or
be bulbous. Accessory piece present, absent. Germarium usually ovate, with
an internal fertilization chamber. Vagina absent. Hooks gyrodactylid. Hooks
16 marginal. Hooks on finger-like projections, or not on peduncles. Ventral
anchor present or absent. Dorsal anchor, bar absent. Ventral anchor
gyrodactylid (long superficial root, knob-like deep root). Deep bar (associat-
ed with knobs of anchors) present when ventral anchor is present.
Accessorius JARA, AN & CONE, 1991
Viviparous. Peduncle conspicuous. Haptor sucker-shaped. Copulatory organ
only represented by MCO. MCO muscular, bulbous, with spines. Ventral
anchor present. Cylindrical haptoral sclerite present. Superficial bar present,
rod-shaped. Deep bar (associated with knobs of anchors) present. Parasites
of external body surface of species of Characidae (Characiformes).
Type and only species: Accessorius peruensis JARA, AN & CONE, 1991 from
Lebiasina bimaculata (Figs. 3-4 A-F).
Anacanthocotyle KRITSKY & FRITTS, 1970
Viviparous. Peduncle conspicuous. Haptor cylindrical or sucker-shaped.
Copulatory organ represented only by MCO, muscular, bulbous, with spines.
Ventral anchor, bars absent. Cylindrical haptoral sclerite absent. Parasites of
the external surface of species of Characidae (Characiformes).
Type and only species: A. anacanthocotyle KRITSKY & FRITTS, 1970 from
Astyanax fasciatus (Figs. 3-5 A-C).
Gyrodactylus NORDMANN, 1832
Synonyms: Paragyrodactyloides; Paragyrodactylus.
Viviparous. Peduncle conspicuous. Copulatory organ represented only by
MCO, muscular, bulbous, with spines. Cylindrical haptoral sclerite absent.
Ventral anchor present. Superficial bar present, rod-shaped, with or without
a shield. Parasites of gills and external surfaces of species of Clupeiformes,
Cyprinodontiformes, Perciformes, Siluriformes, Characiformes.
G. anisopharynx POPAZOGLO & BOEGER, 2000 from Corydoras paleatus, Corydoras
ehrhardtii (Callichthyidae, Siluriformes).
G. bimaculatus AN, JARA & CONE, 1991 from Lebiasina bimaculata (Characidae,
Characiformes).
G. bullatarudis TURNBULL, 1956 from Poecilia sphenops, Poecilia reticulata (Poecili-
idae, Cyprinodontiformes).
G. costaricensis KRITSKY & FRITTS, 1970 from Poecilia sphenops (Poeciliidae,
Cyprinodontiformes).
G. gemini FERRAZ, SHINN & SOMMERVILLE, 1994 from Semaprochilodus taeniurus
(Curimatidae, Characiformes).
G. geophagensis BOEGER & POPAZOGLO, 1995 from Geophagus brasiliensis (Cichl-
idae, Perciformes) (Figs. 3-6 A-E).
G. lebiasinus AN, JARA & CONE, 1991 from Lebiasina bimaculata (Characidae,
Characiformes).
G. milleri HARRIS & CABLE, 2000 from Poecilia caucana (Poecilidae, Cyprinodon-
tiformes).
G. neotropicalis KRITSKY & FRITTS, 1970 from Astyanax fasciatus (Characidae,
Characiformes).
G. pictae CABLE, VAN OOSTERHOUT, BARSON & HARRIS, 2005 from Poecilia picta
(Poeciliidae, Cyprinodontiformes).
G. pimelodellus AN, JARA & CONE, 1991 from Pimelodella yuncensis (Pimelodidae,
Siluriformes).
G. poeciliae HARRIS & CABLE, 2000 from Poecilia caucana (Poecilidae, Cyprin-
odontiformes).
G. samirae POPAZOGLO & BOEGER, 2000 from Corydoras ehrhardtii, Corydoras
paleatus (Callichthyidae, Siluriformes).
G. slendrus AN, JARA & CONE, 1991 from Lebiasina bimaculata (Characidae,
Characiformes).
G. superbus POPAZOGLO & BOEGER, 2000 from Corydoras paleatus, Corydoras
ehrhardtii (Callichthyidae, Siluriformes).
G. trairae BOEGER & POPAZOGLO, 1995 from Hoplias malabaricus (Erythrinidae,
Characiformes).
G. turnbulli HARRIS & LYLES, 1992 from Poecilia reticulata (Poeciliidae, Cyprin-
odontiformes).
Hyperopletes BOEGER, KRITSKY & BELMONT-JÉGU, 1994

Oviparous. Peduncle conspicuous. Copulatory organ represented only by
MCO, muscular, sac-like, with spines. Only one egg in uterus. Ventral anchor
present. Cylindrical haptoral sclerite absent. Superficial bar present, without
posterior median projection. Deep bar (associated with knobs of anchors)
present. Parasites of external surface of species of Loricariidae (Siluriformes).
Type and only species: H. malmbergi BOEGER, KRITSKY & BELMONT-JÉGU,
1994 from Rhineloricaria sp (Figs. 3-7 A-D).
Nothogyrodactylus KRITSKY & BOEGER, 1991
Oviparous. Peduncle conspicuous. Copulatory organ composed of MCO,
accessory piece. Accessory piece one to three, non-articulated. MCO muscu-
lar, elongate, lacking spines. Eggs in uterus never more than one. Ventral
anchor present. Cylindrical haptoral sclerite absent. Superficial bar present,
lacking a shield. Deep bar (associated with knobs of anchors) present.
Parasites of external surface of species of Loricariidae (Siluriformes).
N. amazonicus KRITSKY & BOEGER, 1991 from Ancistrus sp.
N. clavatus KRITSKY & BOEGER, 1991 from Ancistrus sp. (Type species).
N. plaesiophallus KRITSKY & BOEGER, 1991 from Ancistrus sp. (Figs. 3-8 A-E).
Oogyrodactylus HARRIS, 1983
Oviparous. Peduncle conspicuous. Copulatory organ composed only of
MCO, muscular, elongate, sac-like, lacking spines. Eggs in uterus never more
than one. Ventral anchor present. Cylindrical haptoral sclerite absent. Superfi-
cial bar present, lacking a shield. Deep bar (associated with knobs of anchors)
present. Parasites of external surface of species of Loricariidae (Siluriformes).
Type and only species: O. farlowellae HARRIS, 1983 from Farlowella amazona
(Figs. 3-9 A-D).
Phanerothecium KRITSKY & THATCHER, 1977
Oviparous. Peduncle conspicuous. Copulatory organ composed only of
MCO, muscular, sac-like, with or without spines. Eggs in uterus more than
2, usually up to 10. Ventral anchor present. Cylindrical haptoral sclerite
absent. Superficial bar present, lacking a shield. Deep bar (associated twith
knobs of anchors) present. Parasites of external surface of species of
Pimelodidae and Loricariidae (Siluriformes).
P. caballeroi KRITSKY & THATCHER, 1977 from Cephalosilurus zungaro (Pimelo-
didae) (Type species) (Fig. 3-10 A-E).
P. harrisi KRITSKY & BOEGER, 1991 from Plecostomus plecostomus (Loricariidae)
P. spinatus BOEGER, KRITSKY & BELMONT-JÉGU, 1994 from Hypostomus punctatus
(Loricariidae).
P. sp. (= P. caballeroi forma major of KRITSKY & THATCHER, 1977) from
Cephalosilurus zungaro (Pimelodidae).
Scleroductus JARA & CONE, 1989
Viviparous. Peduncle conspicuous. Copulatory organ composed only of
MCO, muscular, bulbous, with spines; one spine significantly longer, serving
as guide for distal vas deferens. Ventral anchor present. Cylindrical haptoral
sclerite absent. Superficial bar present, with double posterior projections

similar to ribbons. Deep bar (associated with knobs of anchors) present.
Parasites of gills and external surface of species of Siluriformes.
S. spp. (of KRITSKY, BOEGER & POPAZOGLO, 1995) from Glanidium melanopter-
um (Auchenipteridae), Pimelodella sp. (Pimelodidae), Parauchenipterus striatu-
lus (Auchenipteridae), Rhamdia quelen (Pimelodidae).
S. yuncensi JARA & CONE, 1989 from Pimelodella yuncensis (Pimelodidae) (Type
species) (Figs. 3-11 A-F).
Order Dactylogyrinea BYCHOWSKY, 1937
Suborder Dactylogyridea BYCHOWSKY, 1937
Diplectanidae MONTICELLI, 1903
Diplectanum DIESING, 1858
Body fusiform. Tegument scaled or smooth. Eyes four or two; accessory
granules present or absent. Pharynx bulbous, one subunit. Ceca non-
confluent. Gonads tandem; germarium anterior to testis, looping right
intestinal caecum. Testis single. Copulatory organ composed only of MCO, or
MCO and accessory piece. MCO J-shaped, or sinuous, or somewhat straight,
or slightly curved. Accessory piece one, when present, non-articulated.
Seminal vesicle fusiform. Prostatic reservoirs short, round to ovate. Germari-
um looping right caecum. Vagina sinistro-marginal, sclerotized or muscular.
Squamodisc present. Hooks dactylogyrid, with shank simple; 14, with ancyro-
cephaline distribution (4 dorsal, 10 ventral). Ventral, dorsal anchor pairs
present. Ventral anchor with point, shaft, roots well defined. Dorsal anchor
with shaft, point, base not clearly separated into roots. Ventral bar lip-shaped;
anterior projection absent, posterior projection present or absent. Groove on
entire length of ventral bar present. Dorsal bar present, double, rod-shaped.
Parasites of the gills of species of Perciformes (in Neotropical freshwater,
only species of Sciaenidae are known to host diplectanids).
D. decorum KRITSKY & THATCHER, 1984 from Plagioscion squamosissimus (Figs. 3-
12 A-F).
D. gymnopeus KRITSKY & THATCHER, 1984 from Plagioscion squamosissimus.
D. hilum KRITSKY & THATCHER, 1984 from Plagioscion sp.
D. pescadae KRITSKY & THATCHER, 1984 from Plagioscion sp., Plagioscion squamosis-
simus.
D. piscinarius KRITSKY & THATCHER, 1984 from Plagioscion squamosissimus.
Dactylogyridae BYCHOWSKY, 1933
Ameloblastella KRITSKY, MENDONZA-FRANCO & SCHOLZ, 2000
Synonyms: Cleidodiscus (in part); Vancleaveus (in part); Urocleidoides (in part)
Body fusiform. Tegument smooth; annulations on body absent. Eyes absent;
eye granules present. Pharynx bulbous, one subunit. Ceca confluent.
Gonads overlapping; testis single. Copulatory organ composed of MCO and
accessory piece. MCO of loose coils, not inflated proximally. Accessory
piece articulated to MCO by copulatory ligament. Seminal vesicle fusiform,
with thin walls, of variable length. Prostatic reservoirs short, round to ovate.
Germarium ovate or elongate. Vagina sinistro-marginal, sclerotized; vaginal

bar, accessory vaginal sclerite absent. Hooks dactylogyrid, shank divided in
two portions clearly defined; 14, with ancyrocephaline distribution (4 dorsal,
10 ventral), not on peduncles; hook pair 2 on the haptor, close to hook pairs
3, 4; hook pair 5 similar to other pairs. Anchors ventral, dorsal pairs. Ventral
anchor with point, shaft, roots well defined; anchor filament delicate, often
inconspicuous; accessory sclerite associated with base of ventral anchor
absent. Dorsal anchor with shaft, point, base composed of two roots; fold
on inner superficial root of dorsal anchor absent. Ventral bar rod-shaped
with antero-median projection; no posterior median projection. Dorsal bar
single, rod-shaped; anterior, posterior projections absent. Parasites of the
gills of species of Pimelodidae (Siluriformes).
A. chavarriai (PRICE, 1938) from Rhamdia sebae, Rhamdia rogersi, Rhamdia quelen.
(Type species) (Figs. 3-13 A-E).
A. mamaevi (KRITSKY & THATCHER, 1976) from Cephalosilurus zungaro.
A. platensis (SURIANO & INCORVAIA, 1995) from Pimelodus claria maculatus.
Amphithecium BOEGER & KRITSKY, 1988
Body strongly flattened or fusiform. Tegument papillated, scaled, or smooth;
annulations on body absent. Eyes four or two; eyes granules absent or
present. Pharynx bulbous, one subunit. Ceca confluent. Gonads overlap-
ping. Copulatory organ composed of MCO, accessory piece; MCO single or
double, somewhat straight or slightly curved rami. Accessory piece articulat-
ed to MCO directly or by copulatory ligament. Seminal vesicle fusiform with
thin walls. Prostatic reservoirs short, round to ovate. Germarium ovate or
elongate. Vagina sclerotized or muscular, double; vaginal apertures dorso-
lateral; vaginal bar, accessory vaginal sclerite absent. Hooks dactylogyrid,
shank divided in two portions clearly defined; 14, with ancyrocephaline
distribution (4 dorsal, 10 ventral), not on peduncles; hook pair 2 on haptor,
close to hook pairs 3, 4; hook pair 5 similar to other pairs. Ventral, dorsal
pairs of anchors. Ventral anchor with point, shaft, well defined roots; anchor
filament delicate, often inconspicuous. Dorsal anchor with shaft, point, base
composed of two roots; fold on inner superficial root absent. Ventral bar
present, slightly V-shaped, slightly U-shaped, or rod-shaped, anterior, poste-
rior projections absent. Dorsal bar slightly V-shaped, slightly U-shaped, or
rod-shaped, anterior, posterior projections absent. Parasites of the gills of
species of Serrasalmidae (Characiformes).
A. brachycirrum BOEGER & KRITSKY, 1988 from Pygocentrus nattereri.
A. calycinum BOEGER & KRITSKY, 1988 from Pygocentrus nattereri (Type species)
(Fig. 3-14 A-F).
A. camelum BOEGER & KRITSKY, 1988 from Pygocentrus nattereri.
A. catalaoensis BOEGER & KRITSKY, 1988 from Pygocentrus nattereri.
A. diclonophallum KRITSKY, BOEGER & JÉGU, 1997 from Serrasalmus elongatus,
Serrasalmus sp., Pristobrycon sp., Serrasalmus compressus, Serrasalmus rhombeus,
Serrasalmus gouldingi.
A. falcatum BOEGER & KRITSKY, 1988 from Pygocentrus nattereri, Pristobrycon sp.,
Serrasalmus elongatus, Serrasalmus compressus, Serrasalmus gouldingi, Serrasalmus
manuelli, Serrasalmus rhombeus, Serrasalmus spilopleura, Serrasalmus sp.
A. junki BOEGER & KRITSKY, 1988 from Pygocentrus nattereri, Serrasalmus
rhombeus.
A. microphallum KRITSKY, BOEGER & JÉGU, 1997 from Pygocentrus nattereri,
Serrasalmus sp.
A. minutum KRITSKY, BOEGER & JÉGU, 1997 from Serrasalmus gouldingi, Pristobry-
con sp., Serrasalmus spilopleura, Pristobrycon eigenmanni (Type species).
A. muricatum KRITSKY, BOEGER & JÉGU, 1997 from Pristobrycon eigenmanni,
Serrasalmus rhombeus, Serrasalmus sp.
A. pretiosum KRITSKY, BOEGER & JÉGU, 1997 from Serrasalmus manueli, Serrasal-
mus gouldingi, Pristobrycon sp.
A. prodotum KRITSKY, BOEGER & JÉGU, 1997 from Catoprion mento, Pristobrycon
striolatus.
A. speirocamarotum KRITSKY, BOEGER & JÉGU, 1997 from Serrasalmus elongatus.
A. unguiculum KRITSKY, BOEGER & JÉGU, 1997 from Serrasalmus spilopleura.
A. verecundum KRITSKY, BOEGER & JÉGU, 1997 from Serrasalmus sp., Pristobrycon
eigenmanni.
Anacanthoroides KRITSKY & THATCHER, 1976
Body strongly flattened. Tegument smooth. Eyes four; eye granules
absent. Pharynx bulbous, one subunit. Ceca confluent. Gonads overlap-
ping. Copulatory organ composed of MCO, accessory piece. MCO coiled.
Accessory piece articulated to MCO by copulatory ligament. Seminal
vesicle fusiform. Germarium ovate, solid. Vagina sclerotized, sinistro-
marginal; vaginal bar, accessory vaginal sclerite absent. Hooks dactylogy-
rid, shank simple; 14 marginal; hook pair 2 on the haptor, close to hook
pairs 3, 4; hook pair 5 similar to other pairs. Two pairs of ventral anchors
modified as 4A hooks (incipient). Bars absent. Parasites from the gills of
species of Prochilodontidae (Characiformes).
Type and only species: A. mizellei KRITSKY & THATCHER, 1976 from
Prochilodus reticulatus (Fig. 3-15 A-C).
Anacanthorus MIZELLE & PRICE, 1965
Body fusiform. Tegument smooth. Eyes four or two; eye granules present or
absent. Pharynx bulbous, one subunit. Ceca confluent. Gonads tandem;
testis anterior to germarium. Copulatory organ composed of MCO, accesso-
ry piece; MCO J-shaped, sinuous, somewhat straight, slightly curved, or
coiled. Accessory piece one, non-articulated or articulated directly to MCO.
Seminal vesicle fusiform. Prostatic reservoirs round to short, ovate. Germa-
rium ovate or elongate; solid. Vagina absent. Hooks dactylogyrid, proximal
portion of shank inflated, divided in two portions clearly defined, or simple.
Hooks 14, with anacanthorine distribution (6 dorsal, 8 ventral). Ventral,
dorsal anchors pairs present modified as 4A hooks (incipient). Bars absent.
Parasites of the gills of species of Characiformes.
A. acuminatus KRITSKY, BOEGER & VAN EVERY, 1992 from Triportheus elongatus,
Triportheus angulatus, Triportheus albus (Characidae).
A. alatus KRITSKY, BOEGER & VAN EVERY, 1992 from Triportheus albus, Triporth-
eus elongatus (Characidae).
A. amazonicus KRITSKY & BOEGER, 1995 from Serrasalmus rhombeus, Serrasalmus
sp., Pristobrycon striolatus (Curimatidae).
A. anacanthorus MIZELLE & PRICE, 1965 from Pygocentrus nattereri (Serrasalmi-
dae) (Type species).
A. andersoni KRITSKY, BOEGER & VAN EVERY, 1992 from Triportheus angulatus
(Characidae).
A. beleophallus KRITSKY, BOEGER & VAN EVERY, 1992 from Pristobrycon eigenman-
ni (Serrasalmidae).
A. bellus KRITSKY, BOEGER & VAN EVERY, 1992 from Triportheus albus, Triporth-
eus elongatus (Characidae) (Figs. 3-17 A).
A. brazilensis MIZELLE & PRICE, 1965 from Pygocentrus nattereri (Serrasalmidae).
A. brevis MIZELLE & KRITSKY, 1969 from Brycon melanopterus (Characidae).
A. calophallus KRITSKY, BOEGER & VAN EVERY, 1992 from Triportheus elongatus
(Characidae) (Fig. 3-17 B).
A. carinatus KRITSKY, BOEGER & VAN EVERY, 1992 from Triportheus angulatus
(Characidae) (Fig. 3-17 C).
A. catoprioni KRITSKY, BOEGER & VAN EVERY, 1992 from Catoprion mento
(Serrasalmidae).
A. chaunophallus KRITSKY, BOEGER & VAN EVERY, 1992 from Triportheus angula-
tus (Characidae).
A. chelophorus KRITSKY, BOEGER & VAN EVERY, 1992 from Triportheus angulatus,
Triportheus sp. (Characidae).
A. cinctus VAN EVERY & KRITSKY, 1992 from Pristobrycon striolatus (Serrasalmi-
dae).
A. cladophallus VAN EVERY & KRITSKY, 1992 from Serrasalmus spilopleura
(Serrasalmidae).
A. colombianus KRITSKY & THATCHER, 1974 from Salminus affinis (Characidae).
A. cornutus KRITSKY, BOEGER & VAN EVERY, 1992 from Triportheus angulatus
(Characidae) (Fig. 3-17 D).
A. crytocaulus VAN EVERY & KRITSKY, 1992 from Pristobrycon striolatus (Serras-
almidae).
A. cuticulovaginus KRITSKY & THATCHER, 1974 from Salminus affinis (Characidae).
A. dipelecinus KRITSKY, BOEGER & VAN EVERY, 1992 from Roeboides myersi
(Characidae) (Fig. 3-17 E).
A. elegans KRITSKY, THATCHER & KAYTON, 1979 from Brycon melanopterus
(Characidae).
A. euryphallus KRITSKY, BOEGER & VAN EVERY, 1992 from Triportheus albus,
Triportheus elongatus, Triportheus angulatus (Characidae) (Fig. 3-17 H).
A. formosus KRITSKY, BOEGER & VAN EVERY, 1992 from Triportheus elongatus,
Triportheus sp. (Characidae).
A. furculus KRITSKY, BOEGER & VAN EVERY, 1992 from Triportheus elongatus
(Characidae) (Fig. 3-17 G).
A. glyptophallus KRITSKY, BOEGER & VAN EVERY, 1992 from Triportheus angulatus
(Characidae) (Fig. 3-17 F).
A. gravihamulatus VAN EVERY & KRITSKY, 1992 from Pristobrycon eigenmanni
Serrasalmus sp. 2, Serrasalmus rhombeus (Serrasalmidae).
A. hoplophallus KRITSKY, BOEGER & VAN EVERY, 1992 from Myleus rubripinnis
(Serrasalmidae) (Fig. 3-17 I).
A. jegui VAN EVERY & KRITSKY, 1992 from Serrasalmus rhombeus, Serrasalmus
spilopleura, Pristobrycon sp., from Serrasalmus sp. 2, Pristobrycon eigenmanni,
Serrasalmus sp. (2n = 58) (Serrasalmidae).
A. kruidenieri KRITSKY, THATCHER & KAYTON, 1979 from Brycon melanopterus
(Characidae).
A. lasiophallus VAN EVERY & KRITSKY, 1992 from Pristobrycon striolatus (Serra-
salmidae).
A. lepyrophallus KRITSKY, BOEGER & VAN EVERY, 1992 from Serrasalmus elongatus,
Serrasalmus sp. (2n = 58), Serrasalmus sp. 1 (Serrasalmidae).
A. lygophallus KRITSKY, BOEGER & VAN EVERY, 1992 from Triportheus angulatus
(Characidae) (Fig. 3-17 M).
A. maltai BOEGER & KRITSKY, 1988 from Pygocentrus nattereri (Serrasalmidae).
A. mastigophallus KRITSKY, BOEGER & VAN EVERY, 1992 from Pristobrycon
eigenmanni (Serrasalmidae) (Fig. 3-17 J).
A. mesocondylus VAN EVERY & KRITSKY, 1992 from Serrasalmus spilopleura,
Pristobrycon eigenmanni, Serrasalmus elongatus, Serrasalmus sp. 1, Pristobrycon sp.
n., Serrasalmus sp. 2 (Serrasalmidae).
A. nanus KRITSKY, BOEGER & VAN EVERY, 1992 from Triportheus angulatus
(Characidae).
A. neotropicalis MIZELLE & PRICE, 1965 from Pygocentrus nattereri (Serrasalmidae).
A. palamophallus KRITSKY, BOEGER & VAN EVERY, 1992 from Pristobrycon sp. n.
(Serrasalmidae).
A. paraspathulatus KRITSKY, BOEGER & VAN EVERY, 1992 from Mylossoma
duriventris (Serrasalmidae).
A. pedanophallus KRITSKY, BOEGER & VAN EVERY, 1992 from Myleus rubripinnis
(Serrasalmidae).
A. pelorophallus KRITSKY, BOEGER & VAN EVERY, 1992 from Triportheus elongatus
(Characidae) (Fig. 3-17 N).
A. penilabiatus BOEGER, HUSAK & MARTINS, 1995 from Piaractus mesopotamicus
(Serrasalmidae).
A. periphallus KRITSKY, BOEGER & VAN EVERY, 1992 from Serrasalmus sp. 1,
Serrasalmus sp. 2 (Serrasalmidae) (Fig. 3-17 L).
A. pithophallus KRITSKY, BOEGER & VAN EVERY, 1992 from Triportheus angulatus
(Characidae) (Fig. 3-17 K).
A. prodigiosus VAN EVERY & KRITSKY, 1992 from Serrasalmus sp., Serrasalmus
rhombeus, Serrasalmus elongatus (Serrasalmidae).
A. quinqueramis KRITSKY, BOEGER & VAN EVERY, 1992 from Triportheus albus,
Triportheus elongatus (Characidae).
A. ramosissimus VAN EVERY & KRITSKY, 1992 from Serrasalmus elongatus (Serra-
salmidae).
A. ramulosus KRITSKY, BOEGER & VAN EVERY, 1992 from Triportheus albus,
A. reginae BOEGER & KRITSKY, 1988 from Pygocentrus nattereri (Serrasalmidae).
A. rondonensis BOEGER & KRITSKY, 1988 from Pygocentrus nattereri (Serrasalmidae).
A. scapanus VAN EVERY & KRITSKY, 1992 from Serrasalmus spilopleura (Serra-
salmidae).
A. sciponophallus VAN EVERY & KRITSKY, 1992 from Serrasalmus sp., Serrasalmus
elongatus, Serrasalmus rhombeus, Serrasalmus spilopleura (Serrasalmidae) (Figs.
3-16 A-E).
A. serrasalmi VAN EVERY & KRITSKY, 1992 from Pristobrycon sp. n., Serrasalmus
sp., Serrasalmus elongatus, Serrasalmus rhombeus, Serrasalmus spilopleura (Serra-
salmidae).
A. spathulatus KRITSKY, THATCHER & KAYTON, 1979 from Colossoma macropo-
mum, Colossoma bidens (Characidae).
A. spinatus KRITSKY, BOEGER & VAN EVERY, 1992 from Myleus rubripinnis
(Serrasalmidae) (Fig. 3-17 P).
A. spiralocirrus KRITSKY, THATCHER & KAYTON, 1979 from Brycon melanopterus
(Characidae).
A. stachophallus KRITSKY, BOEGER & VAN EVERY, 1992 from Pygocentrus nattereri
(Serrasalmidae).
A. stagmophallus KRITSKY, BOEGER & VAN EVERY, 1992 from Myleus rubripinnis
(Serrasalmidae).
A. strongylophallus KRITSKY, BOEGER & VAN EVERY, 1992 from Triportheus
elongatus (Characidae).
A. thatcheri BOEGER & KRITSKY, 1988 from Pygocentrus nattereri (Serrasalmidae).
A. tricornis KRITSKY, BOEGER & VAN EVERY, 1992 from Triportheus angulatus,
A. xaniophallus KRITSKY, BOEGER & VAN EVERY, 1992 from Pristobrycon sp. n.,
Pristobrycon eigenmanni (Serrasalmidae) (Fig. 3-17 O).
Ancistrohaptor AGARWAL & KRITSKY, 1998
Body fusiform. Tegument smooth. Eyes four; eyes granules absent or
ping. Copulatory organ composed of MCO, accessory piece. MCO
sclerotized, coiled. Accessory piece two, one non-articulated, one articulat-
ed to MCO. Seminal vesicle fusiform with thin walls. Prostatic reservoirs
unknown. Germarium ovate or elongate, solid. Vagina dextro-marginal or
ventro-dextral. Hooks dactylogyrid, shank divided in two portions clearly
defined, 14, with ancyrocephaline distribution (4 dorsal, 10 ventral).
Ventral, dorsal anchor pairs present. Ventral anchor with point, elongate
shaft, roots well defined; anchor filament delicate, often inconspicuous.
Dorsal anchor with shaft, point, base composed of two roots. Ventral bar
rod-shaped with anterior transversal groove, anterior, posterior projections
absent. Dorsal bar slightly V-shaped or rod-shaped; anterior, posterior
projections absent. Parasites of the gills of Triportheus spp. (Characidae,
Characiformes).
A. falcatum AGARWAL & KRITSKY, 1998 from Triportheus elongatus.
A. falciferum AGARWAL & KRITSKY, 1998 from Triportheus elongatus, Triportheus
angulatus, Triportheus albus Triportheus sp. (Type species) (Figs. 3-18 A-C).
A. falcunculum AGARWAL & KRITSKY, 1998 from Triportheus albus, Triportheus
elongatus, Triportheus angulatus.
Annulotrematoides KRITSKY & BOEGER, 1995
Body fusiform. Tegument smooth, annulations present. Eyes four; eyes
granules absent or present. Pharynx bulbous, one subunit. Ceca confluent.
Gonads overlapping. Copulatory organ composed of MCO, accessory
piece. MCO sclerotized, J-shaped. Accessory piece articulated to MCO by
copulatory ligament. Seminal vesicle fusiform. Prostatic reservoirs round
to short, ovate. Germarium ovate or elongate. Vagina sinistro-marginal,
sclerotized. Hooks dactylogyrid, shank divided in two portions clearly
defined; 14, with ancyrocephaline distribution (4 dorsal, 10 ventral).
Ventral, dorsal anchor pairs present. Ventral anchor with point, shaft, roots
well defined; anchor filament delicate, often inconspicuous. Dorsal anchor
with shaft, point, base composed of two roots. Ventral bar rod-shaped;
antero-median projection; posterior median projection absent. Dorsal bar
slightly V-shaped; anterior, posterior projections absent. Parasites of the
gills of species of Characiformes.
A. amazonicus KRITSKY & BOEGER, 1995 from Psectrogaster rutiloides (Curima-
tidae) (Type species) (Figs. 3-19 A-B).
A. bryconi CUGLIANNA, CORDEIRO & LUQUE, 2003 from Brycon cephalus
(Characidae).
Aphanoblastella KRITSKY, MENDONZA-FRANCO & SCHOLZ, 2000
Synonyms: Urocleidoides (in part); Cleidodiscus (in part)
present. Pharynx bulbous, one subunit. Ceca confluent. Gonads tandem;
germarium anterior to testis. Copulatory organ composed of MCO,
accessory piece, sinuous or coiled. Accessory piece one, non-articulated.
Seminal vesicle fusiform. Prostatic reservoirs round to short, ovate.
Germarium ovate or elongate. Vagina muscular, sinistro-marginal. Hooks
dactylogyrid, shank simple; 14, with ancyrocephaline distribution (4 dorsal,
10 ventral). Ventral, dorsal anchor pairs present. Ventral anchor with point,
shaft, roots well defined; anchor filament delicate, often inconspicuous.
slightly V-shaped, anterior projection absent; posterior projections present.
Dorsal bar slightly V-shaped; anterior, posterior projections absent. Para-
sites of the gills of species of Pimelodidae (Siluriformes).
A. mastigatus SURIANO, 1986 from Rhamdia sapo.

A. robustus (MIZELLE & KRITSKY, 1969) from Rhamdia sp.
A. travassosi (PRICE, 1938) from Rhamdia rogersi, Rhamdia sebae, Pimelodella
laticeps, Rhamdia quelen (Type species) (Figs. 3-20 A-D).
Cacatuocotyle BOEGER, DOMINGUES & KRITSKY, 1997
Body robust, strongly flattened dorso-ventrally. Tegument smooth. Eyes
four; eyes granules present. Pharynx bulbous, one subunit. Ceca confluent.
Gonads tandem; germarium anterior to testis. Copulatory organ com-
posed of MCO, accessory piece. MCO coiled. Accessory piece articulated
to MCO by copulatory ligament. Seminal vesicle C-shaped or fusiform.
Prostatic reservoirs round to short, ovate. Germarium ovate or elongate.
Vagina sinistro-marginal, sclerotized. Anterior margins of haptor heavily
muscular. Hooks dactylogyrid with shank simple; 14, 12 ventral, on
posterior margin of haptor, 2 central. Ventral pair of anchors with point,
shaft, roots well defined. Dorsal anchor pair absent. Ventral bar rod-
shaped; anterior, posterior projections absent. Dorsal bar absent. Parasites
of gills of species of Characidae (Characiformes).
Type and only species: C. paranaensis BOEGER, DOMINGUES & KRITSKY, 1997
from Characidium pterostictum, Characidium lanei (Figs. 3-21 A-B).
Calpidothecioides KRITSKY, BOEGER & JÉGU, 1997
Synonyms: Urocleidus (in part).
Body fusiform. Tegument smooth. Eyes two; eye granules absent, present.
Pharynx bulbous, one subunit. Ceca confluent. Gonads overlapping.
Copulatory organ composed of MCO, accessory piece. MCO J-shaped,
somewhat straight, or slightly curved. Accessory piece articulated directly
to MCO or articulated to MCO by copulatory ligament. Seminal vesicle C-
shaped or fusiform. Germarium ovate or elongate. Vagina double, muscu-
lar, openings one marginal, one dorsal. Hooks dactylogyrid with shank
divided in two portions clearly defined; 14, with ancyrocephaline distribu-
tion (4 dorsal, 10 ventral). Ventral, dorsal anchor pairs present. Ventral
anchor with point, shaft, roots well defined. Dorsal anchor with shaft,
point, base composed of two roots. Ventral bar rod-shaped, with slightly
projected margin, with or without medial cleft; posterior projection absent.
Dorsal bar slightly V-shaped; anterior, posterior projections absent. Para-
sites of the gills of species of Serrasalmidae (Characiformes).
C. orthus (MIZELLE & PRICE, 1965) KRITSKY, BOEGER & KRITSKY, 1997 from
Pygocentrus nattereri.
C. pygopristi KRITSKY, BOEGER & JÉGU, 1997 from Pygopristis denticulata (Type
species) (Figs. 3-22 A-B).
Calpidothecium KRITSKY, BOEGER & JÉGU, 1997
Synonyms: Urocleidus (in part); Cleidodiscus (in part).
Body fusiform. Tegument smooth; annulations on body absent or present.
Eyes two; eye granules absent. Pharynx bulbous, one subunit. Ceca
confluent. Gonads overlapping. Copulatory organ composed of MCO,
accessory piece. MCO straight or slightly curved. Accessory piece articu-

lated to MCO by copulatory ligament. Seminal vesicle fusiform. Prostatic
reservoirs round to short, ovate. Germarium ovate or elongate. Vagina
sinistro-marginal, sclerotized. Hooks dactylogyrid with shank divided in
two portions clearly defined; 14, with ancyrocephaline distribution (4
dorsal, 10 ventral). Ventral, dorsal anchor pairs present. Ventral anchor
with point, shaft, roots well defined. Dorsal anchor with shaft, point, base
composed of two roots. Ventral bar slightly V-shaped or rod-shaped;
anterior, posterior projections absent. Dorsal bar V-shaped; anterior,
posterior projections absent. Parasites of the gills of species of Serrasalm-
idae (Characiformes).
C. crescentis (MIZELLE & PRICE, 1965) KRITSKY, BOEGER & JÉGU, 1997 from
Pygocentrus nattereri, Pygopristis denticulata (Type species) (Figs. 3-23 A-C).
C. serrasalmus (MIZELLE & PRICE, 1965) KRITSKY, BOEGER & JÉGU, 1997 from
Pygopristis denticulata, Pygocentrus nattereri.
Cosmetocleithrum KRITSKY, THATCHER & BOEGER, 1986
Body fusiform. Tegument smooth. Eyes absent; eyes granules present.
Pharynx bulbous, one subunit. Ceca confluent. Gonads tandem; germari-
um anterior to testis. Copulatory organ composed of MCO, accessory
piece. MCO J-shaped or coiled. Accessory piece one, non-articulated.
Germarium ovate or elongate. Vagina sinistro-marginal or sinistro-ventral,
sclerotized. Hooks dactylogyrid with shank simple; 14, with ancyrocepha-
line distribution (4 dorsal, 10 ventral). Ventral, dorsal anchor pairs present.
Ventral anchor with point, shaft, roots well defined. Dorsal anchor with
shaft, point, base composed of two roots. Ventral bar slightly V-shaped,
slightly U-shaped, or strongly V-shaped, anterior projection absent, poste-
rior projections present or absent. Dorsal bar slightly V-shaped, slightly U-
shaped, strongly V-shaped, or rod-shaped; anterior projection absent,
posterior projections on dorsal bar two, ribbon-like. Parasites of species of
Siluriformes, especially species of Doradidae.
C. bulbocirrus KRITSKY, THATCHER & BOEGER, 1986 from Pterodoras granulosus
(Doradidae).
C. confusus KRITSKY, THATCHER & BOEGER, 1986 from Oxydoras niger (Doradidae).
C. gussevi KRITSKY, THATCHER & BOEGER, 1986 from Oxydoras niger (Doradidae)
(Figs. 3-24 A-F).
C. longivaginatum SURIANO & INCORVAIA, 1995 from Pimelodus albicans (Pimelo-
didae).
C. parvum KRITSKY, THATCHER & BOEGER, 1986 from Oxydoras niger (Dora-
didae).
C. rarum KRITSKY, THATCHER & BOEGER, 1986 from Oxydoras niger (Dora-
didae).
C. sobrinus KRITSKY, THATCHER & BOEGER, 1986 from Oxydoras niger (Dora-
didae).
Curvianchoratus HANEK, MOLNAR & FERNANDO, 1974

Synonym: Notodiplocerus SURIANO, 1980
Body short, robust, strongly flattened. Tegument smooth. Eyes four; eye
posed of MCO, accessory piece. MCO J-shaped or coiled. Accessory piece
articulated directly to MCO. Seminal vesicle fusiform. Prostatic reservoirs
unknown. Germarium ovate or elongate. Vagina dextro-marginal, sclero-
tized. Hooks dactylogyrid with shank simple; 14, with ancyrocephaline
distribution (4 dorsal, 10 ventral). Ventral, dorsal anchor pairs. Ventral
anchor with point, shaft, roots well defined. Dorsal anchor greatly
modified, base apparently elongate, distorted, distal extremity sometimes
hook-like. Ventral bar strongly V-shaped; anterior, posterior projections
absent. Dorsal bar double, strongly distorted. Parasites of the gills of
species of Curimatidae (Characiformes).
C. hexacleidus HANEK, MOLNAR & FERNANDO, 1974 from Curimata argentea.
C. singularis (SURIANO,1980) from Pseudocurimata gilberti (Figs. 3-25 A-C).
Dawestrema PRICE & NOWLIN, 1967
germarium anterior to testis. Copulatory organ composed of MCO and
accessory piece. MCO coiled. Accessory piece articulated directly to MCO.
Seminal vesicle two, fusiform. Prostatic reservoirs round to short, ovate.
Germarium ovate or elongate. Vagina sinistro-marginal, sclerotized.
Hooks dactylogyrid, with shank divided in two portions clearly defined;
14, arranged concentrically around haptor dorso-ventrally. Ventral, dorsal
anchor pairs. Ventral anchor with point, shaft, roots well defined. Dorsal
anchor dactylogyrid, with shaft, point, base composed of two roots.
Ventral bar present rod-shaped, antero-median projection present, posteri-
or projection absent. Dorsal bar slightly U-shaped; anterior, posterior
projections absent. Parasites of the gills of species of Arapaima gigas,
Osteoglossidae (Osteoglossiformes).
D. cycloancistrioides KRITSKY, BOEGER & THATCHER, 1985 from Arapaima gigas.
D. cycloancistrium PRICE & NOWLIN, 1967 from Arapaima gigas (Type species)
(Figs. 3-26 A-C).
D. punctatum PRICE & NOWLIN, 1967 from Arapaima gigas.
Demidospermus SURIANO, 1983
Synonyms: Omothecium KRITSKY, THATCHER & BOEGER, 1987; Paramphoclei-
thrum SURIANO & INCORVAIA, 1995.
Body fusiform. Tegument smooth. Eyes four or two; eye granules absent
or present. Pharynx bulbous, one subunit. Ceca confluent. Gonads
tandem; germarium anterior to testis. Copulatory organ composed of
MCO, accessory piece. MCO J-shaped, somewhat straight, slightly curved,
coiled. Accessory piece one, non-articulated or articulated directly to
MCO. Seminal vesicle fusiform. Prostatic reservoirs round to short, ovate.

Germarium ovate or elongate. Vagina sinistro-marginal, sclerotized or
muscular. Hooks dactylogyrid, entire shank inflated, conferring a robust
morphology to hook, or proximal portion of shank inflated (circular,
ovate), or shank divided in two portions clearly defined, or shank simple;
14, with ancyrocephaline distribution (4 dorsal, 10 ventral). Ventral, dorsal
anchor pairs present. Ventral anchor with point, shaft, roots not defined or
well defined. Dorsal anchor with shaft, point, base not clearly separated in
roots or base composed of two roots. Ventral bar strongly V-shaped;
anterior, posterior projection absent. Dorsal bar strongly V-shaped; anteri-
or projection absent; posterior projection present. Parasites of the gills of
species of Siluriformes, especially of Pimelodidae.
D. anus SURIANO, 1983 from Loricaria anus (Loricariidae). (Type species).
D. armostus KRITSKY & GUTIÉRREZ, 1998 from Pimelodus albicans, Pimelodus
clarias (Pimelodidae).
D. bidiverticulatum (SURIANO & INCORAVIA, 1995) from Pimelodus clarias, Pimelodus
albicans, Pimelodus clarias maculatus (Pimelodidae).
D. cornicinus KRITSKY & GUTIÉRREZ, 1998 from Iheringichthys westermanni
(Pimelodidae) (Figs. 3-27 A-F).
D. idolus KRITSKY & GUTIÉRREZ, 1998 from Pimelodus albicans (Pimelodidae).
D. labrosi FRANÇA, ISSAC, PAVANELLI & TAKEMOTO, 2003 from Iheringinchthys
labrosus (Pimelodidae).
D. leptosynophallus KRITSKY & GUTIÉRREZ, 1998 from Iheringichthys westermanni
(Pimelodidae).
D. luckyi (KRITSKY, THATCHER & BOEGER, 1987) from Pinirampus pirinampu
(Pimelodidae).
D. majusculus KRITSKY & GUTIÉRREZ, 1998 from Pimelodus albicans (Pimelodidae).
D. mandi FRANÇA, ISSAC, PAVANELLI & TAKEMOTO, 2003 from Iheringinchthys
labrosus (Pimelodidae).
D. paravalenciennesi GUTIERREZ & SURIANO, 1992 from Pimelodus clarias (Pimelo-
didae).
D. pinirampi (KRITSKY, THATCHER & BOEGER, 1987) from Pirinampus pirinampu
(Pimelodidae).
D. uncusvalidus GUTIERREZ & SURIANO, 1992 from Pimelodus clarias, Paraucheni-
pterus galeatus (Auchenipteridae).
D. valenciennesi GUTIERREZ & SURIANO, 1992 from Parapimelodus valenciennesi
(Pimelodidae).
Diaphorocleidus JOGUNOORI, KRITSKY & VENKATANARASAIAH, 2004
Synonym: Urocleidoides sensu lato (in part)
Body fusiform. Tegument smooth. Eyes four; eyes granules absent, or
present. Ceca confluent. Gonads overlapping. Copulatory organ com-
posed of MCO and accessory piece. MCO with loose coils, not inflated
proximally. Accessory piece one, non-articulated. Seminal vesicle fusiform.
Prostatic reservoirs round to short, ovate. Germarium ovate or elongate;
solid. Vagina sclerotized, single, sinistro-ventral. Vaginal bar, accessory

vaginal sclerite absent. Hooks dactylogyrid; shank divided in two portions
clearly defined; 14, with ancyrocephaline distribution (4 dorsal, 10 ventral).
well defined. Dorsal anchor with shaft, point, base composed of two roots.
Ventral bar slightly V-shaped, with antero-median projection. Dorsal bar
slightly V-shaped; anterior, posterior projections absent. Parasites of the
gills of species of Characidae (Characiformes).
D. affinis (MIZELLE, KRITSKY & CRANE, 1968) from Creatochanes affinis
D. armillatus JOGUNOORI, KRITSKY & VENKATANARASAIAH, 2004 from Gymnoc-
orymbus ternetzi (Type species) (Figs. 3-28 A-C).
D. kabatai (MOLNAR, HANEK & FERNANDO, 1974) from Astyanax fasciatus.
D. microstomus (MIZELLE, KRITSKY & CRANE, 1968) from Hemigrammus microsto-
mus.
Enallothecium KRITSKY, BOEGER & JÉGU, 1998
Synonym: Notothecium (in part).
Body short, robust, or fusiform. Tegument scaled or smooth. Eyes four or
two; eyes granules absent or present. Ceca confluent. Gonads overlapping.
Copulatory organ composed of MCO, accessory piece. MCO straight or
slightly curved. Accessory piece articulated to MCO by copulatory
ligament. Seminal vesicle C-shaped. Prostatic reservoirs round to short,
ovate. Germarium ovate or elongate. Vagina sinistro-dorsal, muscular.
Hooks dactylogyrid, shank divided in two portions clearly defined; 14,
with ancyrocephaline distribution (4 dorsal, 10 ventral). Ventral, dorsal
anchor pairs present. Ventral anchor with point, shaft, roots well defined.
slightly U-shaped; anterior, posterior projections absent. Dorsal bar
slightly U-shaped; anterior, posterior projections absent. Parasites of the
gills of species of Serrasalmidae (Characiformes).
E. aegidatum (BOEGER & KRITSKY, 1988) from Serrasalmus spilopleura, Serrasalmus
sp., Serrasalmus rhombeus, Serrasalmus compressus, Serrasalmus gouldingi, Serra-
salmus elongatus, Pristobrycon sp., Pygocentrus nattereri, Serrasalmus sp. (Type
species) (Fig. 3-29).
E. cornutum KRITSKY, BOEGER & JÉGU, 1998 from Serrasalmus gouldingi, Pristobry-
con eigenmanni, Serrasalmus sp., Serrasalmus sp., Pristobrycon sp., Serrasalmus
rhombeus, Serrasalmus compressus.
E. umbelliferum KRITSKY, BOEGER & JÉGU, 1998 from Serrasalmus compressus,
Serrasalmus rhombeus, Serrasalmus sp.
E. variabilum KRITSKY, BOEGER & JÉGU, 1998 from Pristobrycon striolatus.
Euryhaliotrema BOEGER & KRITSKY, 2002
Body fusiform. Tegument smooth. Eyes four; eye granules absent or
germarium anterior to testis. Copulatory organ composed of MCO, accesso-
ry piece. MCO sinuous, or somewhat straight, or slightly curved, or coiled,
with inflated proximal portion. Seminal vesicle fusiform. Prostatic reservoirs

round to short, ovate. Germarium ovate or elongate. Vagina dextro-
marginal, sclerotized. Hooks dactylogyrid, shank simple; 14, with ancyro-
cephaline distribution (4 dorsal, 10 ventral). Ventral anchor pair present;
dorsal anchor pairs present or absent. Ventral anchor with point, shaft, roots
Ventral bar slightly U-shaped, or rod-shaped, with short anterior sclerotized
muscle-attachment; posterior projection absent or shield like. Dorsal bar
present or absent, slightly U-shaped or rod-shaped; anterior projection
absent; posterior projection absent or shield like. Parasites of freshwater and
marine species of species of Sciaenidae (Perciformes).
E. chaoi KRITSKY & BOEGER, 2002 from Plagioscion squamosissimus (Type species)
(Figs. 3-30 A-B).
E. dontykoleos FEHLAUER & BOEGER, 2005 from Pachyurus junki
E. lovejoyi KRITSKY & BOEGER, 2002 from Plagioscion squamosissimus.
E. monacanthus KRITSKY & BOEGER, 2002 from Plagioscion squamosissimus.
E. potamocetes KRITSKY & BOEGER, 2002 from Plagioscion squamosissimus.
E. succedaneus KRITSKY & BOEGER, 2002 from Plagioscion squamosissimus.
E. thatcheri KRITSKY & BOEGER, 2002 from Plagioscion squamosissimus.
Gonocleithrum KRITSKY & THATCHER, 1983
Body fusiform. Tegument smooth. Eyes four; eye granules absent, or
ping, or tandem; germarium anterior to testis. Copulatory organ composed
of MCO, accessory piece. Accessory piece articulated directly to MCO.
Germarium ovate or elongate. Vagina sinistro-marginal, sclerotized or
muscular; gonadal bar present. Hooks dactylogyrid, shank divided in two
portions clearly defined; 14, with ancyrocephaline distribution (4 dorsal,
shaft, roots well defined. Dorsal anchor with shaft, point, base composed
of two roots. Ventral bar slightly U-shaped; anterior projection present or
absent, posterior projection absent. Dorsal bar slightly U-shaped, or
strongly V-shaped, or rod-shaped; anterior, posterior projections absent.
Parasites of the gills of species of Osteoglossum (Osteoglossidae, Osteoglo-
ssiformes) – presently only known from Osteoglossum biccirhosum.
G. aruanae KRITSKY & THATCHER, 1983.
G. coenoideum KRITSKY & THATCHER, 1983.
G. cursitans KRITSKY & THATCHER, 1983.
G. planacroideum KRITSKY & THATCHER, 1983.
G. planacrus KRITSKY & THATCHER, 1983 (Type species) (Figs. 3-31 A-D).
Gussevia KOHN & PAPERNA, 1964
Synonym: Longihaptor MIZELLE & KRITSKY, 1969
Body robust, strongly flattened, or fusiform. Tegument smooth. Eyes four;
eye granules absent or present. Pharynx bulbous, one subunit. Ceca
confluent. Gonads overlapping. Copulatory organ composed of MCO and

accessory piece. MCO coiled. Accessory piece one, non-articulated, or
articulated directly to MCO. Seminal vesicle fusiform. Prostatic reservoirs
round to short, ovate. Germarium ovate or elongate. Vagina ventral, or
dextro-marginal, or sinistro-marginal, sclerotized or muscular. Hooks dacty-
logyrid, proximal portion of shank inflated (circular, ovate), or shank simple;
14, with ancyrocephaline distribution (4 dorsal, 10 ventral); hook pair 5
morphologically distinct from other pairs. Ventral, dorsal anchor pairs
present. Ventral anchor with point, shaft, roots defined or not well defined.
slightly U-shaped, or rod-shaped; anterior margin smooth or with slightly
projected, with or without medial cleft, or with anterior transversal groove;
posterior projection absent. Dorsal bar rod-shaped; anterior, posterior
projection absent. Parasites of Cichlidae (Perciformes).
G. alii (MOLNAR, HANEK & FERNANDO, 1974) from Cichlasoma bimaculatum.
G. alioides KRITSKY, THATCHER & BOEGER, 1986 from Cichlasoma severum.
G. arilla KRITSKY, THATCHER & BOEGER, 1986 from Cichla ocellaris.
G. asota KRITSKY, THATCHER & BOEGER, 1989 from Astronotus ocellatus (Figs. 3-
32 A-D).
G. astronoti KRITSKY, THATCHER & BOEGER, 1989 from Astronotus ocellatus.
G. cichlasomatis (MOLNAR, HANEK & FERNANDO, 1974) from Cichlasoma bimacu-
latum.
G. dispar KRITSKY, THATCHER & BOEGER, 1986 from Cichlasoma severum.
G. disparoides KRITSKY, THATCHER & BOEGER, 1986 from Cichlasoma severum.
G. dobosi (MOLNAR, HANEK & FERNANDO, 1974) from Cichlasoma bimaculatum.
G. elephus KRITSKY, THATCHER & BOEGER, 1986 from Uaru amphiacanthoides.
G. herotilapiae VIDAL-MARTÍNEZ, SCHOLZ & AGUIRRE-MACEDO, 2001 from Hero-
tilapia multispinosa.
G. longihaptor (MIZELLE & KRITSKY, 1969) from Cichla ocellaris.
G. obtusa KRITSKY, THATCHER & BOEGER, 1986 from Uaru amphiacanthoides.
G. rogersi KRITSKY, THATCHER & BOEGER, 1989 from Astronotus ocellatus.
G. spiralocirra KOHN & PAPERNA, 1964 from Pterophyllum scalare (Type species).
G. tucunarense KRITSKY, THATCHER & BOEGER, 1986 from Cichla ocellaris.
G. undulata KRITSKY, THATCHER & BOEGER, 1986 from Cichla ocellaris.
Heterothecium KRITSKY, BOEGER & JÉGU, 1997
Body fusiform. Tegument smooth; annulations on body present or absent.
Eyes four; eyes granules absent or present. Pharynx bulbous, one subunit.
Ceca confluent. Gonads overlapping. Copulatory organ composed of
MCO, accessory piece. MCO straight or slightly curved. Accessory piece
articulated to MCO by copulatory ligament. Seminal vesicle fusiform.
Vagina sinistro-dorsal, muscular. Hooks dactylogyrid, shank divided in two
of two roots. Ventral bar rod-shaped; anterior, posterior projections
absent. Dorsal bar slightly U-shaped; anterior, posterior projections
absent. Parasites of species of Serrasalmidae (Characiformes).
H. dicrophallum KRITSKY, BOEGER & JÉGU, 1997 from Catoprion mento.
H. globatum KRITSKY, BOEGER & JÉGU, 1997 from Serrasalmus gouldingi (Type
species) (Fig. 3-33).
Heterotylus JOGUNOORI, KRITSKY & VENKATANARASAIAH, 2004
Body fusiform. Tegument smooth. Eyes absent; eye granules present or
straight or slightly curved. Accessory piece one, non-articulated. Seminal
vesicle fusiform. Prostatic reservoirs round to short, ovate. Germarium
ovate or elongate. Vagina ventral, sclerotized. Hooks dactylogyrid, shank
simple; 14, with ancyrocephaline distribution (4 dorsal, 10 ventral).
not defined. Dorsal anchor, with shaft, point, base composed of two roots.
Ventral bar inverted U-shaped, with tapering extremities; anterior, posteri-
or projections absent. Dorsal bar slightly U-shaped; anterior, posterior
projections absent. Parasites of species of Loricariidae (Siluriformes).
Type and only species: H. heterotylus JOGUNOORI, KRITSKY & VENKATANARA-
SAIAH, 2004 from Hypostomus sp. (Figs. 3-34 A-D).
Jainus MIZELLE, KRITSKY & CRANE, 1968
Body short, robust, or fusiform. Tegument smooth. Eyes four; eye
Gonads overlapping. Copulatory organ composed of MCO, accessory
piece. MCO J-shaped, or sinuous, or coiled. Accessory piece one, non-
articulated. Seminal vesicle fusiform. Prostatic reservoirs round to short,
ovate. Germarium ovate or elongate. Vagina ventral, or sinistro-ventral,
sclerotized. Hooks dactylogyrid; 14, with ancyrocephaline distribution (4
dorsal, 10 ventral); hook pair 5 similar to other pairs, or significantly
distinct from others, reduced. Ventral, dorsal anchor pairs present. Ventral
anchor robust, with elongate superficial, deep roots. Dorsal anchor with
shaft, point, base composed of two roots. Ventral bar rod-shaped; antero-
median projection present or absent; posterior median projection present.
Dorsal bar slightly U-shaped, or strongly V-shaped, or rod-shaped;
anterior, posterior projections absent. Parasites of gills of species of
Characidae (Characiformes).
J. amazonensis KRITSKY, THATCHER & KAYTON, 1980 from Brycon melanopterus.
J. hexops KRITSKY & LEIBY, 1972 from Astyanax fasciatus.
J. jainus MIZELLE, KRITSKY & CRANE, 1968 from Chalceus macrolepidotus (Type
species) (Figs. 3-35 A-D).
J. robustus MIZELLE, KRITSKY & CRANE, 1968 from Creatochanes affinis.
Kritskyia KOHN, 1990

Body fusiform. Tegument smooth. Eyes four; eye granules present or
Accessory piece more than two, non-articulated, or one, non-articulated to
MCO. Seminal vesicle fusiform. Prostatic reservoirs round to short, ovate.
Germarium ovate or elongate. Vagina sinistro-marginal or sinistro-ventral.
Hooks dactylogyrid with shank divided in two portions clearly defined; 14
on posterior margin of haptor. Ventral, dorsal anchors and bars absent.
Parasites of the excretory system of species of Characiformes and
Siluriformes.
K. annakohnnae BOEGER, TANAKA & PAVANELLI, 2001 from Serrasalmus spilopleu-
ra, Serrasalmus marginatus (Serrasalmidae, Characiformes).
K. boegeri TAKEMOTO, LIZAMA & PAVANELLI, 2002 from Prochilodus lineatus
(Prochilodontidae, Characiformes).
K. eirasi GUIDELLI, TAKEMOTO & PAVANELLI, 2003 from Leporinus lacustris
(Anostomidae, Characiformes).
K. moraveci KOHN, 1990 from Rhamdia quelen (Pimelodidae, Siluriformes)
(Type species) (Figs. 3-36 A-C).
Linguadactyloides THATCHER & KRITSKY, 1983
present. Pharynx two bulbous subunits. Ceca confluent. Gonads tandem;
testes anterior to germarium. Testis multiple, pre-germarian.. Copulatory
organ composed of MCO, accessory piece. MCO coiled. Accessory piece
Prostatic reservoirs very long, often looping posteriorly. Germarium ovate
or elongate. Vagina dextro-marginal, sclerotized. Hooks dactylogyrid with
entire shank inflated, conferring a robust morphology to hook, or shank
Ventral, dorsal anchor pairs present. Ventral anchor short robust, deep
root longer than superficial root. Dorsal anchor robust, deep root longer
than superficial root. Ventral bar rod-shaped; anterior projection absent;
posterior projection present. Dorsal bar absent. Parasites of the gills of
species of Serrasalmidae (Characiformes).
Type and only species: L. brinkmanni THATCHER & KRITSKY, 1983 from
Colossoma macropomum (Figs. 3-37 A-D)
Monocleithrium PRICE & MCMAHON, 1966
Body fusiform. Tegument smooth. Eyes four; eye granules absent or present.
Pharynx bulbous, one subunit. Ceca confluent. Gonads tandem; germarium
anterior to testis. Copulatory organ composed of MCO, accessory piece.
MCO coiled. Accessory piece articulated directly to MCO. Seminal vesicle
unknown. Prostatic reservoirs unknown. Germarium ovate or elongate.
Vagina ventral. Hooks dactylogyrid, with shank divided in two portions clearly
defined; 14, with ancyrocephaline distribution (4 dorsal, 10 ventral). Ventral,
dorsal anchor pairs present. Ventral anchor with point, shaft, roots well
defined. Dorsal anchor with shaft, point, base composed of two roots. Ventral
bar rod-shaped; anterior, posterior projections absent. Dorsal bar absent.
Parasites of the gill of species of Hemiodontidae (Characiformes).
Type and only species: M. lavergneae PRICE & MCMAHON, 1966 from
Hemiodus semitaeniatus (Fig. 3-38).
Mymarothecium KRITSKY, BOEGER & JÉGU, 1996
straight or slightly curved. Accessory piece articulated directly to MCO, or
Prostatic reservoirs short, round to ovate. Germarium ovate or elongate.
Vagina dextro-dorsal, muscular. Hooks dactylogyrid, shank divided in two
of two roots. Ventral bar rod-shaped; anterior, posterior projections
present or absent. Dorsal bar slightly U-shaped or rod-shaped; anterior
projection present or absent, posterior projection absent. Parasites of gills
of species of Serrasalmidae (Characiformes).
M.boegeri COHEN & KOHN, 2005 from Colossoma macropomum (Characiformes,
Characidae).
M.dactylotum KRITSKY, BOEGER & JÉGU, 1996 from Serrasalmus sp. (2n = 58),
Serrasalmus sp. (2 of Jégu), Pristobrycon sp., Serrasalmus rhombeus (Type
species) (Figs. 3-39 A-C).
M.galeolum KRITSKY, BOEGER & JÉGU, 1996 from Pristobrycon eigenmanni,
Serrasalmus gouldingi, Pygocentrus nattereri, Pristobrycon sp., Serrasalmus rhombeus.
M.perplanum KRITSKY, BOEGER & JÉGU, 1996 from Serrasalmus spilopleura.
M.viatorum BOEGER, PIASECKI & SOBECKA, 2002 from Piaractus brachypomus.
M.whittingtoni KRITSKY, BOEGER & JÉGU, 1996 from Serrasalmus rhombeus, from
Serrasalmus sp. (2n = 58).
Notothecium BOEGER & KRITSKY, 1988
Body fusiform. Tegument smooth. Eyes absent; eye granules absent or
ping. Copulatory organ composed of MCO, accessory piece. MCO single, or
double, sclerotized, J-shaped, or straight or slightly curved. Accessory piece
articulated to MCO by copulatory ligament. Seminal vesicle C-shaped.
Vagina dextro-dorsal, muscular. Hooks dactylogyrid, shank divided in two
portions clearly defined; 14, with ancyrocephaline distribution (4 dorsal, 10
ventral). Ventral, dorsal anchor pairs present. Ventral anchor with point,
shaft, roots well defined. Dorsal anchor with shaft, point, base composed of
two roots. Ventral bar slightly U-shaped or rod-shaped; anterior, posterior
projections absent. Dorsal bar strongly V-shaped; anterior, posterior projec-

tions absent. Parasites of gills of species of Serrasalmidae (Characiformes).
N. circellum KRITSKY, BOEGER & JÉGU, 1998 from Serrasalmus gouldingi, Pristobry-
con sp.
N. cyphophallum KRITSKY, BOEGER & JÉGU, 1998 from Serrasalmus rhombeus,
Pristobrycon eigenmanni, Serrasalmus compressus, Serrasalmus elongatus, Serrasalmus
gouldingi, Serrasalmus sp.
N. deleastoideum KRITSKY, BOEGER & JÉGU, 1998 from Serrasalmus sp.
N. deleastum KRITSKY, BOEGER & JÉGU, 1998 from Serrasalmus elongatus, Serrasal-
mus rhombeus, Serrasalmus gouldingi, Serrasalmus sp.
N. mizellei BOEGER & KRITSKY, 1988 from Pygocentrus nattereri (Type species)
(Figs. 3-40 A-B).
N. modestum KRITSKY, BOEGER & JÉGU, 1998 from Serrasalmus spilopleura.
N. phyleticum KRITSKY, BOEGER & JÉGU, 1998 from Serrasalmus rhombeus.
N. reduvium KRITSKY, BOEGER & JÉGU, 1998 from Serrasalmus sp.
Notothecioides KRITSKY, BOEGER & JÉGU, 1997
Pharynx bulbous, one subunit. Ceca confluent. Gonads overlapping. Copula-
tory organ composed of MCO and accessory piece. MCO straight or slightly
curved. Accessory piece articulated to MCO by copulatory ligament. Seminal
vesicle fusiform. Prostatic reservoirs round to short, ovate. Germarium ovate
or elongate. Vagina middorsal, looping left caecum. Hooks dactylogyrid, shank
divided in two portions clearly defined; 14, with ancyrocephaline distribution
(4 dorsal, 10 ventral). Ventral, dorsal anchor pairs present. Ventral anchor with
point, shaft, roots well defined. Dorsal anchor with shaft, point, base
composed of two roots. Ventral bar strongly V-shaped; anterior, posterior
projections absent. Dorsal bar rod-shaped; anterior, posterior projections
absent. Parasites of the gills of species of Serrasalmidae (Characiformes).
Type and only species: N. llewellyni KRITSKY, BOEGER & JÉGU, 1997 from
Myleus torquatus, M. rubripinnis (Fig. 3-41).
Notozothecium BOEGER & KRITSKY, 1988
Pharynx bulbous, one subunit. Ceca confluent. Gonads overlapping. Copula-
tory organ composed of MCO and accessory piece. MCO coiled (usually less
than one ring) or J-shaped. Accessory piece articulated to MCO by copulatory
ligament. Seminal vesicle fusiform. Prostatic reservoirs round to short, ovate.
Germarium ovate or elongate. Vagina dorso-dextral, sclerotized, looping right
caecum. Hooks dactylogyrid, shank divided in two portions clearly defined;
14, with ancyrocephaline distribution (4 dorsal, 10 ventral). Ventral, dorsal
anchor pairs present. Ventral anchor with point, shaft, roots well defined.
Dorsal anchor with shaft, point, base composed of two roots. Ventral bar rod-
shaped; anterior projection present; posterior projection absent. Dorsal bar
rod-shaped or slightly V-shaped; anterior, posterior projections absent. Para-
sites of the gills of species of Serrasalmidae (Characiformes).
N. bethae KRITSKY, BOEGER & JÉGU, 1996 from Mylesinus paucisquamatus,

Mylesinus paraschomburgkii, Myleus pacu, Myleus rhomboidalis.
N. euzeti KRITSKY, BOEGER & JÉGU, 1996 from Acnodon normani.
N. foliolum KRITSKY, BOEGER & JÉGU, 1996 from Pristobrycon sp.
N. janauachensis BELMONT-JÉGU, DOMINGUES & LATERÇA, 2004 from Colossoma
macropomum.
N. minor BOEGER & KRITSKY, 1988 from Pygocentrus nattereri.
N. penetrarum BOEGER & KRITSKY, 1988 from Pygocentrus nattereri (Type species)
(Figs. 3-42 A-D).
N. robustum KRITSKY, BOEGER & JÉGU, 1996 from Pristobrycon striolatus.
N. teinodendrum KRITSKY, BOEGER & JÉGU, 1996 from Serrasalmus elongatus,
Serrasalmus rhombeus, Serrasalmus gouldingi, Pristobrycon eigenmanni, Serrasalmus
sp., Serrasalmus manuelli, Pristobrycon sp.
Odothecium KRITSKY, BOEGER & JÉGU, 1997
straight or slightly curved. Accessory piece articulated to MCO by
copulatory ligament. Seminal vesicle C-shaped. Prostatic reservoirs round
to short, ovate. Germarium ovate or elongate. Vagina middorsal, looping
left caecum, sclerotized. Hooks dactylogyrid, shank divided in two
of two roots. Ventral bar rod-shaped, with slightly projected anterior
margin, with or without medial cleft; posterior projection absent. Dorsal
bar slightly U-shaped; anterior, posterior projections absent. Parasites of
gills of species of Serrasalmidae (Characiformes).
Type and only species: O. raphidiophallum KRITSKY, BOEGER & JÉGU, 1997
from Catoprion mento (Figs. 3-43 A-B)
Pavanelliella KRITSKY & BOEGER, 1998
present. Haptor not differentiated from trunk. Pharynx bulbous, one
subunit. Ceca confluent. Gonads overlapping. Copulatory organ com-
posed of MCO, accessory piece. MCO coiled. Accessory piece one, non-
ovate. Germarium ovate or elongate. Vagina sinistro-marginal, sclerotized.
Hooks dactylogyrid, shank divided in two portions clearly defined; 14 on
posterior margin of haptor. Anchors, bars absent. Parasites of the nasal
fossae of species of Pimelodidae (Siluriformes).
P. pavanellii KRITSKY & BOEGER, 1998 from Callophysus macropterus, Pseudoplatys-
toma coruscans (Type species) (Fig. 3-44).
P. scaphiocotylus KRITSKY & MENDOZA-FRANCO, 2003 from Rhamdia guatemalen-
sis.
Philocorydoras SURIANO, 1986

Body fusiform. Tegument smooth. Eyes two; eye granules absent or present.
Pharynx bulbous, one subunit. Ceca confluent. Gonads overlapping. Copu-
latory organ composed of MCO, accessory piece. MCO straight or slightly
curved. Accessory piece articulated to MCO by copulatory ligament.
Seminal vesicle fusiform. Prostatic reservoirs round to short, ovate. Germa-
rium ovate or elongate. Vagina ventral, muscular. Hooks dactylogyrid, shank
simple; 14, with ancyrocephaline distribution (4 dorsal, 10 ventral). Ventral,
dorsal anchor pairs present. Ventral anchor with point, shaft, roots well
defined. Dorsal anchor with shaft, point, base composed of two roots.
Ventral bar slightly U-shaped or rod-shaped; anterior projection absent;
posterior projection present. Dorsal bar slightly U-shaped or rod-shaped;
small subterminal knob at each extremity; posterior projections absent.
Parasites of gills of species of Callichthyidae (Siluriformes).
Type and only species: P. platensis SURIANO, 1986 from Corydoras paleatus
(Figs. 3-45 A-B).
Pithanothecium KRITSKY, BOEGER & JÉGU, 1997
Synonyms: Urocleidus (in part); Cleidodiscus (in part).
Body fusiform. Tegument scaled; annulations on body present. Eyes four;
eye granules absent or present. Pharynx bulbous, one subunit. Ceca
confluent. Gonads overlapping. Copulatory organ composed of MCO,
accessory piece. MCO double. MCO straight or slightly curved. Accessory
piece articulated to MCO by copulatory ligament. Seminal vesicle fusiform.
Vagina dextro-marginal sclerotized. Hooks dactylogyrid, shank divided in
two portions clearly defined; 14, with ancyrocephaline distribution (4 dorsal,
two roots. Ventral bar slightly U-shaped; anterior, posterior projections
absent. Dorsal bar slightly U-shaped; anterior, posterior projections absent.
Parasites of the gills of species of Serrasalmidae (Characiformes).
P. amazonensis (MIZELLE & PRICE, 1965) from Pygopristis denticulata, Pygocentrus
nattereri, Catoprion mento, Pristobrycon striolatus.
P. piranhus (KRITSKY, MIZELLE & PRICE, 1965) from Pygopristis denticulata,
Pygocentrus nattereri, Pristobrycon striolatus, Catoprion mento (Type species) (Fig.
3-46).
Protorhinoxenus DOMINGUES & BOEGER, 2002
Accessory piece articulated to MCO by copulatory ligament. Seminal
vesicle fusiform. Prostatic reservoirs unknown. Germarium ovate or
elongate. Vagina dextro-marginal, sclerotized. Hooks dactylogyrid, shank

anchor with point, shaft, base very elongated with roots not defined;
accessory sclerite associated with base of ventral anchor. Dorsal anchor
with point, shaft, base very elongate with roots not defined. Ventral bar
rod-shaped; anterior, posterior projections absent. Dorsal bar rod-shaped;
anterior, posterior projections absent. Parasites of the gills of species of
Prochilodus lineatus Prochilodontidae (Characiformes).
Type and only species: P. prochilodi DOMINGUES & BOEGER, 2002 from
Prochilodus lineatus (Fig. 3-47 A-D).
Pseudovancleaveus FRANÇA, ISSAC, PAVANELLI & TAKEMOTO, 2003
Synonyms: Vancleaveus (in part).
Body strongly flattened or fusiform. Tegument smooth. Eyes absent; eye
posed of MCO, accessory piece. MCO coiled. Accessory piece articulated
to MCO by copulatory ligament. Seminal vesicle fusiform. Prostatic
sinistro-ventral, muscular. Hooks dactylogyrid, shank simple; 14, with
ancyrocephaline distribution (4 dorsal, 10 ventral). Ventral, dorsal anchor
pairs present. Ventral anchor with point, shaft, roots well defined. Dorsal
anchor with shaft, point, base composed of two roots. Ventral bar slightly
V-shaped; anterior projection absent; posterior projection present. Dorsal
bar slightly U-shaped; anterior, posterior projections absent. Parasites of
gills of Pimelodidae (Siluriformes).
P. paranaensis FRANÇA, ISSAC, PAVANELLI & TAKEMOTO, 2003 from Iheringinchthys
labrosus (Type species) (Fig. 3-48).
P. platensis (SURIANO & INCORVAIA, 1995) from Iheringinchthys labrosus, Pimelodus
clarias maculatus.
Rhinonastes KRITSKY, THATCHER & BOEGER, 1988
Body short, robust, or strongly flattened. Tegument smooth. Eyes four;
eye granules absent or present. Pharynx bulbous, one subunit. Gonads
overlapping. Copulatory organ composed of MCO, accessory piece. MCO
coiled. Accessory piece articulated directly to MCO. Seminal vesicle
fusiform. Prostatic reservoirs round to short, ovate. Germarium ovate or
elongate. Vagina sinistro-marginal, sclerotized. Hooks dactylogyrid, shank
divided in two portions clearly defined; 14, 12 ventral distributed along
margin of haptor, 2 central between anchors. Ventral anchor pair present;
dorsal anchors absent. Ventral anchor with point, shaft, base comprising a
distinct subunit, robust, long, lacking definition of roots. Ventral bar rod-
shaped; two submedian elongate processes on anterior margin; posterior
median projection present. Parasites of nose of species of Prochilodon-
tidae (Characiformes).
Type and only species: R. pseudocapsaloideum KRITSKY, THATCHER & BOEGER,
1988 from Prochilodus nigricans (Figs. 3-49 A-E).
Rhinoxenus KRITSKY, BOEGER & THATCHER, 1988

Body short, robust, or fusiform. Tegument smooth. Eyes four; eye granules
absent or present. Pharynx bulbous, one subunit. Ceca confluent. Gonads
coiled. Accessory piece articulated directly to MCO. Seminal vesicle fusi-
form. Prostatic reservoirs round to short, ovate. Germarium ovate or
elongate. Vagina sinistro-marginal, sclerotized. Hooks dactylogyrid, shank
anchor with point, shaft, roots well defined or not defined. Dorsal anchor
spike like. Ventral bar rod-shaped; anterior, posterior projections absent.
Dorsal bar absent. Parasites of nasal fossae of species of Characiformes.
R. anaclaudiae DOMINGUES & BOEGER, 2005 from Triportheus cf. nematurus,
Brycon sp., Triportheus sp. (Characidae, Characiformes).
R. arietinus KRITSKY, BOEGER & THATCHER, 1988 from Rhytiodus argenteofuscus,
Schizodon fasciatum (Anostomidae).
R. bulbovaginatus BOEGER, DOMINGUES & PAVANELLI, 1995 from Salminus
maxillosus (Characidae).
R. curimbatae DOMINGUES & BOEGER, 2005 from Prochilodus cf. lineatus (Prochil-
odontidae, Characiformes).
R. euryxenus DOMINGUES & BOEGER, 2005 from Serrasalmus marginatus, S.
gouldingi, S. rhombeus, S. pilopleura, S. striolatus (Serrasalmidae, Characi-
formes), Leporinus agassizii (Anostomatidae, Characiforems).
R. guianensis DOMINGUES & BOEGER, 2005, from Curimata cyprinoides (Curima-
tidae, Characiformes).
R. nyttus KRITSKY, BOEGER & THATCHER, 1988 from Schizodon fasciatum (Anos-
tomidae).
R. piranhas KRITSKY, BOEGER & THATCHER, 1988 from Pygocentrus nattereri
(Serrasalmidae) (Type species) (Figs. 3-50 A-C).
Sciadicleithrum KRITSKY, THATCHER & BOEGER, 1989
Body short, robust, or cylindrical, or fusiform. Tegument smooth. Eyes
four or two; eye granules absent or present. Pharynx bulbous, one subunit.
Ceca confluent. Gonads overlapping. Copulatory organ composed of
MCO, accessory piece. MCO sinuous, or coiled. Accessory piece one, non-
ovate. Germarium ovate or elongate. Vagina ventral, or dextro-marginal,
or sinistro-marginal; sclerotized or muscular. Hooks dactylogyrid, shank
Ventral bar slightly V-shaped, or slightly U-shaped, or rod-shaped; with
double umbelliform membranes on anterior margin; posterior projection
absent. Dorsal bar rod-shaped; anterior, posterior projections absent.
Parasites of gills of species of Cichlidae (Perciformes).
S. aequidens (PRICE & SCHLUETER, 1989) from Aequidens maroni.

S. bicuense VIDAL-MARTÍNEZ, SCHOLZ & AGUIRRE-MACEDO, 2001 from Archocen-
trus nigrofasciatus.
S. bravohollisae KRITSKY, THATCHER & BOEGER, 1989 from Cichlasoma synspilum,
Petenia splendida, Cichlasoma pearsei.
S. cavanaughi (PRICE, 1966) from Aequidens maroni.
S. ergensi KRITSKY, THATCHER & BOEGER, 1989 from Cichla ocellaris.
S. geophagi KRITSKY, THATCHER & BOEGER, 1989 from Geophagus surinamensis.
S. iphthimum KRITSKY, THATCHER & BOEGER, 1989 from Pterophyllum scalare
(Figs. 3-51 A-F).
S. maculicaudae VIDAL-MARTÍNEZ, SCHOLZ & AGUIRRE-MACEDO, 2001 from
Cichlasoma maculicauda.
S. mexicanum KRITSKY, VIDAL-MARTÍNEZ & RODRIGUES-CANUL, 1994 from
Cichlasoma urophthalmus.
S. nicaraguense VIDAL-MARTÍNEZ, SCHOLZ & AGUIRRE-MACEDO, 2001 from
Amphilophus alfari.
S. splendidae KRITSKY, VIDAL-MARTÍNEZ & RODRIGUES-CANUL, 1994 from
Petenia splendida.
S. tortrix KRITSKY, THATCHER & BOEGER, 1989 from Uaru amphiacanthoides.
S. umbilicum KRITSKY, THATCHER & BOEGER, 1989 from Cichla ocellaris.
S. uncinatum KRITSKY, THATCHER & BOEGER, 1989 from Cichla ocelaris (Type
species).
S. variabilum (MIZELLE & KRITSKY, 1969) from Symphysodon discus.
S. meekii MENDOZA-FRANCO, SCHOLZ & VIDAL-MARTÍNEZ, 1997 from Archocen-
trus nigrofasciatus.
Telethecium KRITSKY, VAN EVERY & BOEGER, 1996
Body fusiform. Tegument smooth. Eyes four; eye granules absent. Haptor not
differentiated from trunk. Pharynx bulbous, one subunit. Ceca confluent.
Gonads overlapping. Copulatory organ composed of MCO, accessory piece.
MCO coiled. Accessory piece articulated directly to MCO, or articulated to
MCO by copulatory ligament. Seminal vesicle fusiform. Prostatic reservoirs
round to short, ovate. Germarium ovate or elongate. Vagina sinistro-marginal,
sclerotized. Hooks dactylogyrid, shank divided in two portions clearly defined;
14 on posterior margin of haptor. Ventral, dorsal anchor pairs absent. Ventral,
dorsal bars absent. Parasites of nose of species of Osteoglossidae (Osteoglo-
ssiformes) and Pristigasteridae (Clupeiformes).
T. nasalis KRITSKY, VAN EVERY & BOEGER, 1996 from Osteoglossum bicirrhosum
(Osteoglossidae) (Type species) (Figs. 3-52 A-B).
T. paniculum KRITSKY, VAN EVERY & BOEGER, 1996 from Pellona flavipinnis
(Pristigasteridae).
Tereancistrium KRITSKY, THATCHER & KAYTON, 1980
Body fusiform. Tegument smooth. Eyes four, or two; eye granules absent
or present. Pharynx bulbous, one subunit. Ceca confluent. Gonads
coiled. Accessory piece one, non-articulated. Seminal vesicle fusiform.

Vagina sinistro-marginal, sclerotized or muscular. Hooks dactylogyrid,
shank simple; 14, with ancyrocephaline distribution (4 dorsal, 10 ventral).
well defined; accessory sclerite associated with base present. Dorsal
anchor with shaft, point, base composed of two roots. Ventral bar rod-
shaped; anterior margin smooth or with slightly projected margin, with or
without medial cleft. Dorsal bar slightly V-shaped, or strongly V-shaped;
anterior projection absent; posterior projection present or absent. Parasites
of gills of species of Characiformes.
T. kerri KRITSKY, THATCHER & KAYTON, 1980 from Brycon melanopterus (Charac-
idae) (Type species) (Figs. 3-53 A-B).
T. ornatus KRITSKY, THATCHER & KAYTON, 1980 from Prochilodus reticulatus
(Curimatidae).
T. parvus KRITSKY, THATCHER & KAYTON, 1980 from Leporinus fasciatus (Anos-
tomidae).
Trinibaculum KRITSKY, THATCHER & KAYTON, 1980
Body short, robust. Tegument smooth. Eyes four; eye granules absent or
germarium anterior to testis. Copulatory organ composed of MCO, accesso-
ry piece. MCO straight or slightly curved. Accessory piece articulated
directly to MCO. Seminal vesicle fusiform. Prostatic reservoirs round to
short, ovate. Germarium ovate or elongate. Vagina dextro-ventral, sclero-
tized. Hooks dactylogyrid, shank simple; 14, with ancyrocephaline distribu-
anchor with point, shaft, roots well defined. Dorsal anchor with shaft, point,
base composed of two roots. Ventral bar slightly V-shaped; anterior,
posterior projections absent. Dorsal bar double, rod-shaped; anterior,
posterior projections absent. Parasites of gills of species of Characiformes.
Type and only species: T. brazilensis KRITSKY, THATCHER & KAYTON, 1980
from Brycon melanopterus (Fig. 3-54).
Trinidactylus HANEK, MOLNAR & FERNANDO, 1974
Body fusiform. Tegument smooth. Annulations on body absent. Eyes
four, eye granules absent or present. Pharynx bulbous, one subunit.
Gonads tandem; germarium anterior to testis. Testis single. Copulatory
organ composed of MCO, accessory piece. MCO coiled, loose coils, with
non-inflated proximal portion. Accessory piece one, non-articulated.
Seminal vesicle unknown. Prostatic reservoirs unknown. Germarium ovate
or elongate, solid. Vagina single, ventral. Vaginal bar, accessory vaginal
sclerite absent. Hooks dactylogyrid. Hook shank simple. Hooks 14, with
ancyrocephaline distribution (4 dorsal, 10 ventral). Hooks not on pedun-
cles. Hook pair 2 on the haptor, close to hooks pairs 3, 4; hook pair 5
similar to other pairs. Ventral, dorsal anchor present. Ventral anchor rod-
shaped; ventral anchor filament delicate, often inconspicuous. Accessory

sclerite associated with base of ventral anchor absent. Dorsal anchor
dactylogyrid, with shaft, point and base composed of two roots. Fold on
inner superficial root of dorsal anchor absent. Ventral bar rod-shaped.,
with smooth surface on anterior margins. Groove along length of ventral
bar absent. Ventral bar without posterior median projection. Dorsal bar
absent. Parasite of species of Cichlidae (Perciformes).
Type and only species: T. cichlasomatis HANEK, MOLNAR & FERNANDO, 1974
from Cichlasoma bimaculatum (Fig. 3-55).
Trinigyrus HANEK, MOLNAR & FERNANDO, 1974
Body fusiform. Tegument smooth. Eyes absent; eye granules absent.
Pharynx bulbous, one subunit. Ceca confluent. Gonads overlapping.
Copulatory organ composed of MCO, accessory piece. MCO sinuous, or
straight, or slightly curved. Accessory piece one, non-articulated. Seminal
vesicle fusiform. Prostatic reservoirs round to short, ovate. Germarium
ovate or elongate. Vagina dextro-marginal, sclerotized or muscular. Hooks
dactylogyrid, shank divided in two portions clearly defined; 14, with
ancyrocephaline distribution (4 dorsal, 10 ventral). Ventral anchor with
point, shaft, roots not defined. Dorsal anchor absent. Ventral bar with
tapering extremities, rod-shaped, anterior absent; posterior projections
present; groove along entire length of bar present. Dorsal bar absent.
Parasites of gills of species of Loricariidae (Siluriformes).
T. acuminatus KRITSKY, BOEGER & THATCHER, 1986 from Acanthicus hystrix
(Figs. 3-56 A-E).
T. hypostomatis HANEK, MOLNAR & FERNANDO, 1974 from Hypostomus robinii
(Type species).
T. mourei BOEGER & JÉGU, 1994 from Hypostomus marginatus.
T. tentaculoides KRITSKY, BOEGER & THATCHER, 1986 from Hypoptopoma thora-
cathum.
Unibarra SURIANO & INCORVAIA, 1995
Body fusiform. Tegument smooth. Eyes absent; eye granules absent.
Pharynx bulbous, one subunit. Ceca confluent. Gonads tandem; germari-
um anterior to testis. Copulatory organ composed of MCO, accessory
piece. MCO straight or slightly curved. Seminal vesicle fusiform. Prostatic
ventral, sclerotized. Hooks dactylogyrid, entire shank inflated, conferring a
robust morphology to hook; 14, with ancyrocephaline distribution (4
point, shaft, roots not defined. Dorsal anchor with shaft, point, base not
clearly separated in roots. Ventral bar rod-shaped; anterior projection
absent; posterior projection present. Dorsal bar absent. Parasites of gills
of species of Pimelodidae (Siluriformes).
Type and only species: U. paranoplatensis SURIANO & INCORVAIA, 1995 from
Pimelodus clarias maculatus, Paulicea luetkeni (Fig. 3-57)
Unilatus MIZELLE & KRITSKY, 1967

Synonym: Diaccessorius
germarium anterior to testis. Copulatory organ composed of MCO, accessory
piece. MCO coiled, with tight coils, cork-screw like. Accessory piece one, non-
ovate. Germarium ovate or elongate. Vagina ventral or sinistro-marginal,
sclerotized or muscular. Hooks dactylogyrid, shank divided in two portions
clearly defined or shank simple; 14, 8 dorsal, 6 ventral. Two pairs of dorsal
anchors, anterior, posterior. Posterior pair of anchors with point, shaft, roots
not defined. Anterior pair of anchors with shaft, point, base composed of two
roots. Posterior bar slightly V-shaped or rod-shaped; anterior projection
absent; posterior projection absent or present. Anterior bar slightly U-shaped
or strongly V-shaped; anterior projection absent; posterior projections present
or absent. Parasites of gills of species of Loricariidae (Siluriformes).
U. anoculus (PRICE, 1968) from Hypostomus bolivianus.
U. brevispinus SURIANO, 1985 from Pterigoplichthys multiradiatus.
U. brittani MIZELLE, KRITSKY & CRANE, 1968 from Hypostomus sp., Plecostomus sp.
U. dissimilis SURIANO, 1985 from Hemiancistris sp.
U. longispinus SURIANO, 1985 from Pterigopliichthys multiradiatus.
U. scaphirhynchae SURIANO, 1985 from Hemiancistrus scaphirhynchae.
U. unilatus MIZELLE & KRITSKY, 1967 from Hypostomus robinii, Plecostomus sp.
(Type species) (Figs. 3-58 A-E).
Urocleidoides sensu stricto MIZELLE & PRICE, 1964
Body fusiform. Tegument smooth. Eyes four, or two, or absent; eye granules
absent or present. Pharynx bulbous, one subunit. Ceca confluent. Gonads
coiled. Accessory piece one, non-articulated, or articulated directly to MCO,
or articulated to MCO by copulatory ligament. Seminal vesicle fusiform.
Vagina dextral or sinistro-marginal; sclerotized or muscular; vaginal accesso-
ry sclerite present. Hooks dactylogyrid, shank divided in two portions clearly
defined, or shank simple; 14, with ancyrocephaline distribution (4 dorsal, 10
ventral). Ventral, dorsal anchor pairs present. Ventral anchor with point,
two roots. Ventral bar slightly V-shaped, or slightly U-shaped, or rod-shaped;
anterior, posterior projections absent. Dorsal bar slightly V-shaped, or
slightly U-shaped; anterior, posterior projections absent. Parasites of species
of Cyprinodontiformes and Characiformes.
U. anops KRITSKY & THATCHER, 1974 from Characidium caucanum (Characidae,
Characiformes).
U. curimatae MOLNAR, HANEK & FERNANDO, 1974 from Curimata argentea
(Curimatidae, Characiformes).
U. eremitus KRITSKY, THATCHER & BOEGER, 1986 from Hoplias malabaricus

(Erythrinidae, Characiformes).
U. paradoxus KRITSKY, THATCHER & BOEGER, 1986 from Rhytiodus microlepis
(Anostomidae, Characiformes) (Figs. 3-59 A-D).
U. reticulatus MIZELLE & PRICE, 1964 from Lebistes reticulata (Poeciliidae,
Cyprinodontiformes) (Type species).
U. vaginoclaustrum JOGUNOORI, KRITSKY & VENKATANARASAIAH, 2004 from
Xiphophorus helleri (Poeciliidae, Cyprinodontiformes).
Vancleaveus KRITSKY, THATCHER & BOEGER, 1986
ping. Copulatory organ composed of MCO, accessory piece. MCO J-
shaped. Accessory piece one, non-articulated. Seminal vesicle fusiform,
with thick walls, very long, extending from level of base of MCO to initial
portion of germarium. Prostatic reservoirs very long, often looping
posteriorly. Germarium ovate or elongate. Vagina ventral or dextro-
ventral, sclerotized. Hooks dactylogyrid, entire shank inflated, conferring a
robust morphology to hook; 14, with ancyrocephaline distribution (4
with point, shaft, roots well defined. Dorsal anchor with shaft, point, base
composed of two roots; fold on inner superficial root of dorsal anchor
present. Ventral bar slightly V-shaped or rod-shaped; anterior projection
present; dorsal projection absent. Dorsal bar rod-shaped; anterior projec-
tion absent or present; posterior projection absent. Parasites of gills of
species of Siluriformes, most commonly on Pimelodidae.
V. cicinnus KRITSKY, THATCHER & BOEGER, 1986 from Pimelodus spp., Phracto-
cephalus hemiliopterus (Pimelodidae).
V. fungulus KRITSKY, THATCHER & BOEGER, 1986 from Pseudoplatystoma sp.,
Pseudoplatystoma fasciatum (Pimelodidae).
V. janauacaensis KRITSKY, THATCHER & BOEGER, 1986 from Pterodoras granulosus
(Doradidae) (Type species) (Figs. 3-60 A-B).
V. platyrhynchi KRITSKY, THATCHER & BOEGER, 1986 from Hemisorubim platyrhyn-
chos (Pimelodidae).
Genera of Dactylogyridae, Ancyrocephalinae not well known or of questionable status.
The diagnosis and/or species composition of the genera listed below is
complicated by inadequate species descriptions, lack of evident diagnostic
features, or represent catchall taxa that require extensive revision.
Amphocleithrium PRICE & ROMERO, 1969
A. paraguayensis PRICE & ROMERO, 1969 from Pseudoplatystoma sp., Pseudoplatysto-
ma coruscans (Pimelodidae, Siluriformes).
Cleidodiscus MUELLER, 1934
C. microcirrus PRICE & SCHLUETER, 1967 from Hemiodus semitaeniatus (Hemi-
odontidae, Characiformes).
Palombitrema PRICE, C.E. & W.A. BUSSING, 1968

Syn: Androspira SURIANO, 1981
P. heteroancistrium (PRICE & BUSSING, 1968) from Astyanax bimaculatus (Charac-
idae, Characiformes).
P. chascomusense (SURIANO, 1981) from Curimata gilberti (Curimatidae, Characi-
formes).
P. triangulum (SURIANO, 1981) from Curitimata gilberti (Curimatidae, Characi-
formes).
Salsuginus BEVERLEY-BURTON, 1984
Salsuginus neotropicalis MENDOZA-FRANCO & VIDAL-MARTINEZ, 2001 from
Belonesox belizanus (Atheriniformes, Poeciliidae).
Urocleidoides sensu lato
U. amazonensis MIZELLE & KRITSKY, 1969 from Phractocephalus hemiliopterus
(Pimelodidae, Siluriformes).
U. astyanacis GIOIA, CORDEIRO & ARTIGAS, 1988 from Astyanax scabripinnis,
Astyanax fasciatus (Characidae, Characiformes).
U. carapus MIZELLE, KRITSKY & CRANE, 1968 from Gymnotus carapo (Gym-
notidae, Gymnotiformes).
U. catus MIZELLE & KRITSKY, 1969 from Phractocephalus hemiliopterus (Pimelo-
didae, Siluriformes).
U. corydori MOLNAR, HANEK & FERNANDO, 1974 from Corydoras aeneus (Callich-
thyidae, Siluriformes).
U. costaricensis (PRICE & BUSSING, 1967) from Curimata argentea (Curimatidae,
Characiformes), Astyanax bimaculatus (Characidae, Characiformes) Asty-
anax fasciatus (Characidae, Characiformes).
U. gymnotus MIZELLE, KRITSKY & CRANE, 1968 from Gymnotus carapo (Gym-
notidae, Gymnotiformes).
U. lebedevi KRITSKY & THATCHER, 1976 from Pimelodus grosskopfi (Pimelodidae,
Siluriformes).
U. margolisi MOLNAR, HANEK & FERNANDO, 1974 from Corydoras aeneus (Calli-
chthyidae, Siluriformes).
U. megorchis MIZELLE & KRITSKY, 1969 from Sorubim lima (Pimelodidae,
Siluriformes).
U. strombicirrus (PRICE & BUSSING, 1968) from Astyanax fasciatus (Characidae,
Characiformes).
U. trinidadensis MOLNAR, HANEK & FERNANDO, 1974 from Astyanax bimaculatus
(Characidae, Characiformes).
U. virescens MIZELLE, KRITSKY & CRANE, 1968 from Eigenmannia virescens
(Sternopygidae, Gymnotiformes).
VIII. Plates of Monogenoidea (Figs. 3-1 to 3-64)
Abreviation used:
ap – accessory piece mg – Mehlis gland
ascl – accessory sclerite ph – pharynx
bo – buccal organ pr – prostatic reservoir
ca – caecum pt – point
cg – cephalic gland sb – superficial bar
cl – copulatory ligament sf – shaft
db – deep bar sk – shank
dr – deep root sl – spinelet
e – embryo sp – spine
eg – egg sr – seminal receptacle
fh – FH loop sr – superficial root
ge – germarium sv – seminal vesicle
he – heel te – testis
ho – head organ to – toe
hs – haptoral sucker va – vagina
mco – male copulatory organ vi – vitellaria
bo
1 mm
50
3-2B
hs
150
3-1B 3-1C
500
3-2A
50
3-1A
sr 3-4B
dr
50
ph
30
sf pt
mco
50
3-3B
sv
3-4C e
250
3-4D
ge
te 3-4F
ca
50
50
1
2
3
4
3-3A 3-4E 3-4A 5
6
8 7
3-1. Paraheteronchocotyle amazonensis: A. entire; B. large sucker sclerite; C. small sucker sclerite; 3-2.
Paranaella luquei: A. entire; B. clamp; 3-3. Potamotrygonocotyle tsalickisi: A. entire; B. anchor; 3-4. Accessorius
peruensis: A. entire; B. accessory sclerite; C. male copulatory organ; D. marginal hook; E. ventral bar-
shield complex; F. anchor; hooks are numbered according to the system proposed by MIZELLE (1936).
All scales in micrometers (µm), except 3-2A.
pt sp 3-6B
sf
15
100
to
he 4
sk
20
fh
sl 3-6C
3-5B 3-5C
20
db 3
3-6D
sb
20
20
40
af
3-5A 3-6E 3-6A
ho
3-8B
25
cg
50
200
sv
25
eg
3-7B
mg
25
25
ge
3-8C 3-8D
3-7C
25
50
3-7D
200
3-8E 3-8A
3-7A
3-5. Anacanthocotyle anacanthocotyle: A. entire; B. marginal hook; C. male copulatory organ; 3-6.
Gyrodactylus geophagensis: A. entire; B. marginal hook; C. male copulatory organ; D. anchor; E. anchor-
superficial bar complex; 3-7. Hyperopletes malmbergi: A. entire; B. male copulatory organ; C. marginal
hook; D. anchor-superficial bar complex; 3-8. Nothogyrodactylus plaesiophallus: A. entire; B. accessory
copulatory sclerites; C. anchor; D. marginal hook; E. anchor-superficial bar complex. All scales in
micrometers (µm).
100
25
20
3-10B
3-9B
50
70
50
25
3-10C 3-10D
70
200
3-9C
3-9A 3-9D 3-10E

3-10A
20
20
3-12B
3-11B 3-11C
40
40
20
50
3-12C 3-12D
3-11D
100
3-12E
40
3-11E
sq
25
3-12F
3-11A 3-11F 3-12A
3-9. Oogyrodactylus farlowellae: A. entire; B. male copulatory organ; C. marginal hook; D. anchor; 3-10.
Phanerothecium caballeroi: A. entire; B. male copulatory organ; C. marginal hook; D. anchor; E. anchor-
superficial bar complex; 3-11. Scleroductus sp.: A. entire (no publish data); B. superficial bar-shield
complex; C-D. male copulatory organ seen from different angle; E. marginal hook; F. anchor; 3-12.
Diplectanum decorum: A. entire (composite); B. vagina; C. copulatory complex; D. ventral anchor; E.
ventral bar; F. dorsal bar. All scales in micrometers (µm).
3-13B 3-14B 3-14C
vg
ap
ap
25
cl
200
30
mco
vi
mco 3-13C 3-14D
cl
3-14E
100
3-13D
1 2
3
1 5 4
2 7 6
3 4 76
3-13A 3-13E 3-14F 5
3-14A
100
3-16B
20
va
3-15B
25
3-16C
vi
te
3-16D
20
100
3-15C 3-16E
3-15A 3-16A
3-13. Ameloblastella chavarrai: A. entire; B. vagina; C. male copulatory organ; D. ventral bar; E. dorsal
bar; 3-14. Amphithecium calycinum: A. entire; B. ventral anchor. C. dorsal anchor; D. male copulatory
organ; E. ventral bar; F. dorsal bar; hooks are numbered according to the system proposed by MIZELLE
(1936); 3-15. Anacanthoroides mizellei: A. entire; B. male copulatory organ; C. hooks; 3-16. Anacanthorus
sciponophallus: A. entire; B-D. hooks; E. male copulatory organ. All scales in micrometers (µm).
3-17B
3-17C 3-17D
3-17A
mco
3-17E
ap
3-17G 3-17H
3-17F
3-17I
3-17J
3-17K 3-17L
3-17M 3-17N 3-17O 3-17P
3-17. Male copulatory organ of Anacanthorus spp.: A. A. bellus; B. A. calophallus; C. A. carinatus; D. A.

cornutus; E. A. dipelecinus; F. A. glyptophallus; G. A. furculus; H. A. euryphallus; I. A. hoplophallus; J. A.
mastigophallus; K. A. pithophallus; L.A. periphallus; M. A. lygophallus; N. A. pelorophallus; O. A. xaniophallus;
P. A. spinatus. All scales = 25 micrometers (µm).
3-18B
25
3-19B
100
100
25
3-18C
3-19A
3-18A 25
3-21B
100
100
sr
3-20B
3-20C
25
3-20D
3-20A 3-21A
3-18. Ancistrohaptor falciferum: A. entire; B. ventral bar; C. male copulatory organ; 3-19. Annulotrematoides
amazonicus: A. entire; B. male copulatory organ; 3-20. Aphanoblastela travassoi: A. entire; B. male
copulatory organ; C. ventral bar; D. dorsal bar; 3-21. Cacatuocotyle paranaensis: A. entire; B. male
copulatory organ. All scales in micrometers (µm).
25
100
3-22B
pr
100
25
3-23B
3-22A 3-23C 3-23A
3-25B
30
200
3-24B
100
sv
3-24C
50
3-24D
30
3-25C
3-24E 3-24F
3-24A
3-25D 3-25A
3-22. Calpidothecioides pygopristi: A. entire; B. ventral bar; 3-23. Calpidothecium crescentis: A. entire; B.
ventral anchor; C. dorsal anchor; 3-24. Cosmetocleithrum gussevi: A. entire; B. ventral bar; C. dorsal bar;
D. male copulatory organ; E. ventral anchor; F. dorsal anchor; 3-25. Curvianchoratus singularis: A.
entire; B-C. dorsal anchor; D. ventral anchor. All scales in micrometers (µm).
25
500
200
3-26B 3-27B
3-26C
25
3-27C
30
3-27D
25
3-28B 3-27E 3-27F
3-26A 3-27A
20
50
200
3-28C
50
50
3-28A 3-30B 3-29 3-30A
3-26. Dawestrema cycloancistrium: A. entire; B. vagina; C. ventral bar; 3-27. Demidospermus cornicinus: A.
entire; B. vagina; C. ventral bar; D. dorsal bar; E-F. hooks; 3-28. Diaphorocleidus armillatus: A. entire; B.
male copulatory organ; C. ventral bar; 3-29. Enallothecium aegidatum; 3-30. Euryhaliotrema chaoi: A.
entire; B. male copulatory organ. All scales in micrometers (µm).
100
3-32B
100
30
3-31B
vag
30
vgb
3-31C 3-32C
3-32D
3-31A 3-31D 3-32A

100
3-34B
50
20
3-34C
3-33 3-34A
3-34D
3-31. Gonocleithrum planacrus: A. entire; B. gonadal bar; C. ventral bar; D. male copulatory organ; 3-32.
Gussevia asota: A. entire; B. dorsal bar; C. ventral anchor; D. male copulatory organ; 3-33. Heterothecium
globatum; 3-34. Heterotylus heterotylus: A. entire; B. ventral bar; C. male copulatory organ; D. ventral
anchor. All scales in micrometers (µm).
10
50
200
3-35B
25
3-36B
10
3-35C
3-36C
3-35A 3-35D 3-36A

10
pr
10
3-37B 3-37C
500
10
3-37D
3-37A
3-35. Jainus sp.: A. entire; B. male copulatory organ; C. hook; D. ventral anchor; 3-36. Kritskyia moraveci:
A. entire; B. hook (ventral view); C. hook; 3-37. Liguadactyloides brinkmanni: A. entire; B. ventral anchor;
C. dorsal anchor; D. hook. All scales in micrometers (µm).
100
20
50
3-39C
30
40
3-38 3-39A 3-40B 3-40A
20
3-39B
300
30
100
3-42B
30
3-42C
3-41
3-42D 3-42A
3-38. Monocleithrium lavergneae; 3-39. Mymarothecium dactylotum: A. entire; B. ventral bar; C. male copu-
latory organ; 3-40. Notothecium mizellei: A. entire; B. dorsal bar; 3-41. Notothecioides llewellyni; 3-42.
Notozothecium penetrarum: A. entire; B. male copulatory organ; C. ventral bar; D. dorsal bar. All scales
in micrometers (µm).
100
25
200
3-43B
25
100
3-45B
3-43A 3-44
3-45A
100
50
100
3-47B
3-47C
50
3-46 3-47D 3-47A
3-43. Odothecium raphidiophallum: A. entire; B. ventral bar; 3-44. Pavanelliella pavanelli; 3-45. Philocorydoras
platensis: A. entire; B. ventral bar; 3-46. Pithanothecium piranhus; 3-47. Protorhinoxenus prochilodi: A. entire;
B. dorsal anchor; C. ventral anchor; D. hook. All scales in micrometers (µm).
100
125
50
3-49B 30 3-49C
20
15
3-48 3-49D
3-49E 3-49A
30
3-51B
30 100
200
3-51C
30
3-51D
30
3-50B
3-51E
3-50C
3-50A 3-51F 3-51A

3-48. Pseudovancleaveus paranaensis; 3-49. Rhinonastes pseudocapsaloideum: A. entire; B. ventral bar; C. anchor;
D. hook; E. male copulatory organ; 3-50. Rhinoxenus piranhas: A. entire; B. dorsal haptorial spike; C.
ventral anchor; 3-51. Sciadicleithrum iphthimum: A. entire; B-F. ventral bar; B. S. ergensi; C. S. uncinatum;
D. S. geophagi; E. S. umbilicum; F. S. tortrix. All scales in micrometers (µm).
100
100
3-52B
100
ascl
20
3-52A 3-53B 3-53A 3-54

30
40
100
3-55 3-56B
30
3-56D
3-56C
30
3-56A
30
3-56E
3-52. Telethecium nasalis: A. entire; B. lateral view of specimen showing relationship of body and
copulatory bag (scale not provide); 3-53. Tereancistrium kerri: A. entire; B. ventral anchor and accessory
anchor sclerite; 3-54. Trinibaculum brazilensis; 3-55. Trinidactylus cichlasomatis (haptor); 3-56. Trinigyrus
acuminatus: A. entire; B. anchor; C. bar; D male copulatory organ; E. vagina and distal portion of
seminal receptacle. All scales in micrometers (µm).
400
3-59B
30
3-58A 3-58B
3-59C
3-58D
100
3-58C
3-57
3-59D
30
3-59A
3-58E
30
100
3-60B
3-60A
3-57. Unibarra paranoplatensis; 3-58. Unilatus unilatus: A. anterior bar; B. anterior anchor; C. posterior
bar; D. posterior anchor; E. accessory piece; 3-59. Urocleioides paradoxus: A. entire; B. hook (pairs 1, 5);
C. hook (pairs 2, 3, 4, 6, 7); D. vaginal sclerite; 3-60. Vancleaveus janauacaensis: A. entire; B. dorsal bar.
All scales in micrometers (µm).
3-61
3-62
3-61. Histological section of gill filament showing haptor of Linguadactyloides with an anchor secured
in the cartilage; 3-62. Linguadactyloides penetrated into gill filament showing epithelial hyperplasia
around point of entry. (Original photographs).
3-63
3-64
250 µm
3-63. Linguadactyloides penetrated within gill filament showing missing ephitelium; 3-64. Undescribed
monogenoid penetrated into the gill filament of Oxydoras niger showing tumeroid caused by the
haptor. (Original photographs).
AGARWAL, N. & D.C. KRITSKY (1998): Neotropical Monogenoidea. 33. Three new species of
Ancistrohaptor n. g. (Dactylogyridae, Ancyrocephalinae) on Triportheus spp. (Teleostei, Charac-
idae) from Brazil, with checklists of ancyrocephalines recorded from Neotropical characi-
form fishes. - Systematic Parasitology 39(1): 59-69.
AN, L., JARA, C.A. & D.K. CONE (1991): Five species of Gyrodactylus NORDMANN, 1832 (Monogenea)
from freshwater fishes of Peru. - Can. J. Zool. 69: 1199-1201.
BELMONT-JÉGU, E., DOMINGUES, M.V. & M.L. MARTINS (2004): Notozothecium janauachensis n. sp.
(Monogenoidea: Dactylogyridae) from wild and cultured tambaqui, Colossoma macropomum
(Teleostei: Characidae: Serrasalminae) in Brazil. - Zootaxa 736: 1-8.
BOEGER, W.A. & E. BELMONT-JÉGU (1994): Neotropical Monogenoidea. 21. Trinigyrus mourei sp. n.
(Dactylogyridae) from the gills of the Amazonian catfish Hypostomus marginatus (Loricariidae).
- Amazoniana 13(1/2): 13-16.
BOEGER, W.A. & D.C. KRITSKY (1988): Neotropical Monogenea. 12. Dactylogyridae from Serrasal-
mus nattereri (Cypriniformes: Serrasalmidae) and aspects of their morphologic variation and
distribution in the Brazilian Amazon. - Proc. Helminthol. Soc. Wash. 55(2): 188-213.
BOEGER, W.A. & F. POPAZOGLO (1995): Neotropical Monogenoidea. 23. Two new species of
Gyrodactylus from cichlid and characid fishes of Brazil. - Mem. Inst. Oswaldo Cruz 90: 689-694.
BOEGER, W.A., DOMINGUES, M.V. & D.C. KRITSKY (1996): Neotropical Monogenoidea. 32.
Cacatuocotyle paranaensis n. g., n. sp. (Dactylogyridae, Ancyrocephalinae) from Characidium spp.
(Teleostei, Characidae) from the State of Paraná, Brazil. - Systematic Parasitology 37: 1-4.
BOEGER, W.A., DOMINGUES, M.V. & G.C. PAVANELLI (1997): Neotropical Monogenoidea. 24. Rhinoxenus
bulbovaginatus n. sp. (Ancyrocephalinae) from the nasal cavity of Salminus maxillosus (Osteichthyes,
Characidae) from the Rio Paraná, Parana, Brazil. - Mem. Inst. Oswaldo Cruz 90: 695-694.
BOEGER, W.A., HUSAK, W.S. & M.L. MARTINS (1995): Neotropical Monogenoidea. 25. Anacanthorus
penilabiatus n. sp. (Dactylogyridae, Anacanthorinae) from Piaractus mesopotamicus (Osteichthyes,
Serrasalmidae), cultivated in the State of São Paulo, Brazil. - Mem. Inst. Oswaldo Cruz 90: 699-701.
BOEGER, W.A., KRITSKY, D.C. & E. BELMONT-JÉGU (1994): Neotropical Monogenoidea. 20. Two new
species of oviparous Gyrodactylidea (Polyonchoinea) from loricariid catfishes (Siluriformes)
in Brazil and the phylogenetic status of Ooegyrodactylidae HARRIS, 1983. - J. Helminthol. Soc.
Wash. 61: 34-44.
BOEGER, W.A., KRITSKY, D.C. & M. PIE (2003): The context of diversification of the viviparous
Gyrodactylidae (Platyhelminthes: Monogenoidea). - Zoologica Scripta 32(5): 437-448.
BOEGER, W.A., PIASECKI, W. & E. SOBECKA (2002): Neotropical Monogenoidea. 44. Mymarothecium
viatorum sp. n. (Ancyrocephalinae) from the gill of Piaractus brachypomus (Serrasalmidae,
Teleostei) captured in a warm-water canal of a power plant in Szczecin, Poland. - Acta
Ichthyologica et Piscatoria 32(2): 158-161.
BOEGER, W.A., TANAKA, L.K. & G.C. PAVANELLI (2001): Neotropical Monogenoidea. 39. A new
species of Kritskyia (Dactylogyridae, Ancyrocephalinae) from the ureters and urinary bladder
of Serrasalmus marginatus and S. spilopleura (Characiformes, Serrasalmidae) from southern
Brazil with an emended generic diagnosis. - Zoosystema 23: 5-10.
BRANDES, G. (1894): Fridericianella ovicola n.g., n.sp. Ein neuer monogenetischer Trematode. - Abh.
Naturf. Ges. Halle 20, Jubiläums-Festschrift: 303-310.
BYCHOWSKY, B.E. (1957): Monogenetic trematodes, their systematics and phylogeny. - Akad. Nauk.
U.S.S.R., Moscow (translated from Russian by AIBS, Washington, D.C.): 509 pp.
CABLE, J.C., VAN OOOSTERHOUT, C., BARSON, N. & P.D. HARRIS (2005): Gyrodactylus pictae n. sp.
(Monogenea: Gyrodactylidae) from the Trinidadian swamp guppy Poecilia picta REGAN, with a
discussion on species of Gyrodactyls VON NORDMANN, 1832 and their poeciliid hosts. -
Systematic Parasitology 60: 159-164.
COHEN, S.C. & A. KOHN (2005): A new species of Mymarothecium and new host and geographical
records for M. viatorum (Monogenea: Dactylogyridae), parasites of freshwater fishes in Brazil.
- Folia Parasitologica 52: 307-310.
CUGLIANNA, A.N., CORDEIRO, N.S. & J.L. LUQUE (2003): Annulotrematoides bryconi sp. n. (Monogenea:
Dactylogyridae) parasitic on Brycon cephalus (Osteichthyes: Characidae) from Brazil. - Folia
Parasitologica 50: 272-274.
DALLWITZ, M. J. (1974): A flexible computer program for generating identification keys. -Systematic.
Zoology 23: 50–7.
DALLWITZ, M. J. (1980): A general system for coding taxonomic descriptions. -Taxon 29: 41–6.
DOMINGUES, M.V. & W.A. BOEGER (2002): Neotropical Monogenoidea. 40. Protorhinoxenus prochilodi
gen. n., sp. n. (Monogenoidea: Ancyrocephalinae), parasite of Prochilodus scrofa (Characi-
formes: Prochilodontidae) from South Brazil. - Folia Parasitologica 49: 35-38.
DOMINGUES, M.V. & W.A. BOEGER (2005): Neotropical Monogenoidea. 47. Phylogeny and coevolu-
tion of species of Rhinoxenus (Plathyhelminthes, Monogenoidea, Dactylogyridae) and their
characiformes hosts (Teloestei, Ostariophysi) with description of four new species. -Zoosyste-
ma 27(3): 441-467.
FERRAZ, E., SHINN, A.P. & C. SOMMERVILLE (1994): Gyrodactylus gemini n. sp. (Monogenea: Gyrodac-
tylidae), a parasite of Semaprochilodus taeniurus (Steindachner) from the Venezuelan Amazon. -
Systematic Parasitology 29: 217-222.
FRANCA, J.G., ISSAC, A., PAVANELLI, G.C. & R.M. TAKEMOTO (2003): Dactylogyridae (Monogenea) from
the gills of Iheringichthys labrosus (Osteichthyes: Pimelodidae) from the upper Paraná River
floodplain, Brazil, with the proposal of Pseudovancleaveus n. g. - Systematic Parasitology 54: 25-31.
GIOIA, I., CORDEIRO, N.S. & P.T. ARTIGAS (1988): Urocleidoides astyanacis n. sp. (Monogenea:
Ancyrocephalinae) from freshwater Characidians of the genus Astyanax. - Mem. Inst.
Oswaldo Cruz 83: 13-15.
GUIDELLI, G.M., TAKEMOTO, R.M. & G.C. PAVANELLI (2003): A new species of Kritskyia (Dactylogy-
ridae, Ancyrocephalinae), parasite of urinary bladder and ureters of Leporinus lacustris (Characi-
formes, Anostomidae) from Brazil. - Acta Scientiarum. Biological Sciences 25: 279-282.
GUTIÉRREZ, P.A. & D.M. SURIANO (1992): Ancyrocephalids of the genus Demidospermus suriano, 1983
(Monogenea) parasites from siluriform fishes in Argentina, with descriptions of three new
species. - Acta Parasitologica 37: 169-172.
HANEK, G., MOLNAR, K. & C.H. FERNANDO (1974): Three new genera of Dactylogyridae
(Monogenea) from freshwater fishes of Trinidad. - Parasit. 60: 911-913.
HARRIS, P.D. (1983): The morphology and life-cycle of the oviparous Oögyrodactylus farlowellae gen. et
sp. nov. (Monogenea, Gyrodactylidae). - Parasit. 87: 405-420.
HARRIS, P.D, & J. CABLE (2000): Gyrodactylus poeciliae n. sp. and G. milleri n. sp. (Monogenea:
Gyrodactylidae) from Poecilia caucana (STEINDACHNER) in Venezuela. - Systematic Parasitology
47(2): 79-85.
HARRIS, P.D. & A.M. LYLES (1992): Infections of Gyrodactylus turnbulli on guppies (Poecilia reticulata)
in Trinidad. - J. Parasitol. 78: 912-914.
JARA, C.A. & D.K. CONE (1989): Scleroductus yuncensi gen. et sp. n. (Monogenea) from Pimelodella
yuncensis (Siluriformes: Pimelodidae) in Peru. - Proc. Helminthol. Soc. Wash. 56: 125-127.
JARA, C., AN, L. & D. CONE (1991): Accessorius peruensis gen. et sp. n. (Monogenea: Gyrodactylidea)
from Lebiasina bimaculata (Characidae) in Peru. - J. Helminthol. Soc. Wash. 58: 164-166.
JOGUNOORI, W., KRITSKY, D.C. & J. VENKATANARASAIAH (2004): Neotropical Monogenoidea. 46. Three
new species from the gills of introduced aquarium fishes in India, the proposal of Heterotylus n. g.
and Diaphorocleidus n. g., and the reassignment of some previously described species of Urocleidoides
MIZELLE & PRICE, 1964 (Polyonchoinea: Dactylogyridae). - Systematic Parasitology 58: 115-124.
KOHN, A. (1990): Kritskyia moraveci n. g., n. sp. (Monogenea: Dactylogyridae) from the urinary
bladder and ureters of Rhamdia quelen (Quoy & Gaimard, 1824) (Pisces: Pimelodidae) in
Brazil. - Systematic Parasitology 17: 81-85.
KOHN, A. & I. PAPERNA (1964): Monogenetic trematodes from aquarium fishes. - Rev. Brasil. Biol.
24:145-149.
KOHN, A. & C.P. SANTOS (1989): Brazilian Monogenea. - List of species, hosts and geographical
distribution. - Rev. Brasil. Biol. 49(3): 809-815.
KOHN A., BAPTISTA-FARIAS, M.D. & S.C. COHEN (2000): Paranaella luquei gen. et sp. n. (Monogenea:
Microcotylidae), a new parasite of Brazilian catfishes. - Folia Parasitologica 47: 279-283.
KRITSKY, D.C. & W.A. BOEGER (1991): Neotropical Monogenea. 16. New species of oviparous
Gyrodactylidea with proposal of Nothogyrodactylus gen. n. (Oogyrodactylidae). - J. Helminthol.
Soc. Wash. 58: 7-15.
KRITSKY, D.C. & W.A. BOEGER (1995): Neotropical Monogenoidea. 26. Annulotrematoides amazonicus,
a new genus and species (Dactylogyridae: Ancyrocephalinae), from the gills of Psectrogaster
rutiloides (KNER) (Teleostei: Characiformes: Curimatidae) from the Brazilian Amazon. - Proc.
Biol. Soc. Wash. 108: 528-532.
KRITSKY, D.C. & W.A. BOEGER (1998): Neotropical Monogenoidea. 35. Pavanelliella pavanelli, a new
genus and species (Dactylogyridae: Ancyrocephalinae) from the nasal cavities of siluriform
fishes in Brazil. - J. Helminthol. Soc. Wash. 65: 160-163.
KRITSKY, D.C. & W.A. BOEGER (2002): Neotropical Monogenoidea. 41. New and previously
described species of Dactylogyridae (Platyhelminthes) from the gills of marine and
freshwater perciform fishes (Teleostei) with proposal of a new genus and a hypothesis on
phylogeny. - Zoosystema 24(1): 7-40.
KRITSKY, D.C. & T.H. FRITTS (1970): Monogenetic trematodes from Costa Rica, with the
proposal of Anacanthocotyle gen. n. (Gyrodactylidae: Isancistrinae). - Proc. Helminthol.
Soc. Wash. 37: 63-68.
KRITSKY, D.C. & P.A. GUTIÉRREZ (1998): Neotropical Monogenoidea. 34. Species of Demidospermus
(Dactylogyridae: Ancyrocephalinae) from the gills of pimelodids (Teleostei, Siluriformes) in
Argentina. - J. Helminthol. Soc. Wash. 65: 147-159.
KRITSKY, D.C. & P.D. LEIBY (1972): Dactylogyridae (Monogenea) from the freshwater fish, Astyanax
fasciatus (CUVIER), in Costa Rica, with descriptions of Jainus hexops sp. n. Urocleidoides costaricensis
and U. heteroancistrium combs. n. - Proc. Helminthol. Soc. Wash. 39: 227-230.
KRITSKY, D.C. & V.E. THATCHER (1974a): Monogenetic trematodes (Monophisthocotylea: Dactylo-
gyridae) from freshwater fishes of Colombia, South America. - J. Helminthol. 48: 59-66.
KRITSKY, D.C. & V.E. THATCHER (1974b): New monogenetic trematodes from freshwater fishes of
Western Colombia with the proposal of Anacanthoroides gen. n. (Dactylogyridae). - Proc.
Helminthol. Soc. Wash. 43: 129-134.
KRITSKY, D.C. & V.E. THATCHER (1976): New monogenetic trematodes from freshwater fishes of
Western Colombia with the proposal of Anacanthoroides gen. n. (Dactylogyridae). - Proc.
KRITSKY, D.C. & V.E. THATCHER (1977): Phanerothecium gen. nov. and Fundulotrema gen. nov.; two new
genera of viviparous Monogenoidea (Gyrodactylidae) with a description of P. caballeroi sp.
nov. and a key to the subfamilies and genera of the family. - Instituto Biológico de México,
Publ. Espec. 4: 53-60.
KRITSKY, D.C. & V.E. THATCHER (1983): Neotropical Monogenea. 5. Five new species from the aruanã,
Osteoglossum bicirrhosum VANDELLI, a freshwater teleost from Brazil, with the proposal of
Gonocleithrum n. gen. (Dactylogyridae: Ancyrocephalinae). - Proc. Biol. Soc. Wash. 96: 581-597.
KRITSKY, D.C. & V.E. THATCHER (1984): Neotropical Monogenea. 6. Five species of Diplectanum
(Diplectanidae) from freshwater teleosts, Plagioscion spp. (Scianidae), in Brazil. - Proc. Biol.
Soc. Wash. 97: 434-443.
KRITSKY, D.C., BOEGER, W.A. & M. JÉGU (1996 a): Neotropical Monogenoidea. 28. Ancyrocephali-
nae (Dactylogyridae) of piranha and their relatives (Teleostei, Serrasalmidae) from Brazil
and French Guiana: species of Notozothecium BOEGER and KRITSKY, 1988, and Mymarothecium
gen. n. - J. Helminthol. Soc. Wash. 63: 153-175.
KRITSKY, D.C., BOEGER, W.A. & M. JÉGU (1997a): Neotropical Monogenoidea. 29. Ancyrocephalinae
(Dactylogyridae) of piranha and their relatives (Teleostei, Serrasalmidae) from Brazil: species
of Amphithecium BOEGER and KRITSKY, 1988, Heterothecium gen. n. and Pithanothecium gen. n. - J.
KRITSKY, D.C., BOEGER, W.A. & M. JÉGU (1997b): Neotropical Monogenoidea. 30. Ancyrocephali-
nae (Dactylogyridae) of piranha and their relatives (Teleostei, Serrasalmidae) from Brazil:
species of Calpidothecium gen. n., Calpidothecioides gen. n., Odothecium gen. n., and Notothecioides
gen. n. - J. Helminthol. Soc. Wash. 64: 208-218.
KRITSKY, D.C., BOEGER, W.A. & M. JÉGU (1998 ): Neotropical Monogenoidea. 31. Ancyrocephalinae
(Dactylogyridae) of piranha and their relatives (Teleostei, Serrasalmidae) from Brazil: species of
Notothecium BOEGER & KRITSKY, 1988, and Enallothecium gen. n. - J. Helminthol. Soc. Wash. 65: 31-49.
KRITSKY, D.C., BOEGER, W.A. & F. POPAZOGLO (1995): Neotropical Monogenoidea. 22. Variation in
Scleroductus species (Gyrodactylidea, Gyrodactylidae) from siluriform fishes of Southeastern
Brazil. - J. Helminthol. Soc. Wash. 62: 53-56.
KRITSKY, D.C., BOEGER, W.A. & V.E. THATCHER (1985): Neotropical Monogenea. 7. Parasites of the
pirarucu, Arapaima gigas (CUVIER), with description of two new species and redescription of
Dawestrema cycloancistrium PRICE & NOWLIN, 1967 (Dactylogyridae, Ancyrocephalinae). - Proc.
Biol. Soc. Wash. 97: 321-331.
KRITSKY, D.C., BOEGER, W.A. & V.E. THATCHER (1986): Neotropical Monogenea. 9. Status of Trinigyrus
HANEK, MOLNAR & FERNANDO, 1974 (Dactylogyridae) with descriptions of two new species
from loricariid catfishes from the Brazilian Amazon. - Proc. Biol. Soc. Wash. 99(3): 392-398.
KRITSKY, D.C., BOEGER, W.A. & V.E. THATCHER (1988): Neotropical Monogenea. 11. Rhinoxenus,
new genus (Dactylogyridae: Ancyrocephalinae) with descriptions of three new species from
the nasal cavities of Amazonian Characoidea. - Proc. Biol. Soc. Wash. 101(1): 87-94.
KRISKY, D.C., BOEGER, W.A. & L.R. VAN EVERY (1992): Neotropical Monogenoidea. 17. Anacantho-
rus MIZELLE and PRICE (Dactylogyridae, Anacanthorinae) from characoid fishes of Central
Amazon. - J. Helminthol. Soc. Wash. 59: 25-51.
KRITSKY, D.C., LEIBY, P.B. & R.J. KAYTON (1978): A rapid stain technique for the haptoral bars of
Gyrodactylus species (Monogenea). - J. Parasitol. 64(1): 172-174.
KRITSKY, D.C., MENDOZA-FRANCO , E.F. & T. SCHOLZ (2000): Neotropical Monogenoidea. 36.
Dactylogyrids from the gills of Rhamdia guatemalensis (Siluriformes: Pimelodidae) from
cenotes of the Yucatan Peninsula, Mexico, with proposal of Ameloblastella gen. n. and
Aphanoblastella gen. n. (Dactylogyridae: Ancyrocephalinae). - Comparative Parasitology
67: 76-84.
KRITSKY, D.C., THATCHER, V.E. & W.A. BOEGER (1986): Neotropical Monogenea. 8. Revision of
Urocleidoides (Dactylogyridae, Ancyrocephalinae). - Proc. Helminthol. Soc. Wash. 53: 1-37.
KRITSKY, D.C., THATCHER, V.E. & W.A. BOEGER (1987): Neotropical Monogenea. 10. Omothecium,
new genus (Dactylogyridae: Ancyrocephalinae) and two new species from the piranambu,
Pinirampus pirinampu (SPIX), (Siluriformes), in Brazil. - Proc. Biol. Soc. Wash. 100: 8-12.
KRITSKY, D.C., THATCHER, V.E. & W.A. BOEGER (1988): Neotropical Monogenea. 13. Rhinonastes
pseudocapsaloideum n. gen., n. sp. (Dactylogyridae, Ancyrocephalinae), a nasal parasite of
curimata, Prochilodus nigricans AGASSIZ (Cypriniformes, Prochilodontidae), in Brazil. - J.
Parasitol. 74: 695-698.
KRITSKY, D.C., THATCHER, V.E. & W.A. BOEGER (1989): Neotropical Monogenea. 15. Dactylogyridae
from the gills of Brazilian Cichlidae with proposal of Sciadicleithrum gen. n. (Dactylogyridae).
- Proc. Helminthol. Soc. Wash. 56: 128-140.
KRITSKY, D.C., THATCHER, V.E. & R.J. KAYTON (1979): Neotropical Monogenoidea. 2. The
Anacanthorinae PRICE, 1967, with the proposal of four new species of Anacanthorus MIZELLE
& PRICE, 1965, from Amazonian fishes. - Acta Amazonica 9: 355-361.
KRITSKY, D.C., THATCHER, V.E. & R.J. KAYTON (1980): Neotropical Monogenoidea. 3. Five new
species from South America with the proposal of Tereancistrum gen. n. and Trinibaculum gen. n.
(Dactylogyridae: Ancyrocephalinae). - Acta Amazonica 10: 411-417.
KRITSKY, D.C., VAN EVERY, L.R. & W.A. BOEGER (1996 b): Neotropical Monogenoidea. 27. Two new
species of Telethecium gen. n. from the nasal cavities of Central Amazonian fishes and a
redescription of Kritskyia moraveci KOHN, 1990 (Dactylogyridae, Ancyrocephalinae). - J.
KRITSKY, D.C., VIDAL-MARTINEZ, V.M. & R. RODRIGUEZ-CANUL (1994): Neotropical Monogenoidea.
19. Dactylogyridae of cichlids (Perciformes) from the Yucatán Peninsula, with descriptions
of three new species of Sciadicleithrum KRITSKY, THATCHER & BOEGER, 1989. - J. Helminthol.
Soc. Wash. 61: 26-33.
MALABARBA, L.R. & I.M.L. ROSA (2003): Preface. - Neotropical Ichthyology 1: 1.
MAYES, M.A., BROOKS, D.R. & T.B. THORSON (1981): Potamotrygonocotyle tsalickisi, new genus and
species (Monogenea: Monocotylidae) and Paraheteronchocotyle amazonensis, new genus and
species (Monogenea: Hexabothriidae) from Potamotrygon circularis GARMAN (Chondrichthyes:
Potamotrygonidae) in Northwestern Brazil. - Proc. Biol. Soc. Wash. 94: 1205-1210.
MENDOZA-FRANCO, E.F. & V.M. VIDAL-MARTINEZ (2001): Salsuginus neotropicalis n. sp. (Monogenea:
Ancyrocephalinae) from the pike killifish Belonesox belizanus (Atheriniformes: Poeciliidae) from
Southeastern Mexico. - Systematic Parasitology 48: 41-45.
MENDOZA-FRANCO, E.F., SCHOLZ, T. & V.M. VIDAL-MARTÍNEZ (1997): Sciadicleithrum meeki sp. n.
(Monogenea: Ancyrocephalinae) from the gills of Cichlasoma meeki (Pisces: Cichlidae) from
cenotes (= sinkholes) of the Yucatan Peninsula, México. - Folia Parasitológica 44: 205-208.
MIZELLE, J.D. (1936): New species of trematodes from the gills of Illinois fishes. American Midland
Naturalist 17: 785-806.
MIZELLE, J.D. & D.C. KRITSKY (1967): Unilatus gen. n., a unique Neotropical genus of Monogenea.
- J. Parasitol. 53: 1113-1114.
MIZELLE, J.D. & D.C. KRITSKY (1969a): Studies on monogenetic trematodes. XXXIX. Exotic species
of Monophisthocotylea with the proposal of Archidiplectanum gen. n. and Longihaptor gen. n. -
American Midland Naturalist 81: 370-386.
MIZELLE, J.D. & D.C. KRITSKY (1969b): Studies on monogenetic trematodes. XL. New species from
marine and freshwater fishes. - American Midland Naturalist 82: 417-428.
MIZELLE, J.D., KRITSKY, D.C. & J.W. CRANE (1968): Studies on monogenetic trematodes. XXXVIII.
Ancyrocephalinae from South America with the proposal of Jainus gen. n. - American
Midland Naturalist 80: 186-198.
MIZELLE, J.D. & C.E. PRICE (1964): Studies on monogenetic trematodes. XXVII. Dactylogyrid
species with the proposal of Urocleidoides gen. n. - J. Parasitol. 50: 579-584.
MIZELLE, J.D. & C.E. PRICE (1965). Studies on monogenetic trematodes. XXVII. Gill parasites of
the piranha with the proposal of Anacanthorus gen. n. - J. Parasitol. 51: 30-36.
MOLNAR, K., HANEK, G. & C.H. FERNANDO (1974): Ancyrocephalids (Monogenea) from freshwater
fishes of Trinidad. - J. Parasitol. 60: 914-920.
NASIR, P. (1983): Occurrence and significance of the monogenean Cycloplectanum americanum (PRICE,
1937) OLIVER, 1968, on a freshwater host. - J. Parasitol. 69: 957-962.
POPAZOGLO, F. & W.A. BOEGER (2000): Neotropical Monogenoidea. 37. Redescription of Gyrodacty-
lus superbus (SZIDAT, 1973) comb. n. and description of two new species of Gyrodactylus
(Gyrodactylidea: Gyrodactylidae) from Corydoras paleatus and C. ehrhardti (Teleostei: Siluri-
formes: Callichthyidae) of Southern Brazil. - Folia Parasitologica 47: 105-110.
PRICE, C.E. (1966): Urocleidus cavanaughi, a new monogenetic trematode from the gills of the
keyhole cichlid, Aequidens maroni (STEINDACHNER). - Bulletin of the Georgia Academy of
Science 24: 117-120.
PRICE, C.E. (1968): Diaccessorius, a new genus of Monogenea from the gills of an Amazon river
teleost. - Acta Biológica Venezuelica 6: 84-89.
PRICE, C.E. & W.A. BUSSING (1967): Monogenean parasites of Costa Rican fishes. Part I. Descriptions
of two new species of Cleidodiscus MUELLER, 1934. - Revista Parasitológica 28: 81-86.
PRICE, C.E. & W.A. BUSSING (1968): Monogenean parasites of Costa Rican fishes. Part II. Proposal
of Palombitrema heteroancistrium n. gen., n. sp. - Proc. Helminthol. Soc. Wash. 35: 54-57.
PRICE, C.E. & T.E. MCMAHON (1966): Monocleithrium, a new genus of Monogenea from an Amazon
River teleost. - Revista Parasitológica 27: 221-226.
PRICE, C.E. & W.J. NOWLIN (1967): Proposal of Dawestrema cycloancistrium n. gen., n.sp. (Trematoda:
Monogenea) from an Amazon River host. - Revista Parasitológica 28: 1-9.
PRICE, C.E. & N.G. ROMERO (1969): First account of a monogenetic trematode from Paraguay:
Amphocleithrium paraguayensis n. gen., n. sp. - Zoologische Jahrbücher 96: 449-452.
PRICE, C.E. & E.A. SCHLUETER (1967): Two new monogenetic trematodes from South America. - J.
Tennessee Acad. Sci. 42: 23-25.
PRICE, E.W. (1938): The monogenetic trematodes of Latin America. - Livro Jubilar do Prof. LAURO
TRAVASSOS, Rio de Janeiro (Brazil) 3: 407-413.
SURIANO, D.M. (1980): Notodiplocerus singularis gen. et sp. nov. (Monogenea: Ancyrocephalinae)
parásito de las branquias de Pseudocurimata gilberti (Pisces: Tetragonopteridae) de la Laguna de
Chascomus, República Argentina. - Neotropica 26: 131-143.
SURIANO, D.M. (1981): Andospira n. g., n. sp. (Monogenea, Ancyrocephalinae) a branchial parasite of
Pseudocurimata gilberti from Lake Chascomus, Argentina. - Neotropica 27: 67-78.
SURIANO, D.M. (1983): Demidospermus anus gen. nov. sp. nov. (Monogenea: Ancyrocephalinae)
parasita branquial de Loricaria (L) anus VALENCIENNES, 1840 (Pisces: Loricariidae) de la Laguna
de Chascomus, Provincia de Buenos Aires, Republica Argentina. - Neotropica 29: 169-172.
SURIANO, D.M. (1985): El género Unilatus MIZELLE y KRITSKY, 1967 (Monogenea: Ancyrocephal-
idae) parásito de Siluriformes (Pisces: Loricariidae) del Rio Negro, Manaus, Brasil. -
Neotropica 31: 163-175.
SURIANO, D.M. (1986a): Philocorydoras platensis gen. n. et. sp. n. (Monogenea: Ancyrocephalidae) from
Corydoras paleatus (Jenyns) (Pisces: Callichthyidae) in Laguna Chascomús - República Argenti-
na. - Helminthologia 23: 249-256.
SURIANO, D.M. (1986b): The genus Urocleidoides MIZELLE and PRICE, 1964 (Monogenea: Ancyroceph-
alidae). Anatomy and systematic position. Urocleidoides mastigatus sp. n. and U. travassosi (PRICE,
1934) MOLNAR, HANEK and FERNANDO, 1974 from Rhamdia sapo (VALENCIENNES, 1840)
EIGENMANN, 1888 and Pimelodella laticeps EIGENMANN, 1917 (Pisces: Siluriformes) in Laguna
Chascomus, República Argentina. - Physis (Buenos Aires), Secc. B, 44: 73-80.
SURIANO, D.M. & I.S. INCORVAIA (1995): Ancyrocephalid (Monogenea) parasites from siluriform
fishes from the paranean-platean ichthyogeographical province in Argentina. - Acta Parasito-
logica 40: 113-124.
SZIDAT, L. (1973): Morphologie und Verhalten von Paragyrodactylus superbus n. g., n. sp., Erreger eines
Fischsterben in Argentinien. - Angew. Parasit. 14: 1-10.
TAKEMOTO, R.M., LIZAURA, R. & G.C. PAVANELLI (2002): A new species of Kritskyia (Dactylogyridae,
Ancyrocephalinae) parasite of urinary bladder of Prochilodus lineatus (Prochilodontidae,
Characiformes) from the floodplain of the high Paraná river, Brazil. - Mem. Inst. Oswaldo
Cruz 97(3): 313-315.
THATCHER, V.E. & D.C. KRITSKY (1983): Neotropical Monogenoidea. 4. Linguadactyloides brinkmanni
gen. et sp. n. (Dactylogyridae: Linguadactyloidinae subfa. nov.) with observations on its
pathology in a Brazilian freshwater fish, Colossoma macropomum (CUVIER). - Proc. Helminthol.
Soc. Wash. 50: 305-311.
TURNBULL, E.R. (1956): Gyrodactylus bullatarudis n. sp. from Lebistes reticulatus PETERS with a study of
its life cycle. - Canadian Journal of Zoology 34: 583-594.
VAN EVERY, L.R. & D.C. KRITSKY (1992): Neotropical Monogenoidea. 18. Anacanthorus MIZELLE and
PRICE 1965 (Dactylogyridae, Anacanthorinae) of piranha (Characoidea, Serrasalmidae) from
the central Amazon, their phylogeny, and aspects of host-parasite coevolution. - J. Helm-
inthol. Soc. Wash. 59: 52-75.
VIDAL-MARTÍNEZ, V.M., SCHOLZ, T. & M.L. AGUIRRE-MACEDO (2001): Dactylogyridae of cichlid
fishes from Nicaragua, Central America, with descriptions of Gussevia herotilapiae sp. n. and
three new species of Sciadicleithrum (Monogenea: Ancyrocephalinae). - Comparative Parasitol-
ogy 68(1): 76-86.
4
TREMATODA (DIGENEA)
The digenetic trematodes are endoparasitic platyhelminths having complex life-cycles. All
are hermaphroditic with the exception of some blood inhabiting forms (Schistosoma-
tidae) and some tissue invaders found in marine fishes (Didymozoidae). All trematode
species reported so far from Neotropical freshwater fishes have functional male and
female organ systems in each individual. Many fish trematodes are less than one
millimeter in length, but most of those treated here would fall into the 2 to 20 millimeter
size range. Although considered to be “flatworms”, not all species are dorso-ventrally
flattened. Some are indeed flat and leaf-like but others are cylindrical, spherical or
pyriform. The body is usually provided with oral and ventral suckers and the external
surface (tegument) may be spined.
The female reproductive system consists of: an ovary; oviduct; yolk glands (vitellar-
ia); oötype; Mehlis’s gland; Laurer’s canal and seminal receptacle (sometimes absent).
Some species also have the distal end of the uterus modified into a muscular tube for
reception of the male copulatory organ (cirrus). This functional vagina in trematodes is
called a metraterm.
The male system includes: testes (usually two but there may be from one to many
testes present according to the species); vas deferens; seminal vesicle and a cirrus and
cirrus sac (which are sometimes absent). The number, size, form and position of the
reproductive organs are important in the classification and identification of these worms.
The nature of the digestive system is also important in the systematics of trematodes.
This system usually consists of: a mouth and oral sucker; prepharynx; pharynx; esophagus
and two intestinal branches called ceca or crura. The digestive system is normally incomplete
in that the intestinal branches end blindly. There are some species, however, in which the
crura open to the exterior through anal pores or open into the excretory bladder.
Spermatozoa produced in the testes reach the seminal vesicle by way of tubules (vasa
efferentia and vas deferens). As in the case in most hermaphroditic invertebrates,
copulation consists of a mutual exchange of sperm cells with each individual inserting the
male organ (cirrus) into the female pore (uterine pore or metraterm) of the other. The
male cells make their way the length of the uterus and are stored in the seminal receptacle,
to be used as needed.
Female cells from the ovary pass one at a time down the oviduct to a muscular
organ called the oötype. There the ovum is fertilized by a sperm from the seminal
receptacle and surrounded by yolk cells from the vitelline glands. Shell material is
provided by the yolk glands as well in the form of a liquid which quickly hardens
around the gamete and yolk cells. Mehlis’ gland, which surrounds the oötype, is thought
to produce a catalytic agent to harden the shell. The finished capsule (egg) is slowly
carried down the length of the uterus and eventually expelled into the environment
through the uterine pore (common genital pore).
Adult trematodes live in the digestive tract, hollow organs, circulatory system or
subcutaneous connective tissue of vertebrates. Copulation and egg production
(sexual reproduction) is carried out in these hosts. The eggs are carried to the exterior
environment with the feces or urine of the host and they usually hatch shortly after
reaching water. Some fish trematodes, especially Paramphistomidae, are ovovivipa-
rous and in these forms, the egg shell is reabsorbed in the uterus and larvae are
liberated from the uterine pore. The first larval stage (miracidium) is ciliated and
swims actively in search of an appropriate species of snail. These larvae cannot feed
so they die in a few hours if they do not reach a suitable host. Upon finding a snail,
the miracidium penetrates the body wall by means of its anterior penetration glands
and makes its way to the digestive gland (hepatopancreas). There it loses its cilia, and
most internal structure and assumes a sac-like form (sporocyst). Within the sporocyst,
the third larval stages (rediae) are produced from germinative cells. Several of these
larvae are produced within the sporocyst which eventually ruptures to free them
within the snail’s digestive gland. The rediae in turn produce larvae called cercariae.
These leave the rediae through birth pores and abandon the snail to seek a second
intermediary host, or to encyst on vegetation. Cercariae do not feed although they
normally have a mouth and digestive system. They have enough stored energy to
swim actively for several hours after which they die if they do not find a suitable host
or a site for encystment. Most cercariae have a posterior tail-like projection to
facilitate swimming. During encystment, the tail is lost and the resultant infective
larva is termed a metacercaria. In some cases cercariae may invade the definitive host
directly. This is the normal means of transmission of blood inhabiting species
(Sanguinicolidae) and of tissue forms (Didymozoidae).
Although little work has been done on the life-cycles of Neotropical fish trematodes,
it is possible to deduce what the mode of transmission must be based on studies done
elsewhere. With the exception of the two families mentioned above, all fish trematodes
enter the host as encysted metacercariae in the food. The families Acanthostomidae,
Allocreadiidae, Fellodistomidae, Heterophyidae and Opisthorchiidae are transmitted to
piscivorous species in the tissues of smaller fishes. Angiodictyidae, Haploporidae,
Paramphistomidae and Zonocotylidae reach their fish hosts in aquatic plants. Echinosto-
matidae are probably transmitted to fish that devour the snail intermediate host, and
Bucephalidae to those that eat bivalve mollusks.
III. Pathology
The pathogenicity of trematodes to their fish hosts varies in accordance with a number of
factors which can be summarized as follows:
1) A trematode that penetrates tissues will be more pathogenic than one that lives in
the intestinal lumen or the interiors of other hollow organs.
2) A trematode that is large in relation to the host or organ invaded will be more
pathogenic than a small trematode would be in the same host or organ.
3) Pathology in the host is usually proportional to the number of trematodes present
(intensity of infection).
4) Trematodes that migrate within the host cause more damage than those remain in
one location.
5) Trematode eggs passing through tissues (Sanguinicolidae) can cause more host
reaction that the adults producing the eggs.
6) Host reaction to tissue invading trematodes is initially an inflammatory one, with
infiltration of lymphocytes, and is followed by the fibrous encapsulation of the
invader. Once the encapsulation process is finished, danger to the host ends.
7) Host reaction to trematodes in the intestinal lumen is also of the inflammatory
type, but is generally limited to the point of fixation of the worms.
8) Trematodes within tubules (such as the bile and pancreatic ducts) can cause
distension of the lumen and hyperplasia of the walls and decrease their flow.
Trematode cercariae that invade the skin of fishes and become encysted there can
become visible to the naked eye if the fish host concentrates pigment cells around them.
This condition is wide-spread and is known as “black-spot disease”. In Amazonian fish,
we have observed this phenomenon in Hoplias malabaricus and Semaprochilodus insignis and it
no doubt occurs in other species as well. Although the presence of these metacercariae in
the skin does not seem to impair the health of the fish, the black spots are sometime so
numerous that they make the fish unappealing to the consumer. In the Neotropics, the
black spots are usually caused by Acanthostomidae or Heterophyidae which mature in
crocodilians, piscivorous birds or predatory fish. A somewhat similar condition is known
internationally as “yellow-spot disease”. The yellow spots are caused by the yellow
coloration of the metarcercariae of the family Clinostomidae. In the Amazon, we have
found the genera Clinostomum and Odhneriotrema to be responsible. The former matures in
birds and the latter in crocodilians.
Another sort of “yellow-spot disease” is found in the Amazonian fish, Plagioscion
squamosissimus, which is a freshwater relative of the Atlantic weakfish. This condition is
caused by adults of Brasicystis bennetti THATCHER, 1979, of the family Didymozoidae (Figs.
4-3 & 4-47). These worms live in the subcutaneous tissues with their slender anterior ends
projecting through small holes that they maintain open. As they mature, their entire
bodies become packed with thousands of small egg capsules which have a yellowish hue.
These masses of eggs can be seen through the skin as yellowish spots or blotches which
are as much as two centimeters in diameter. The most common site for penetration is the
inner surface of the gill covers, but the worms can also be found in the gill filaments, fins,
eyes and generally distributed throughout the skin of the body. Some fish become so
heavily infected as to have an overall yellowish appearance. Local fishermen in the

Brazilian Amazon have a special name for such fish and many regard them as a different
species. In some parts of the Amazon, prevalence rates reach 10 % but the worms do not
seem to inconvenience the fish.
Another world wide problem in fish caused by larval trematodes is known as
“eye fluke disease”. Infected fish can be seen to have larvae moving about within the
eye. There is usually no inflammatory reaction, but the presence of the worms
impedes the vision of the fish. These larvae are less well developed than metacercar-
iae and do not become encysted. They are known as mesocercariae or diplostomula
and usually belong to the genera Diplostomum or Alaria of the superfamily Strigeoidea.
Diplostomula are much flattened and have the general appearance of the sole of a
tennis shoe (Figs. 4-48 & 4-49).
Mesocercariae of species that infect fishes can be found in any organ, but they tend
to concentrate in the eyes and optic nerves. In these sites, they can blind the fish and make
it easy prey for piscivorous birds. Thus, the parasite actively participates in the completion
of its life-cycle and the passage from the second intermediate to the definitive host. Such
larvae have been reported from the human eye (ASHTON et al. 1969). In human beings,
these migrating larvae can cause extensive tissue destruction, anaphylactic shock and
death if present in large numbers (FREEMAN et al. 1976).
THATCHER & VARELLA (1980) found that larval trematodes were involved with
branchial carcinoma in an Amazonian fish, Chaetobranchus semifasciatus. In this fish a large
primary neoplasm (27 x 18 x 12 mm) and a small satellite tumor (3 mm in diameter) were
found attached to the first branchial arch (Fig. 4-51). These were both dense tumors
covered by a smooth epithelium and containing blood vessels, collagen fibers, pigment
cells and irregular masses of cartilage. In these two tumors, there were no trematode
larvae, but in each of 19 additional tumorous growths in the gill filaments there was a
single metacercaria of Ascocotyle sp. (Figs. 4-52 & 4-53). In four of these tumoroids, the gill
filaments showed repetitious growth with parts of filaments projecting in varios
directions (Fig. 4-54).
Prevention of trematode infection in fish consists of eliminating intermediate hosts in the

environment. This is easy enough to do in aquaria and small intensive pisciculture
operations but frequently proves to be impossible in larger tanks and ponds. Introducing
snails and aquatic plants collected in nature to aquaria with tropical fish can be hazardous
and should be avoided. For a balanced aquarium, snails and plants grown in aquaria
should be utilized.
If no infected snails are present in an aquatic system, black-spot and yellow-spot
diseases can not occur. Of course, healthy snails can become infected at any time that
vertebrate feces containing trematode eggs enter the system. For example, in outdoor
ponds birds may defecate into the water and initiate an infective cycle. In an aquarium, a
fish carrying adult trematodes can potentially infect all the others if snails are present.
The feeding of wild caught minnows to captive predatory fish should be avoided
since such small fish frequently contain encysted metacercariae. If pieces of fish or
crustaceans are used as food in aquaria or ponds, such material should be deep-frozen for
at least 48 hours before use to kill any metacercariae present.
No treatment is possible for encysted metacercariae but adult trematodes can be
removed from the intestinal tracts of fish with Di-N-Butyl Tin Oxide which can be mixed
with the ration at 0.3 % of body weight administered for from one to five days.
Trematodes are best collected alive from freshly killed fish. To do so, wild caught fish can
be brought in alive and maintained in aquaria until used. If they are deprived of food for
10-24 hours prior to necropsy, their digestive tracts will be nearly clean and the trematodes
will be easier to see. This is especially important for bottom feeders that ingest sand since
some trematodes are about the same size as sand grains. If it is not possible to transport
the fish alive, they may be brought in on ice and kept in the refrigerator. Trematodes will
often remain alive for several hours in cool fish. The hosts should not be deep-frozen,
however, as this will not only kill the worms but damage them as well. Living trematodes
are easier to find than dead ones for the simple reason that they move.
Since many fish trematodes are less than one millimeter in length, some skill and care
are required to find them in the host viscera. All hollow organs (stomach, intestine, swim
bladder, heart, etc.) should be opened with scissors and washed in finger bowls by
enthusiastic agitation. After removing the washed organs from the bowls, the dirty water
should be slowly decanted and gradually replaced by clean water. This process of hand
sedimentation should be repeated until the water in the bowl is clear and the floating
debris has been eliminated. In this way, it is possible to concentrate the live worms in the
bottom of the dish where they can be found by careful examination with a dissecting
microscope.
Trematodes should be killed in a relaxed position before fixation and this can best be
accomplished by the application of gentle heat. A rapid method is to place one, or several
specimens, on a slide in water with a coverglass in place and pass a lighted match several
times underneath. When the worms no longer show movement, they can be fixed in place
with AFA fixative (85 parts of 85 % alcohol: 10 parts of commercial formaldehyde: 5
parts of glacial acetic acid). To do this, it is necessary to place drops of AFA on one side
of the coverglass while withdrawing the water from the other side with filter paper. If
large numbers of trematodes must be fixed in a limited time, they can be placed in water
and left in an embedding oven for 10-15 minutes. After they no longer move, they can be
transferred directly to AFA or fixed on slides with a little coverglass pressure.
After fixation in AFA for several hours, permanent preparations for study should be
prepared by means of the following procedure:
1. Wash the specimens in running water for 30 minutes to remove the fixative.
2. Place the worms in a staining solution containing 3 drops of Mayer’s Carmalum
stock solution to each 10 cm3 of water. Leave them in this stain until they are wine
colored throughout (the time varies from 10 minutes to 10 hours depending on

the size of the worms).
3. Pass the worms to a destaining solution (35 % alcohol to which hydrochloric acid
has been added in the ratio of 6 drops per liter). Leave them in destain until the
bodies are seen as pale pink and the organs are still wine or red (the time will vary
from a few seconds to several hours depending on size).
4. After destaining, dehydrate the worms gradually by serial passage through alcohol
at 50, 70, 85, 95 and 100 %. They should be left for a minimum of 10 minutes in
each alcohol. It is advisable to leave them for 30 minutes or more in the absolute
alcohol to insure complete dehydration.
5. After dehydration, the worms should be passed to a clearing agent such as methyl
salicylate or xylol.
6. The worms should clear completely in a few minutes and they can then be
mounted directly in Canada balsam, Clarite, Piccolyte or other similar mounting
media.
Mayer’s Carmalum stock solution can be mixed in the following way:

Mix 1 g of carminic acid stain, 10 g of alum and 200 cm3 of distilled water in a
beaker and boil for 30 minutes. Afterwards, filter the stain and store in the
refrigerator.
If carminic acid stain is unavailable, it can be made in the laboratory as follows:
1. Mix 10 g of carmine stain with 100 cm3 of glacial acetic acid in a beaker.
2. Heat the beaker to the boiling point and then let it cool.
3. Filter the liquid and save the material that sticks to the filter paper.
4. Scrape the material off of the filter paper and heat it in a beaker to dry it. This will
evaporate the excess acetic acid. The material should be warmed carefully while
stirring. This drying will eventually produce a dry dark red powder without the
odor of acetic acid. This resultant powder is carminic acid stain and can be used
as above to make Mayer’s Carmalum.
Trematodes have few hard parts other than the circumoral spines found in certain genera.
Identification is therefore based mainly on the number, size, form and position of the
suckers, pharynx and internal organs. It is necessary to distinguish the ovary, testes,
viteline glands and cirrus sac and note their position in the body and in relation to each
other. One must also be able to tell the difference between a cirrus sac and a
hermaphroditic sac. In a well stained specimen, it will be possible to see the uterus
entering the proximal end of the hermaphroditic sac. There may be visible eggs within the
sac as well. If the structure is a cirrus sac, the uterus will be seen to have an opening to the
exterior in common with it. Figures 4-1 to 4-3 will serve as guides to the identification of
trematode organs.
Key to the Neotropical freshwater fish trematodes
I. Anterior attachment organ a rhynchus; mouth ventral; gut saccular .............................................................................................. Bucephalidae
A. Rhynchus oval, without projections ................................................................................................................................ Bellumcorpus (Fig. 4-4)
B. Rhynchus with cap, opening subterminal ................................................................................................................................... Pararhipidocotyle
C. Rhynchus plug or funnel-shaped.
1. Mouth in anterior third of body; gut long ............................................................................................................ Paurorhynchus (Fig. 4-5)
2. Mouth in middle third of body; gut short .............................................................................................................Prosorhynchus (Fig. 4-6)
D. Rhynchus sucker-like ................................................................................................................................ Glandulorhynchus (Figs. 4-91 & 4-92)
II. Anterior attachment organ a sucker (when present); mouth anterior, terminal or subterminal; gut divided into two crura.
A. Oral sucker absent; parasites of circulatory system .................. Sanguinicolidae ................................................... Sanguinicola (Fig. 4-32)
B. Oral sucker present; ventral sucker (acetabulum) absent.
1. Body divided into slender anterior and swollen posterior portions; parasites of subcutaneous tissues ..........................................
....................................................................................................... Didymozoidae .......................................... Brasicystis (Figs. 4-3 & 4-47)
2. Body undivided; parasites of digestive tract ...................................................................................................................... Angiodictyidae
ABLA Vol. 1 – Thatcher: Amazon Fish Parasites

a. Body without posterior projections; cirrus sac small ......................................................................................... Curumai (Fig. 4-44)
b. Body with posterior projections; cirrus sac absent.
1) Body with 4 posterior projections; esophageal bulb absent ................................................................... Denticauda (Fig. 4-45)
2) Body with 2 posterior projections; esophageal bulb present .......................................................... Pseudoparabaris (Fig. 4-46)
C. Both suckers present; acetabulum at or near posterior extremity ......................................................................................... Paramphistomata
1. Acetabulum subterminal, with or without horizontal ridges on floor.
a. Acetabulum subterminal, with ridges on floor.
1) One testis present ........................................................................................................................................................ Zonocotylidae
a) Vitelline glands compact, subspherical .................................................................................................. Zonocotyle (Fig. 4-41)
b) Vitelline glands compact, V-shaped .................................................................................................. Zonocotyloides (Fig. 4-42)
2) Two testes present .............................................. Cladorchiidae ................... Dadayius (Figs.4-28, 4-65 A-B, 4-73 to 4-76 & 4-84)
b. Acetabulum without ridges on floor.
1) One testis present ................................................................................................................................... Pseudodiplodiscus (Fig. 4-29)
2) Two testes present.
123
a) Oral diverticula absent; testes tandem..............................................................................................Travassosinia (Fig. 4-27)
124
b) Oral diverticula absent; testes parallel .......................................................................................... Micramphistoma (Fig. 4-95)
c) Oral diverticula small, within wall of oral sucker.
ABLA Vol. 1 – Thatcher: Parásitos de Peces Amazónicos

(1) Anterior extremity with collar-like expansion.
(a) Testes parallel ................................................................................................................................... Kalitrema (Fig. 4-39)
(b) Testes tandem, compact ..................................................................................................... Pronamphistoma (Fig. 4-96)
(c) Testes tandem, lobate ......................................................................................................... Anavilhanatrema (Fig. 4-97)
(2) Anterior extremity without collar-like expansion.
(a) Vitellline glands compact ....................................................................................................... Colocladorchis (Fig. 4-40)
(b) Vitelline glands follicular .........................................................................................Pseudocladorchis (Figs. 4-30, 4-62)
d) Oral diverticula large, external to wall of oral sucker.
1) Collar-like expansion present.
a) Testes diagonal, subspherical .................................................................................................. Curimatrema (Fig. 4-64)
b) Testes tandem, lobate ...................................................................................................... Gammamphistoma (Fig. 4-57)
2) Collar-like expansion absent.
a) Tegument papillate ventrally.
(1) Five pairs of muscular puckers present laterally ...... Annelamphistoma (Figs. 4-67 A-C, 4-77 to 4-79, 4-89)
(2) One pair of muscular puckers present near oral sucker. ...................................... Inpamphistoma (Figs. 4-66)
b) Tegument smooth.
(1) Testes parallel ................................................................................................................... Betamphistoma (Fig. 4-55)
(2) Testes tandem.
(a) Large genital sucker present ............................................................................... Pacudistoma (Fig. 4-60 A-C)
(b) Acetabulum with papillae .................................................................................... Myleustrema (Fig. 4-59 A-B)
(c) Body rim folded ventrally ........................................................................... Alphamphistoma (Fig. 4-61 A-B)
(d) Acetabulum with posterior notch on rim ................................................... Zetamphistoma (Fig. 4-63 A-B)
(e) Vitellaria extensive; testes large, lobate ............................................................... Deltamphistoma (Fig. 4-56)
(f) Vitellaria less extensive; testes small, spherical .............................................. Doradamphistoma (Fig. 4-99)
D. Both suckers present; acetabulum not near posterior extremity.
1. Nine testes present ........................................................................... Cryptogonimidae ......................................... Iheringtrema (Fig. 4-22)
2. One testis present; hermaphroditic bursa present ............................................................................................................... Haploporidae
a. Crura fused together forming single bi-lobed sac ............................................................................................................... Unicoelium
b. Crura short, not reaching level of testis.
1) Acetabulum pre-equatorial ........................................................................................................ Chalcinotrema (Figs. 4-26 & 4-33)
2) Acetabulum post-equatorial .................................................................................................................. Lecithobotrioides (Fig. 4-94)
c. Crura longer reaching to level of testis.
1) Vitelline follicles extensive ..................................................................................................................... Lecithobotrioides (Fig. 4-31)
2) Vitelline follicles of limited extent..
(a) Testis elongate ................................................................................................................................. Megacoelium (Fig. 4-24 A-B)
(b) Testis subspherical ...................................................................................................... Saccocoelioides (Figs. 4-34, 4-35 & 4-43)
d. Crura long, far surpassing testis posteriorly.
1) Hermaphroditic bursa small .................................................................................................................. Paralecithobotrys (Fig. 4-36)
2) Hermaphroditic bursa large ......................................................................................................................... Rondotrema (Fig. 4-93)
3. One testis present; cirrus sac present .................................................................................................................................... Monorchiidae
a. Cirrus sac nearly half as long as body .............................................................................. Genolopa magnacirrus (Figs. 4-70 & 4-72)

b. Cirrus sac small, barely surpassing acetabulum.
1) Testis near posterior extremity .......................................................................................... Sphericomonorchis spinulosus (Fig. 4-69)
2) Testis far from posterior extremity .......................................................................... Paraproctotrema delicata (Figs. 4-68 & 4-71)
4. Two testes present; cirrus sac present or absent.
a. Ovary posterior to testes.
1) Vitelline glands posterior to ovary.
a) Vitelline glands near posterior extremity ............................................................................................................. Halipegidae
(1) Vitelline glands of large follicles grouped together into two masses ........................................ Halipegus (Fig. 4-21)
(2) Vitelline follicles smaller, in two groups .......................................................................................................... Gonocercella
b) Vitelline glands anterior to acetabulum ......................................................................................................... Fellodistomidae
(1) Testes intercecal .................................................................................................................................... Antorchis (Fig. 4-13)
(2) Testes extracecal ....................................................................................................................................... Bacciger (Fig. 4-98)
b. Ovary between testes.
1) Mouth surrounded by papillae .......................................... Cryptogonimidae ............................. Palaeocryptogonimus (Fig. 4-23)
125
2) Mouth simple ........................................................................ Callodistomidae ......................................... Prosthenhystera (Fig. 4-7)
126
c. Ovary anterior to testes.
1) Oral sucker provided with spines.

a) Vitelline glands lateral to acetabulum ...................... Heterophyidae .................................................... Parspina (Fig. 4-20)
b) Vitelline glands posterior to acetabulum ................. Acanthostomidae ..................................... Acanthostomum (Fig. 4-1)
2) Oral sucker provided with spinous collar ....................... Echinostomatidae ................................... Caballerotrema (Fig. 4-16)
3) Oral sucker with membranous extensions ..................... Allocreadiidae.
a) Oral sucker with lobes ....................................................................................................................... Crepidostomum (Fig. 4-10)
b) Oral sucker with cap-like structure ..................................................................................................... Creptotrema (Fig. 4-11)
4) Oral sucker simple.
a) Vitelline glands consisting of two compact masses in mid-body region .................................................. Gorgoderidae
(1) Vitelline masses elongate ................................................................................................................. Dendrorchis (Fig. 4-19)
(2) Vitelline masses smaller than testes ........................................................................................ Amazonadistoma (Fig. 4-2)
(3) Vitelline masses larger than testes ......................................................................................... Phyllodistomoides (Fig. 4-25)
b) Vitelline glands consisting of follicles in limited fields posterior to acetabulum.
(1) Testes near posterior extremity; suckers subequal.
(a) Prepharynx long ............................................... Opisthorchiidae .............................................. Witenbergia (Fig. 4-8)
(b) Prepharynx short ............................................. Allocreadiidae ............................................ Procaudotestis (Fig. 4-14)
(2) Testes near posterior extremity; acetabulum much smaller than oral sucker ..............................................................
................................................................................... Opisthorchiidae ................................................ Cladocystis (Fig. 4-9)
(3) Testes in posterior half of body; acetabulum much larger than oral sucker; pharynx large, with finger-like
anterior extensions ................................................. Fellodistomidae ........................................... Kalipharynx (Fig. 4-15)
c) Vitelline glands consisting of follicles which are large and limited to two fields lateral to acetabulum ......................
.......................................................................................... Allocreadiidae ......................................... Magnivitellinum (Fig. 4-17)
d) Vitelline glands consisting of follicles that are small and extensive, especially in posterior half of body.
(1) Lateral vitelline fields uniting anteriorly and posteriorly to acetabulum ......................................................................
................................................................................... Allocreadiidae ................................................. Crassicutis (Fig. 4-18)
(2) Lateral vitelline fields not uniting anteriorly ...... Lepocreadidae ................................................ Eocreadium (Fig. 4-12)
Clave para identificación de Tremátodos, parásitos de peces Neotropicales
I. Organo de fijación anterior, rincodeo; boca ventral; intestino en forma de saco ..................................................................... Bucephalidae
A. Rincodeo oval, sin proyecciones .....................................................................................................................................Bellumcorpus (Fig. 4 -4)
B. Rincodeo con tapa, abertura subterminal .................................................................................................................................. Pararhipidocotyle
C. Rincodeo en forma de tapón o embudo.
1. Boca en el tercio anterior del cuerpo; intestino largo ......................................................................................... Paurorhynchus (Fig. 4-5)
2. Boca en el tercio central del cuerpo; intestino corto ........................................................................................... Prosorhynchus (Fig. 4-6)
D. Rincodeo en forma de ventosa .............................................................................................................. Glandulorhynchus (Figs. 4-91 & 4-92)
II. Organo de fijación anterior, una ventosa (cuando presente); boca anterior, terminal o subterminal; intestino dividido en 2 ramales
A. Ventosa oral ausente; parásitos del sistema circulatorio ................. Sanguinicolidae ............................................. Sanguinicola (Fig. 4-32)
B. Ventosa oral presente; ventosa ventral (acetábulo) ausente.
1. Cuerpo dividido en porción anterior delgada y porción posterior hinchada; parásitos de tejidos subcutáneos. ............................
.............................................................................................................. Didymozoidae ................................... Brasicystis (Figs. 4-3 & 4-47)
2. Cuerpo no dividido; parásitos del tracto digestivo ........................................................................................................... Angiodictyidae

a. Cuerpo sin proyecciones posteriores; saco del cirro pequeño ......................................................................... Curumai (Fig. 4-44)
b. Cuerpo con proyecciones posteriores; saco del cirro ausente.
1) Cuerpo con 4 proyecciones posteriores; bulbo esofágico ausente ....................................................... Denticauda (Fig. 4-45)
2) Cuerpo con 2 proyecciones posteriores; bulbo esofágico presente ............................................... Pseudoparabaris (Fig. 4-46)
C. Ambas ventosas presentes; acetábulo en o cerca de la extremidad posterior ............................................................... Paramphistomata
1. Acetábulo subterminal, con o sin surcos horizontales en el piso.
a. Acetábulo subterminal, con surcos en el piso.
1) Un teste presente ........................................................................................................................................................ Zonocotylidae
(a) Glándulas vitelinas compactas, subesféricas ......................................................................................... Zonocotyle (Fig. 4-41)
(b) Glándulas vitelinas compactas, forma de V ................................................................................... Zonocotyloides (Fig. 4-42)
2) Dos testes presentes ............................................... Orchiidae ...................... Dadayius (Figs. 4-28, 4-65 A-B, 4-73 a 4-76 & 4-84)
b. Acetábulo sin surcos en el piso.
1) Um teste presente ................................................................................................................................... Pseudodiplodiscus (Fig. 4-29)
2) Dos testes presentes.
127
(a) Divertículo oral ausentes; testes uno delante del otro .................................................................. Travassosinia (Fig. 4-27)
128
(b) Divertículo oral ausentes; testes paralelos ................................................................................... Micramphistoma (Fig. 4-95)
(c) Divertículo oral pequeño, dentro de la pared de la ventosa oral

(1) Extremidad anterior con expansión en forma de collar.
a) Testes paralelos ................................................................................................................................ Kalitrema (Fig. 4-39)
b) Testes uno delante del otro, compactos .......................................................................... Pronamphistoma (Fig. 4-96)
c) Testes uno delante del otro, lobulados ...........................................................................Anavilhanatrema (Fig. 4-97)
(2) Extremidad anterior sin expansión en forma de collar
a) Glándulas vitelinas compactas .............................................................................................. Colocladorchis (Fig. 4-40)
b) Glándulas vitelinas foliculares ................................................................................Pseudocladorchis (Figs. 4-30, 4-62)
(d) Divertículos orales grandes, externo a la pared de la ventosa oral.
(1) Expansión en forma de collar presente.
a) Testes diagonal, sub-esférico .................................................................................................. Curimatrema (Fig. 4-64)
b) Testes uno detrás del otro, lobulado ............................................................................ Gammamphistoma (Fig. 4-57)
(2) Expansión en forma de collar ausente.
a) Tegumento con papilas ventrales.
1. Cinco pares de pliegues musculares laterales ............ Annelamphistoma (Figs. 4-67 A-C, 4-77 a 4-79, 4-89)
2. Un par de pliegues musculares cerca de la ventosa oral ......................................... Inpamphistoma (Fig. 4-66)
b) Tegumento liso.
1. Testes paralelos ................................................................................................................ Betamphistoma (Fig. 4-55)
2. Testes uno detrás del otro.
(a) Ventosa genital grande presente ........................................................................ Pacudistoma (Fig. 4-60 A-C)
(b) Acetábulo con papilas .......................................................................................... Myleustrema (Fig. 4-59 A-B)
(c) Borde del cuerpo doblado ventralmente ................................................. Alphamphistoma (Fig. 4-61 A-B)
(d) Acetábulo con ranura posterior en el borde .............................................. Zetamphistoma (Fig. 4-63 A-B)
(e) Vitelaria extensa; testes grandes, lobulados ....................................................... Deltamphistoma (Fig. 4-56)
(f) Vitellaria menos extensa; testes pequeños, esféricos .................................... Doradamphistoma (Fig. 4-99)
D. Ambas ventosas presentes; acetábulo alejado de la extremidad posterior.
1. Nueve testes presentes ..................................................................... Cryptogonimidae ......................................... Iheringtrema (Fig. 4-22)
2. Un teste presente; bursa hermafrodita presente .................................................................................................................. Haploporidae
a. Ramales intestinales fusionados formando un único saco bilobado ................................................................................ Unicoelium
b. Ramos intestinales cortos, no alcanza el nivel de los testes.
(1) Acetábulo pre-ecuatorial ............................................................................................................ Chalcinotrema (Figs. 4-26 & 4-33)
(2) Acetábulo post-ecuatorial ...................................................................................................................... Lecithobotrioides (Fig. 4-94)
c. Ramos intestinales largos alcanzando el nivel de los testes.
(1) Folículos vitelinos extensos ................................................................................................................... Lecithobotrioides (Fig. 4-31)
(2) Folículos vitelinos con extensión limitada.
(a) Teste alargado .................................................................................................................................. Megacoelium (Fig. 4-24 A-B)
(b) Teste sub esférico ....................................................................................................... Saccocoelioides (Figs. 4-34, 4-35 & 4-43)
d. Ramos intestinales largos, pasando con largueza los testes posteriomente.
1) Bursa hermafrodita pequeña ................................................................................................................. Paralecithobotrys (Fig. 4-36)
2) Bursa hermafrodita grande ........................................................................................................................... Rondotrema (Fig. 4-93)
3. Un teste presente, saco del cirro presente ............................................................................................................................ Monorchiidae
a. Saco del cirro aproximadamente la mitad del largo del cuerpo .................................. Genolopa magnacirrus (Figs. 4-70 & 4-72)

b. Cirro pequeño, apenas pasando el acetábulo.
1) Teste cerca de la extremidad posterior ............................................................................ Sphericomonorchis spinulosus (Fig. 4-69)
2) Teste alejado de la extremidad posterior ................................................................ Paraproctotrema delicata (Figs. 4-68 & 4-71)
4. Dos testes presentes; saco del cirro presente o ausente.
a. Ovario posterior a los testes.
1) Glándulas vitelinas posteriores al ovario.
a) Glándulas vitelinas cerca de la extremidad posterior .................................................................................................. Halipegidae
(1) Glándulas vitelinas de grandes folículos agrupadas en dos masas ............................................ Halipegus (Fig. 4-21)
(2) Folículos vitelinos más pequeños, en dos grupos .......................................................................................... Gonocercella
b) Glándulas vitelinas anteriores al acetábulo ................................................................................................... Fellodistomidae
(1) Testes intercecales .................................................................................................................................. Antorchis (Fig.4-13)
(2) Testes extracecales ................................................................................................................................... Bacciger (Fig. 4-98)
b. Ovario entre los testes.
1) Boca rodeada por papilas ................................................... Cryptogonimidae ............................. Palaeocryptogonimus (Fig. 4-23)
129
2) Boca simple ........................................................................... Callodistomidae ......................................... Prosthenhystera (Fig. 4-7)
130
c. Ovario anterior a los testes.
1) Ventosa oral provista con espinas.

a) Glándulas vitelinas laterales al acetábulo ................ Heterophyidae .................................................... Parspina (Fig. 4-20)
b) Glándulas vitelinas posteriores al acetábulo ........... Acanthostomidae ..................................... Acanthostomum (Fig. 4-1)
2) Ventosa oral provista de collar espinoso ........................ Echinostomatidae ................................... Caballerotrema (Fig. 4-16)
3) Ventosa oral con extensiones membranosas ................. Allocreadiidae.
a) Ventosa oral con lóbulos .................................................................................................................. Crepidostomum (Fig. 4-10)
b) Ventosa oral con estructura parecida a gorra .................................................................................... Creptotrema (Fig. 4-11)
4) Ventosa oral simple.
a) Glándulas vitelinas consisten de dos masas compactas en región central ................................................ Gorgoderidae
(1) Masas vitelinas alongadas ................................................................................................................ Dendrorchis (Fig. 4-19)
(2) Masas vitellinas menores que los testes ................................................................................. Amazonadistoma (Fig. 4-2)
(3) Masas vitelinas mayores que los testes ................................................................................. Phyllodistomoides (Fig. 4-25)
b) Glándulas vitelinas consisten de folículos en áreas limitadas posteriores al acetábulo.
(1) Testes cerca de la extremidad posterior. Ventosas desiguales.
(a) Prefaringe larga ................................................ Opisthorchiidae .............................................. Witenbergia (Fig. 4-8)
(b) Prefaringe corta ................................................ Allocreadiidae ............................................ Procaudotestis (Fig. 4-14)
(2) Testes cerca de la extremidad posterior; acetábulo mucho más pequeño que la ventosa oral ................................
................................................................................... Opisthorchiidae ................................................ Cladocystis (Fig. 4-9)
(3) Testes en la mitad posterior del cuerpo; acetábulo más largo que la ventosa oral; faringe grande, con extensiones
anteriores similares a dedos .................................. Fellodistomidae ........................................... Kalipharynx (Fig. 4-15)
c) Glándulas vitelinas consisten de folículos grandes y limitados a dos áreas laterales al acetábulo .................................
.......................................................................................... Allocreadiidae ......................................... Magnivitellinum (Fig. 4-17)
d) Glándulas vitelinas consisten de folículos pequeños y extensos, especialmente en la mitad posterior del cuerpo.
(1) Áreas laterales vitelinas unidas anterior y posteriormente al acetábulo ........................................................................
................................................................................... Allocreadiidae ................................................. Crassicutis (Fig. 4-18)
(2) Áreas laterales vitelinas no unidas anteriormente .... Lepocreadidae ................................................ Eocreadium (Fig. 4-12)
VII. Checklist of Trematoda (Digenea) from Neotropical freshwater fishes
Measurements are given in millimeters except for the circumoral spines and eggs which
are in micrometers (µm). Abreviations used are: A = acetabulum; B = body size; CS =
cirrus sac; E = esophagus; EB = esophagus bulb; FB = forebody; G = gut; GS = genital
sac; HB = hindbody; HC = head collar; HS = hermaphroditic sac; Mir = miracidia; OS =
oral sucker; Ph = pharynx; Rhy = rhynchus.
Acanthostomidae POCHE, 1926

Body long and cylindrical. Oral sucker well developed; circumoral spines
normally present; pharynx present. Acetabulum small. Cirrus sac absent.
Ovary anterior to two testes. Excretory vesicle Y-shaped. Intestinal tract
of marine and freshwater fish and reptiles.
Acanthostostomum LOOSS, 1899
Oral sucker terminal, funnel-shaped; circumoral spines present; prephar-
ynx long; esophagus short; intestinal crura open through anal pores. Testes
near posterior extremity. Genital pore preacetabular. Intestinal tract of fish
and aquatic reptiles.
A. gnerii SZIDAT, 1954: Rhamdia quelen and R. rogersi: Costa Rica & Argentina.
(Fig. 4-1). B = 1.6-2.0; 21-23 circumoral spines; OS = 0.25 x 0.20; Ph =
0.12 x 0.10; Vitelline glands large, 7-9 per side; Egg = 29-37 x 14-16 µm.
Allocreadiidae (LOOSS, 1902) STOSSICH, 1903
Oral sucker subterminal, sometimes with projections. Acetabulum simple,
pre-equatorial. Prepharynx, pharynx and esophagus usually present; ceca
usually long. Cirrus sac present or absent. Testes tandem, postequatorial.
Ovary pretesticular. Genital pore median or submedian. Excretory vesicle
short. Intestine of fish, amphibia and aquatic reptiles.
Crassicutis MANTER, 1936
Body elongate, flattened; tegument thickened. Oral sucker large; esophagus
short; ceca long. Acetabulum smaller than oral sucker. Cirrus sac absent. Genital
pore median, just anterior to acetabulum. Ovary submedian. Seminal receptacle
present. Uterus with few large eggs, Intestine of freshwater and marine fish.
C. chuscoi (PEARSE, 1920) PETERS, 1957: Aequidens pulcher: South America. B =
0.56 x 0.29.
C. cichlasomae MANTER, 1936: Cichlasoma mayorum: México (Yucatan), Costa Rica
and Cuba (Fig. 4-18). B = 0.88-1.3 x 0.48-0.83; Egg = 110-114 x 64-66 µm.
C. opisthoseminis BRAVO-HOLLIS & ARROYO, 1961: Cichlasoma sp.: Costa Rica. B
= 3.3 x 1.6; Egg = 120 x 82 µm.
C. wallini (PEARSE, 1920) PETERS, 1957: Crenicichla geayi: Venezuela. B = 1.55 x
0.74; Egg = 118-130 x 66-71 µm.
Crepidostomum BRAUN, 1900.
Oral sucker with 6 lobes; prepharynx present; ceca long. Cirrus sac
elongate. Ovary submedian; seminal receptacle present. Intestinal tract of
freshwater fishes and reptiles.
C. macrorchis SZIDAT, 1954: Roeboides bonariensis: Argentina. (Fig. 4-10). B = 1.5

x 0.5; OS = 0.18 x 0.20; A = 0.15; Ph = 0.07 x 0.05; Egg = 55 x 29 µm.
C. platense SZIDAT, 1954: Pimelodus clarias and Iheringichthys labrosus: Argentina. B
= 0.8 x 0.2; OS = 0.12; A = 0.12; Egg = 75 x 39 µm.
Creptotrema TRAVASSOS, ARTIGAS & PEREIRA, 1928.
Oral sucker large, with two dorsal lobes. Pharynx present; esophagus short;
ceca long. Acetabulum similar in size to oral sucker; pre-equatorial. Cirrus
sac long. Genital pore medial, near bifurcation. Ovary between testes and
acetabulum. Vitelline follicles large; lateral, extending from pharynx to
posterior extremity. Eggs few, large. Excretory vesicle tubular. Intestine of
freshwater fish.
C. creptotrema TRAVASSOS, ARTIGAS & PEREIRA, 1928: Leporinus elongatus: Brazil.
(Fig. 4-11). B = 0.47 x 0.54 x 0.24-0.28; OS = 0.11-0.13; A = 0.16; Ph =
0.04; Egg = 72-78 x 44-50 µm.
C. díspar TEXEIRA DE FREITAS, 1941: Triportheus paranensis: Brazil. B = 1.8 x
0.67; OS = 0.19-0.23; A = 0.26 x 0.27; Ph = 0.06; Egg = 80 x 46 µm.
Magnivitellinum KLOSS, 1966.
Oral sucker small; esophagus and ceca short. Acetabulum subequal to oral
sucker. Testes diagonal; in middle third of body. Cirrus sac small, curved.
Genital pore median, preacetabular. Ovary between testes and acetabulum.
Vitelline follicles large; extending length of ceca. Intestine of freshwater fish.
M.simplex KLOSS, 1966: Astyanax bimaculatus: Brazil. (Fig. 4-17). B = 2.7-3.5 x
0.74-1.1; OS = 0.20-0.27; A = 0.23-0.30; Ph = 0.086-0.11; Egg = 36-47 x
21-23 µm.
Procaudotestis SZIDAT, 1954
Body tapering posteriorly. Oral sucker larger than acetabulum. Genital
pore median, preacetabular. Cirrus absent; seminal vesicle large. Repro-
ductive organs in posterior half of body; ovary pretesticular; vitellaria
follicular, lateral. Excretory vesicle tubular. Intestine of freshwater fish.
P. uruguayensis SZIDAT, 1954: Loricaria vetula: Uruguay (Fig. 4-14). B = 2.2 x 0.65; OS
= 0.25 x 0.30; A = 0.22 x 0.25; Ph = 0.11 x 0.10; Egg = 80 x 92 x 40-44 µm.
Angiodictyidae LOOSS, 1902.
Oral sucker with or without lumenal diverticula; esophagus and esophageal
bulb present; ceca long. Cirrus sac usually absent. Genital pore median,
near anterior extremity. Ovary post-testicular. Lymphatic ( = circulatory)
system present in association with ceca. Excretory system reticulate. Eggs
large. Intestine of fish and turtles.
Curumai TRAVASSOS, 1961
Body flattened; without posterior projections. Oral sucker small; pharynx
absent; esophagus short; ceca long. Acetabulum absent. Cirrus sac small,
near bifurcation. Testes large, irregular, diagonal; pre-equatorial. Genital
pore anterior to bifurcation. Ovary spherical, post-testicular. Vitelline
follicles forming “V” from cecal ends to posterior testis. Excretory pore
subterminal. Intestine of fish.
C. curumai TRAVASSOS, 1961: Myleus sp.: Brazil (Fig. 4-44). B = 6.5-8.0 x 1.0-2.0;
OS = 0.15-0.19 x 0.12-0.16; E = 0.43-0.78; Egg = 100-104 x 64-68 µm.
Denticauda FUKUI, 1929
Body with 4 projections and median notch posteriorly. Oral sucker simple;
esophageal bulb absent; ceca medium long. Testes tandem. Cirrus sac
absent. Genital sucker present; pore postbifurcal. Ovary median, post-
testicular. Excretory pore in median notch. Intestine of fish.
D. quadrangulata (DADAY, 1907): Colossoma bidens: Brazil. (Fig. 4-45). B = 1.7 x
0.65; OS = 0.13-0.14; Egg = 60 x 50 µm.
Pseudoparabaris YAMAGUTI, 1958
Body with posterior median notch. Oral sucker simple, terminal; esoph-
ageal bulb present; ceca medium long. Testes tandem, lobate. Seminal
vesicle long; cirrus sac absent. Genital pore at level of bifurcation. Ovary
median; vitelline follicles forming “V” with apex near ovary. Intestine of
freshwater fish.
P. parabaris (TRAVASSOS, 1922): Colossoma bidens: Brazil. (Fig. 4-46). B = 10.5 x
4.0; OS = 0.20; Ph = 0.29 x 0.27; Egg = 134 x 78µm.
Bucephalidae POCHE, 1907
Anterior attachment organ a rhynchus. Oral sucker and acetabulum
absent. Pharynx and esophagus present; mouth ventral; gut saccular or
tubular. Cirrus sac near posterior extremity; genital pore terminal or
subterminal. Vitelline follicles usually anterior. Position of testes and ovary
variable. Intestine of fish and amphibians.
Bellumcorpus KOHN, 1962
Body elongate; rhynchus oval, without projections. Pharynx small, in
middle of body; gut large, directed posteriorly. Testes entire or indented,
diagonal, post-equatorial. Cirrus sac small; genital pore terminal. Ovary
opposite posterior testis; vitelline follicles in 4 – 5 lateral groups. Excretory
vesicle long, tubular. Stomach of freshwater fish.
B. major KOHN, 1962: Salminus hilarii: Brazil. (Fig. 4-4). B = 3.54-7.36 x 1.08-
2.25; Rhy = 0.42-0.63 x 0.18-0.28; Ph = 0.19-0.31 x 0.18-0.28; G = 1.37-
2.74 x 0.42-1.08; CS = 0.58-1.66 x 0.22 0.33; Egg = 22-26 x 18 µm.
Glandulorhynchus THATCHER, 1999
Body stout, flattened, unspined; rhynchus sucker-like, provided with
numerous glands. Pharynx small, near equator or posterior thereto; gut
saccular, mostly posterior to pharynx. Testes two, spherical, diagonal,
posterior to pharynx; cirrus sac small, at posterior extremity; genital pore
subterminal. Ovary spherical, anterior to pharynx; vitellaria in grape-like
clusters extending anterolaterally in anterior half of body; uterus tubular
forming massive uterine reservoir on side of body; eggs small, numerous.
Intestinal parasites of freshwater fishes. Type species: Glandulorhynchus
turgidus THATCHER, 1999
G. turgidus THATCHER, 1999: Hydrolycus sp.: Rondonia State, Brazil. (Figs. 4-91
& 4-92). B = 5.0-7.3 x 2.5-3.5; Rhy = 0.56-0.81 x 0.54-0.81; Ph = 0.21-0.31
x 0.25- 0.35; G = 0.60-1.9 x 0.46-0.62; CS = 1.2-1.5 x 0.27-0.42; Egg = 18-

20 x 9-11 µm.
Pararhipidocotyle KOHN, 1970
Body elongate, spinous. Rhynchus with cap; opening subterminal. Mouth
in middle of body; gut directed posteriorly. Testes parallel or diagonal, in
middle third of body. Cirrus sac elongate; genital pore ventral. Ovary post-
testicular. Vitelline follicles prececal. Excretory pore terminal. Intestine of
freshwater fish.
P. jeffersoni KOHN, 1970: Salminus maxillosus: Brazil. B = 0.90-1.04 x 0.22-0.29;
Rhy = 0.11-0.17 x 0.11-0.16; Ph = 0.054-0.063 x 0.57-0.060; E = 0.045-
0.075; G = 0.13-0.14 x 0.078-0.087; CS = 0.27-0.32 x 0.06-0.07; Egg = 30-
40 x 15-21 µm.
Paurorhynchus DICKERMAN, 1954
Body subcylindrical. Rhynchus small, funnel-shaped. Mouth in anterior
third; gut long, directed posteriorly. Testes large, lobate, diagonal, post-
equatorial. Cirrus sac small; genital atrium small; pore ventral. Ovary
lobate; opposite anterior testis. Vitellaria laterial, acinous. Excretory vesicle
long. Gall bladder of freshwater fish.
P. schubarti KOHN, 1963: Salminus maxillosus: Brazil. (Fig. 4-5). B = 7.73 x 2.8;
Rhy = 0.50 x 0.29; Ph = 0.29-0.32; G = 2.13-0.67; CS = 2.4-0.33; Egg =
not found.
Prosorhynchus ODHNER, 1905
Body rounded to elongate; spinous. Rhynchus plug or funnel-shaped,
without projections. Mouth in middle third; gut short. Testes tandem or
diagonal, in middle third or more posterior. Genital pore ventral or
terminal. Intestine of freshwater and marine fish.
P. costai TRAVASSOS, ARTIGAS & PEREIRA, 1928: Acestrorhamphus sp., Astyanax
fasciatus and Salminus hilarii: Brazil (Fig. 4-6). B = 1.0 x 0.2; Rhy = 0.13 x
0.12; Ph = 0.046; G = 0.15; Egg = 42-46 x 15 µm.
P. piranhus THATCHER, 1999: Serrasalmus rhombeus: Rondonia State, Brazil. (Fig.
4-90). B = 0.78-1.1 x 0.42-0.75; Rhy = 66-119 x 88-132 µm; Ph = 66-83 x
66-83 µm; G = 88-198 x 73-110 µm; CS = 0.18-0.49 x 0.07-0.11; Egg =
44-55 x 22-31 µm.
Callodistomidae POCHE, 1926
Body rounded or elongate. Oral sucker, pharynx and acetabulum present.
Ceca of variable length. Cirrus sac and genital pore preacetabular. Two
testes and ovary present; of variable position. Vitellaria lateral. Uterus
extensive; eggs small, numerous. Intestine and other organs of fish.
Prosthenhystera TRAVASSOS, 1920
Body elliptical to oval. Oral sucker and pharynx small; esophagus short;
ceca long. Acetabulum small, pre-equatorial. Testes parallel; in middle
third. Cirrus sac elliptical, postbifurcal. Genital pore just postbifurcal.
Ovary between acetabulum and left testis. Vitellaria in lateral groups. Gall
bladder of freshwater fish.
P. obesa (DIESING, 1850) Salminus, Leporinus, Boulengerella, Acestrorhamphus,

Galeocharax, Pseudopimelodus, Pimelodus and Astyanax: Mexico, Colombia &
Brazil. (Fig. 4-7). B = 14 x 8; OS = 1.0-1.2; A = 0.95-1.3; Ph = 0.34; E =
1.3; Egg = 63-70 x 35-49 µm.
Cryptogonimidae CIUREA, 1933
Body small, oval or elongate. Oral sucker sometimes provided with
circumoral spines; pharynx present; esophagus short; ceca variable. Ace-
tabulum small, embedded. Testes single or multiple; post-equatorial.
Cirrus sac frequently absent. Ovary usually lobate. Vitellaria variable.
Excretory vesicle V or Y-shaped. Intestine of freshwater and marine fish
and aquatic reptiles.
Iheringtrema TRAVASSOS, 1948
Body small, scaled. Oral sucker large; pharynx present; esophagus short:
ceca thin, medium long. Acetabulum small, within genital atrium; in anterior
quarter. Testes 9 (4 on one side and 5 on the other); in posterior half.
Seminal vesicle long, sigmoid, postacetabular; cirrus sac absent. Genital pore
median; just preacetabular. Ovary of 3 notched lobes; pretesticular. Seminal
receptacle present. Vitellaria extensive. Intestine of fish.
I. iheringi TRAVASSOS, 1948: Pseudopimelodus roosevelti: Brazil. (Fig. 4-22). B = 3.5 x 4.0
x 1.0-1.2; OS = 0,38-0.39; A = 0.17-0.18; Ph = 0.15-0.18; Egg = 38 x 22 µm.
Palaeocryptogonimus SZIDAT, 1954
Body tapered posteriorly; scaled anteriorly. Oral sucker terminal, retractile,
without spines; prepharynx present; pharynx large; esophagus short; ceca
medium long. Acetabulum near anterior extremity; covered by circular
pad. Testes nearly parallel; in midregion. Seminal vesicle tripartite; postac-
etabular. Cirrus sac absent. Genital pore between acetabulum and circular
pad. Ovary median, pretesticular. Vitellaria of large follicles; lateral in
acetabular region. Seminal receptacle present. Excretory vesicle. Y-shaped,
pore terminal. Stomach and intestine of fish.
P. claviformis SZIDAT, 1954; Rhinodoras dorbignyi: Argentina. (Fig. 4-23). B = 1.25
x 0.45; OS = 0.12; Ph = 0.09 x 0.06; Egg = not found.
Didymozoidae POCHE, 1907
Hemaphroditic or partly gonochoristic. Usually encysted in pairs in tissue.
Body form variable; frequently thread-like, or with slender forebody and
expanded hindbody. Pharynx and acetabulum present or absent. Repro-
ductive organs usually elongate, tubular, winding. Uterus extensive; eggs
small, numerous. Excretory vesicle long, pore terminal. Tissues of marine,
and rarely freshwater fish.
Brasicystis THATCHER, 1979
Hermaphroditic; encysted in pairs in subcutaneous tissue. Body divided
into slender forebody and expanded hindbody; hindbody laterally flat-
tened, flexed ventrally in anterior part and curved ventro-laterally in
posterior part. Oral sucker, pharynx and esophagus present; ceca long.
Acetabulum absent. Testis single, slender, extends to posterior extremity.
Seminal vesicle present; cirrus sac absent. Genital pore lateral to oral
sucker. Ovary long, tubular; shorter than testis. Vitelline gland single,
tubular, extends to posterior extremity on dorsal side. Uterus extensive,
eggs small. Subcutaneous cysts in freshwater fish.
B. bennetti THATCHER, 1979: Plagioscion squamosissimus: Brazil (Figs. 4-3 & 4-47).
FB = 1.2-2.8 x 0.10-0.22; HB = 9.5-16 x 1.0-1.2; OS = 0.04-0.06 x 0.02-
0.03; Ph = 0.03-0.05; Egg = 14-16 x 8-9 µm.
Echinostomatidae POCHE, 1926
Body elongate, normally with head collar bearing 1-2 rows of spines;
tegument spined or scaled. Oral sucker, prepharynx and esophagus
present; ceca long. Acetabulum large; in anterior or middle third. Testes
tandem or diagonal. Genital pore median; preacetabular. Ovary pretesticu-
lar; seminal receptacle absent. Vitelline follicles lateral and posterior. Eggs
large. Excretory vesicle Y-shaped. Intestine of reptiles, birds, mammals
and rarely fish.
Caballerotrema PRUDHOE, 1960, THATCHER, 1980
Body long, cylindrical. Head collar wide and flattened, with single row of
spines; ventral lobe (on each side) with 4 spines in 2 pairs. Oral sucker
small, with membranous extension antero-dorsally and laterally; esophagus
long. Acetabulum large, anterior. Cirrus sac large; extending posterior to
acetabulum; external seminal vesicle present. Gonads in midbody. Vitelline
follicles dorsolateral from preovarian region to cecal ends. Intestine of
freshwater fish.
C. arapaimense THATCHER, 1980: Arapaima gigas: Brazil (Amazonia). (Fig. 4-16).
B = 3,8-8.3 x 0.62-0.77; HC = 0.83-1.02; large spines = 84-200 x 27-48
µm; small spines = 67-139 x 18-36 µm ; OS = 0.14-0.30; E = 0.31-0.36; Ph
= 0.26-0.36 x 0.11-0.27; A = 0.33-0.87 x 0.29-0.56; CS = 0.47-0.82 x 0.22-
0.42; Egg = 72-83 x 28-58 µm.
C. aruanense THATCHER, 1980: Osteoglossum bicirrhosum: Brazil (Amazônia). B =
5,8-9.1 x 0.39-0.62; HC = 0.63-0.88; large spines = 72-98 x 24-32 µm;
small spines = 47-65 x 28 µm ; OS = 0.11-0.16; E = 0.30-0.42; Ph = 0.18-
0.22 x 0.12-0.16; A = 0.36-0.45 x 0.31-0.42; CS = 0.74-1.2 x 0.21-0.25; Egg
= 70-97 x 43-60 µm.
C. brasiliense PRUDHOE, 1960, THATCHER, 1980: Arapaima gigas: Brazil (Amazo-
nia). B = 4.6-11.3 x 0.51-0.74; HC = 0.60-1.0; large spines = 67-90 x 26-47
µm; small spines = 34-60 x 14-35 µm ; OS = 0.11-0.19; E = 0.25-0.45; Ph
= 0.19-0.30 x 0.09-0.14; A = 0.29-0.41 x 0.25-0.40; CS = 0.38-0.64 x 0.26-
0.35; Egg = 80-84 x 48-56 µm.
Fellodistomidae NICOLL, 1913
Body shape variable. Oral sucker, pharynx and esophagus present; ceca of
variable length, sometimes forming ring. Acetabulum medium to large.
Testis or testes in posterior half. Cirrus and genital pore usually preacetab-
ular. Ovary variable, usually pretesticular. Vitellaria lateral, variable in
form. Intestine of marine and freshwater fish.
Antorchis LINTON, 1911

Body fusiform spinous. Oral sucker large, cup-shaped with notched
ventral lip; pharynx small; esophagus long; ceca short. Acetabulum in
middle third. Testes parallel, anterolateral to acetabulum. Cirrus sac and
genital pore preacetabular. Ovary dorsal or posterior to acetabulum.
Vitelline follicles pretesticular, extracecal. Uterus occupying hindbody.
Excretory vesicle V or Y-shaped. Intestine of freshwater and marine fish.
A. lintoni TRAVASSOS, ARTIGAS & PEREIRA, 1928: Astyanax fasciatus: Brazil. (Fig.
4-13). B = 0.9 x 0.4; OS = 0.15; A = 0.15; Ph = 0.06; Egg = 30 x 15 µm.
Bacciger NICOLL, 1914
Body rounded to oval, spinous. Oral sucker ventro-terminal; pharynx well
developed; esophagus short; ceca reaching to or slightly beyond testes.
Acetabulum equatorial or pre-equatorial. Testes spherical or ovoid, sym-
metrical or subsymmetrical at postacetabular level. Cirrus sac pyriform to
oval or elliptical, preacetabular, enclosing saccular seminal vesicle, prostat-
ic complex and ejaculatory duct. Genital pore median, postbifurcal. Ovary
rounded or weakly lobate, inter or post-testicular; seminal receptacle and
Laurer’s canal present; vitelline follicles in symmetrical groups outside
ceca; uterine coil occupying most of hindbody; Eggs small, numerous.
Intestinal parasites of marine and freshwater teleosts.
B. pellonae THATCHER, 1992: Pellona castelnaeana: Guaporé River, Rondônia
State, Brazil. (Fig. 4-98). B = 0.60-0.74 x 0.35-0.49; A = 50-94 µm; OS =
72-88 x 82-105 µm; CS = 82-116 x 5-82 µm; Eggs = 28-44 x 19-28 µm.
Kalipharynx BOEGER & THATCHER, 1984
Body elongate, tapering posteriorly, tegument spinous. Oral sucker subter-
minal, large, but with weak musculature; pharynx large, strongly muscled,
with digitiform processes anteriorly; esophagus short; ceca long. Acetabu-
lum large, shallow; pre-equatorial. Testes diagonal, post-acetabular, posto-
varian. Cirrus sac long, thin. Genital pore ventro-lateral; between suckers.
Vitelline follicles lateral, postacetabular. Uterus in posterior half of body;
eggs small, numerous. Excretory pore terminal. Intestine of fish.
K. piramboae BOEGER & THATCHER, 1984: Lepidosiren paradoxa: Brazil. (Fig. 4-
15). B = 1.65-3.25 x 0.73-0.88; OS = 0.13-0.20 x 0.23-0.33; Ph = 0.18-0.25
x 0.25-0.30 (with 6 anterior digitiform processes); A = 0.53-0.83; CS =
0.49-0.94 x 0.06-0.10; Egg = 25-40 x 13-20 µm.
Gorgoderidae LOOSS, 1901
Body narrowed anteriorly, expanded posteriorly; tegument unspined. Oral
sucker present; pharynx often absent; ceca long, may be sinuous, some-
times forming ring. Acetabulum large, projecting. Testes two, or more,
postacetabular, parallel or diagonal. Cirrus sac absent. Genital pore
medium, between bifurcation and acetabulum. Ovary submedian, postace-
tabular. Vitellaria double, compact or lobed, postacetabular. Uterus exten-
sive in hindbody; eggs small. Excretory pore terminal. Instestine or urinary
bladder of fishes, amphibians and reptiles.
Amazonadistoma THATCHER, 1979

Body cylindrical and tapered anteriorly, rounded and flattened posteriorly;
tegument unspined but with muscular corrugations postero-laterally. Oral
sucker large; esophagus short; ceca long, with diverticula and microvilli;
bifurcation near posterior margin of oral sucker. Acetabulum large, pre-
equatorial. Testes parallel or diagonal, postequatorial. Seminal vesicle
small, sinuous. Genital pore and genital sucker between oral sucker and
acetabulum. Ovary submedian, pretesticular. Vitelline glands, two parallel
masses; pretesticular. Uterus extensive, sinuous; eggs small. Excretory
vesicle long, tubular; pore terminal. Intestine of freshwater fish.
A. negrensis THATCHER, 1979: Gymnorhamphichthys hypostomus: Brazil. (Fig. 4-2). B
= 5.3-6.3 x 2.2-2.6; OS = 0.66-0.76; A = 0.53-0.59; Egg = 30 x 23 µm.
Dendrorchis TRAVASSOS, 1926
Body pyriform, flattened. Oral sucker ventro-terminal; bifurcation nearer
oral sucker than acetabulum; ceca long. Acetabulum similar in size to oral
sucker. Testes diagonal, branched. Cirrus sac small, transverse. Genital
pore submedian, between bifurcation and acetabulum. Ovary submedian
at level of anterior testis; vitelline glands transversely oval, between
acetabulum and ovary. Uterus extensive. Excretory vesicle tubular, pore
terminal. Swimbladder of fish.
D. neivai TRAVASSOS, 1926; Brycon lundi: Brazil. (Fig. 4-19). B = 6-8 x 4-5; OS =
0.6-0.7; A = 0.67-0.9; E = 0.4-0.8; Egg = 35-53 x 30-38 µm.
Phyllodistomoides BROOKS, 1977.
Body pyriform, without spines. Oral sucker subterminal; prepharynx
absent; pharynx present; esophagus and ceca long. Acetabulum pre-
equatorial. Testes compact, intercecal, postacetabular. Ovary submedian,
pretesticular, postacetabular. Vitelline glands lateral to acetabulum. Uterus
extensive. Swimbladder of fish.
P. ducani BROOKS, 1977: Astyanax sp.: Colombia. (Fig. 4-25). B = 1.7 x 0.85; OS
= 0.28-0.34; Ph = 0.96 x 0.11; E = 0.96; A = 0.18 x 0.20; Egg = 46 x 29 µm.
Halipegidae POCHE, 1926
Body rounded or elongate, without ecsoma. Oral sucker subterminal;
pharynx small; esophagus short; ceca long, sometimes forming ring.
Acetabulum large. Testes parallel or diagonal, postacetabular. Seminal
vesicle tubular or saccular; hermaphroditic duct short, usually within sac or
genital cone; genital atrium near bifurcation. Ovary near posterior extrem-
ity; vitelline glands of two masses or several lobes, postovarian. Uterus
extensive; eggs sometimes filamented. Excretory branches united anterior-
ly. Stomach of marine and freshwater fish and amphibians.
Gonocercella MANTER, 1940
Body subcylindrical; tegument transversely annulated. Oral sucker subter-
minal, large with preoral lobe; esophagus short; ceca long. Acetabulum
larger than oral sucker, postequatorial. Testes diagonal, postacetabular;
seminal vesicle coiled, anterior to acetabulum; prostatic vesicle large;
hermaphroditic duct within genital cone which projects into conical

genital atrium; genital pore median, postbifurcal. Ovary post-testicular;
vitellaria of two compact or lobed masses, postovarian. Egg without
filaments. Stomach of freshwater fish.
G. magnifica SZIDAT, 1954: Hypostomus plecostomus and H. commersoni: Argentina. B =
5 x 2; OS = 0.45; A = 0.66-0.75; Ph = 0.20 x 0.22; Egg = 50-55 x 20-22 µm.
Halipegus LOOS, 1899
Body elongate, unspined. Oral sucker with preoral lobes; pharynx present;
esophagus short; ceca long. Acetabulum large. Testes nearly parallel, postac-
etabular. No cirrus sac and no hermaphroditic bursa; seminal vesicle
saccular. Genital pore postero-ventral to pharynx. Ovary near posterior
extremity; vitelline glands postovarian, of two compact lobes or two groups
of 4 – 5 lobes each. Uterus extensive; eggs with polar filaments. Excretory
branches uniting near anterior extremity. Digestive tract of frogs and fish.
H. dubius KLEIN, 1905: Salminus maxillosus and Cynopotamus humeralis: Brazil.
(Fig. 44-20). B = 3.3-4.0 x 1.1-1.5; OS = 0.36-0.49 x 0.46-0.53; A = 0.54-
0.66 x 0.61-0.66; Egg = 42-52 x 17-22 µm. (filaments = 138-221 µm).
H. genarchella (TRAVASSOS, ARTIGAS & PEREIRA, 1928) Astyanax, Acestrorhamphus,
Cynopotamus, Moenklausia and Roeboides: Brazil. B = 2.1- 4 x 0.78-1.6; OS =
0.36-0.52; A = 0.68-1.0; Ph = 0.13-0.18; Egg = 48-58 x 24 µm. (polar
filaments present).
H. parvus (TRAVASSOS, ARTIGAS & PEREIRA, 1928): Acestrorhamphus and Salminus:
Brazil. B = 2.1-2.6 x 0.52-0.53; OS = 0.42-0.65; A = 0.36-0.39; Ph = 0.1;
Egg = 85 x 57 µm.
H. tropicus (MANTER, 1936): Astyanax, Charax, Leporinus, Luciopimelodus, Loricaria
and Pimelodus: México, Brazil & Argentina. B = 2.27-4.2 x 0.45-0.7; OS = 0.29-
0.43; Ph = 0.12-0.16; A = 0.48-0.58 x 0.52-0.66; Egg = 44-52 x 23-29 µm.
Haploporidae NICOLL, 1914
Body small, rounded or conical; spinous. Oral sucker subterminal;
prepharynx present; esophagus long; ceca usually short or saccular.
Acetabulum in middle third of body. Testis postacetabular; seminal vesicle
internal and external to hermaphroditic sac. Genital pore median, preace-
tabular. Ovary small, median, anterior to testis; vitellaria of paired lobes or
bunches, lateral or posterior to ovary. Uterus extensive; eggs large,
miracidial eyespot frequently visible. Excretory vesicle variable, pore
terminal. Intestine of freshwater and marine fish.
Chalcinotrema TEXEIRA DE FREITAS, 1947
Body elongate, spinous. Oral sucker subterminal; prepharynx present;
esophagus long; ceca short to medium. Acetabulum similar in size to oral
sucker. Testis near posterior extremity; hermaphroditic sac and external
seminal vesicle present. Genital pore submedian, preacetabular. Ovary far
anterior to testis; seminal receptacle present; vitellaria of elongate lobes,
lateral, in posterior half of body. Uterus extensive; eggs numerous.
Excretory vesicle tubular. Intestine of freshwater fish.
C. lucieni BROOKS 1977: Leporinus muyscorum: Colombia. (Fig. 4-26). B = 4.44-

5.15 x 1.6-1.9; OS = 0.29-0.37 x 0.40-0.47; A = 0.44-0.55; Ph = 0.20-0.24
x 0.19-0.23; E = 0.66-0.88; Egg = 104-120 x 75-89 µm.
C. ruedasueltensis THATCHER, 1978: Astyanax fasciatus: Colombia. (Fig. 4-33). B =
1.4-1.5 x 0.46-0.47; OS = 0.15-0.16; Ph = 0.081-0.082; A = 0.14-0.15; HS
= 0.15-0.16 x 0.075-0.10; Egg = 90-102 x 50-60 µm.
C. salobrensis TEXEIRA DE FREITAS, 1947: Tryportheus paranensis: Brazil. B = 4.2 x
1.4; OS = 0.36; A = 0.35 x 0.32; Ph = 0.26; Egg = 96 x 66 µm.
Lecithobotrioides THATCHER & DOSSMAN, 1974
Body cylindrical, spinous. Acetabulum in anterior half of body. Oral
sucker subterminal; prepharynx short; pharynx smaller than sucker;
esophagus long; ceca medium long. Testis in posterior half of body;
hermaphroditic sac present; external seminal vesicle elongate, curved.
Genital pore preacetabular. Ovary spherical or ovoid, in middle third;
seminal receptacle absent; vitellaria variable, extensive, lateral. Uterus
posterior to genital pore; eggs few, large without miracidial eyespots.
Excretory vesicle saccular, pore terminal. Intestine of freshwater fish.
L. mediacanoensis THATCHER & DOSSMAN, 1974. Prochilodus reticulatus: Colombia.
(Fig. 4-31). B = 2.1-3.7 x 0.63-0.67; OS = 0.21-0.27; Ph = 0.13-0.18; A =
0.22-0.27; HS = 0.37-0.78 x 0.11-0.26; Egg = 74-100 x 45-52 µm.
L. elongatus THATCHER, 1999: Prochilodus nigricans: Guaporé River, Rondonia
State, Brazil. (Fig. 4-94). B = 3.0-3.9 x 0.83-1.0; OS = 0.17-0.22 x 0.19-
0.30; Ph = 0.16-0.17 x 0.17-0.24; A = 0.37-0.39 x 0.37-0.39; HS = 0.26-
0.59 x 0.19-0.30; Egg = 90-118 x 57-73 µm.
Megacoelium SZIDAT, 1954, THATCHER & VARELLA, 1981.
Body elongate, tapered posteriorly, compressed laterally. Tegument with
horizontal rows of spines to posterior extremity; spines usually present
inside both suckers. Oral sucker large, subterminal; prepharynx short;
pharynx large; esophagus short; ceca medium long, laterally compressed.
Acetabulum large, pre-equatorial. Testis elongate, sometimes sinuous,
postequatorial; hermaphroditic sac pyriform or oval, containing hermaph-
roditic cirrus; external seminal vesicle tubular; genital pore median or
submedian. Ovary spherical, median, anterodorsal to testis; uterus exten-
sive; uterine seminal receptacle sometimes present. Eggs numerous, with
short polar projection opposite operculum; with or without miracidial
eyespots. Excretory pore terminal. Stomach of freshwater fish.
M.plecostomi SZIDAT, 1954: Hypostomus plecostomus: Argentina B = 2.7 x 1.7; Egg
= 130-138 x 70 µm.
M.spinicavum THATCHER & VARELLA, 1981: Pterygoplichthys pardalis: Brazil
(Amazônia). (Fig. 4-24 A-B). B = 3.5-4.9 x 1.2-1.7; OS = 0.52-0.58 x 0.48-
0.55; Ph = 0.27-0.37; A = 0.64-0.75 x 0.55-0.79; HS = 0.37-0.64 x 0.2-
0.30; Egg = 95-110 x 50-60 µm.
M.spinispecum THATCHER & VARELLA, 1981: Pterygoplichthys sp.: Brazil. B = 5.4-

7.7 x 2.7-3.0; OS = 0.88-0.95 x 0.81-0.88; Ph = 0.45-0.47; A = 1.2; HS =
0.64-1.0 x 0.33-0.45; Egg = 83-100 x 43-57 µm.
Paralecithobotrys TEXEIRA DE FREITAS, 1947
Body elongate, unspined. Oral sucker subterminal; prepharynx absent;
pharynx present; esophagus and ceca long. Acetabulum slightly larger than
oral sucker, pre-equatorial. Testis in midbody; hermaphroditic sac U-
shaped, preacetabular, genital pore submedian. Ovary immediately pretes-
ticular; vitellaria of few scattered follicles in midbody; uterus extensive;
eggs with miracidial eyespots. Excretory vesicle Y-shaped, pore terminal.
Intestine of freshwater fish.
P. brasiliensis TEXEIRA DE FREITAS, 1947. Lahilliela kneri and Leporinus elongatus.:
Brazil. (Fig. 4-36). B = 2.1-4.7 x 0.74-1.4; OS = 0.25-0.41 x 0.28-0.41; Ph =
0.14-0.22 x 0.10-0.16; A = 0.33-0.55 x 0.36-0.53; Egg = 87-113 x 52-70 µm.
Rondotrema THATCHER, 1999
Body small, stout, tapering towards both extremities, not flattened; tegument
not spinous. Oral sucker large, subterminal; prepharynx short; pharynx large,
spherical; esophagus long; ceca medium long, slender. Acetabulum large,
pre-equaorial. Testis small, single, spherical or irregular, postequatorial;
hermaphroditic sac large, pyriform, containing internal seminal vesicle,
cirrus and distal part of uterus; genital pore medial, immediately preacetab-
ular. Ovary ovoid, pretesticular, lateral to midline; vitellaria of two grape-like
clusters, one on each side, lateral to testis but intercecal; uterus with
descending and ascending loops; eggs large, containing miracidia with eye-
spots. Excretory vesicle tubular, pore terminal. Intestinal parasites of
freshwater fishes. Type species: Rondotrema microvitellarum, THATCHER, 1999.
R. microvitellarum THATCHER, 1999; Hemiodus microlepis: Guaporé River, Ron-
donia State, Brazil. (Fig. 4-93). B = 0.63-1.2 x 0.31-0.59; OS = 0.11-0.16 x
0.13-0.21; Ph = 55-79 x 66-92 µm; Ceca = 22-66 µm in diameter; A =
0.11-0.17 x 0.14-0.22; Egg = 42-66 x 22-26 µm.
Saccocoelioides SZIDAT, 1954, THATCHER, 1978
Body fusiform or elongate, tegument spined or not. Oral sucker subtermi-
nal; prepharynx short; pharynx similar in size to oral sucker; esophagus long;
ceca medium or long. Acetabulum nearly equatorial, similar to oral sucker in
size. Testis spherical or oval, postequatorial; hermaphroditic sac larger than
pharynx; internal and external seminal vesicles present; genital pore median
or submedian, between acetabulum and pharynx. Ovary spherical, immedi-
ately pretesticular; vitelline follicles dorso-lateral, usually extending from
acetabulum to posterior margin of testis; uterine coils on both side of body,
from genital pore to testis; eggs with or without miracidial eyespots.
Excretory vesicle Y-shaped, pore terminal. Intestine of freshwater fish.
S. leporinodus THATCHER, 1978: Leporinodus vittatus: Colombia. (Fig. 4-34). B =
8.0-8.6 x 0.37-0.46; OS = 0.093-0.13; Ph = 0.09-0.13; A = 0.10-0.12; HS =
0.093-0.11 x 0.11-0.14; Egg = 62-80 x 29-34 µm.
S. magniovatus SZIDAT, 1954: Leporinus obtusidens: Argentina. B = 0.7 x 0.36; OS

= 0.11; Ph = 0.09; A = 0.12; Egg = 95-114 x 41-60 µm.
S. magnorchis THATCHER, 1978: Saccodon caucae: Colombia. B = 0.54-0.96 x 0.24-
0.51; OS = 0.075-0.10; Ph = 0.053-0.12; A = 0.094-0.11; HS = 0.067-0.10
x 0.11-0.19; Egg = 68-72 x 26-30 µm.
S. nanii SZIDAT, 1954: Prochilodus lineatus: Argentina. B = 0.72 x 0.24 – 0.30; OS
= 0.1; Ph = 0.055; A = 0.1; Egg = 85 x 38 µm.
S. octavus SZIDAT, 1970: Astyanax fasciatus: Argentina. B = 1.2 x 0.32; OS =
0.11; Ph = 0.08; A = 0.08; Egg = 110-150 x 60 µm.
S. quintus SZIDAT, 1954: Loricaria anus: Argentina. B = 1.12-1.13 x 0.49-0.50;
OS = 0.09; Ph = 0.06; A = 0.13; Egg = not given.
S. rotundus THATCHER & JÉGU, 1996: Mylesinus paraschomburgkii: Trombetas
River, Pará State, Brazil. (Fig. 4-58). B = 0.53-0.90 x 0.34-0.47; A = 0.12-
0.19 x 0.13-0.19; OS = 0.10-0.16 x 0.13-0.18; Ph = 53-115 x 80-103 µm.
HB = 0.14-0.18 x 0.078-0.10; Eggs = 71-109 x 32-42 µm.
S. saccodontis THATCHER, 1978: Saccodon caucae: Colombia. (Fig. 4-35). B = 0.74-
1.2 x 0.31-0.38; OS = 0.80-0.10; Ph = 0.082-0.11; A = 0.11-0.13; HS =
0.082-0.11 x 0.11-0.15; Egg = 64-75 x 29-32 µm.
S. szidati TRAVASSOS, TEXEIRA DE FREITAS & KOHN, 1969: Schizodon fasciatus: Brazil.
B = 1.5 x 0.7; OS = 0.16; Ph = 0.14; A = 0.16; Egg = 110-118 x 48-53 µm.
Unicoelium THATCHER & DOSSMAN, 1975
Body fusiform, spined. Oral sucker subterminal, prepharynx short; phar-
ynx smaller than suckers; ceca united to form bilobed sac, equatorial.
Acetabulum in midbody. Testis large, oval, in posterior third of body;
hermaphroditic sac preacetabular; external seminal vesicle pyriform; geni-
tal pore submedian, posterior to pharynx. Ovary small, spherical, just pre-
testicular; uterine seminal receptacle prominent, looped laterally from
ovary to near pharynx; uterus looped laterally on both sides, from pharynx
to testis; eggs large, with miracidial eyespots visible in distal portion of
uterus. Excretory vesicle Y-shaped. Intestine of freshwater fish.
U. prochilodorum THATCHER & DOSSMAN, 1975: Prochilodus reticulatus: Colombia
(Fig. 4-43). B = 0.96-1.62 x 0.37-0.66; OS = 0.14-0.19; Ph = 0.10-0.15; A =
0.15-0.24; HS = 0.13-0.22 x 0.052-0.15; Egg = 73-74 x 29 x 37 µm.
Heterophyidae ODHNER, 1914
Body small, with scale-like spines. Oral sucker and pharynx present; ceca
of variable length. Acetabulum usually present, median or submedian,
often enclosed in genital atrium. Testes single or double, variable in form
and location; cirrus sac absent; seminal vesicle well developed; genital pore
median or submedian, postbifurcal. Ovary usually pretesticular, seminal
receptacle normally present; vitellaria lateral, uterus extensive, eggs small.
Excretory pore terminal. Intestinal tract of vertebrates.
Parspina PEARSE, 1920
Body small, rounded, spined. Oral sucker terminal, with single circle of
spines, prepharynx and pharynx present, esophagus short; ceca long.
Acetabulum smaller than oral sucker, pre-equatorial. Testes nearly parallel,

postequatorial; genital sac encloses large seminal vesicle and short ejacula-
tory duct; latter unites with distal end of uterus forming short hermaphro-
ditic duct which opens on anterior margin of acetabulum. Ovary spherical,
antero-ventral to testes; seminal receptacle present; vitellaria acinous,
extend along ceca from pharynx to ovary; uterus extensive; eggs small.
Excretory vesicle Y-shaped, pore terminal. Intestine of freshwater fish.
P. argentinensis SZIDAT, 1954: Pimelodus clarias: Argentina (Fig. 4-21). B = 0.86-
0.94 x 0.42; OS = 0.18 x 0.15 (with circle of 26 spines); Ph = 0.08 x 0.06;
Egg = 31 x 30 µm.
Lepocreadiidae NICOLL, 1935
Body variable, often oculate. Oral sucker occasionally provided with lobes;
prepharynx usually present; pharynx and esophagus present; ceca long.
Acetabulum pre-equatorial. Testes postequatorial; cirrus sac present;
genital pore variable in position. Ovary median or submedian, pretesticu-
lar; seminal receptacle present; vitellaria extensive. Excretory pore termi-
nal. Digestive tract of marine and freshwater fish.
Eocreadium SZIDAT, 1954
Body spatulate, with rounded ends; unspined. Oral sucker subterminal;
prepharynx and esophagus short; ceca long. Acetabulum smaller than oral
sucker, pre-equatorial. Testes parallel, in middle third; cirrus sac slender,
dorsal to acetabulum; external seminal vesicle present; genital pore just
anterior to acetabulum. Ovary trilobed, submedian, pretesticular; vitelline
follicles small, extensive; eggs large, few. Excretory vesicle tubular, pore
terminal. Stomach of freshwater fish.
E. intermedium SZIDAT, 1954: Hypostomus plecostomus: Argentina. (Fig. 4-12). B =
2.8-3.0 x 1.0; OS = 0.25-0.30; Ph = 0.15-0.18; A= 0.20-0.22; Egg = 110-
120 x 60-68 µm.
Monorchiidae ODHNER, 1911.
Body small, spinous. Oral sucker and acetabulum usually small. Pharynx
present; ceca usually long. Testes usually single, occasionally double
frequently located in hindbody. Cirrus sac well developed; metraterm well
developed, spinous. Parasites of marine and sometimes freshwater fishes
Genolopa LINTON, 1910
Body fusiform, spinous. Oral sucker funnel-shaped; pharynx well devel-
oped; esophagus short; ceca not reaching posterior extremity. Acetabulum
pre-equatorial. Testis single, postacetabular. Cirrus sac large, extending
posterior to acetabulum; genital atrium spined, opening between intestinal
bifurcation and acetabulum. Ovary compact or lobed, pretesticular;
vitellaria forming lateral bunches; uterus filling hindbody; metraterm
spined distally. Excretory vesicle small, with terminal pore. Parasites of
marine and freshwater teleosts.
G. magnacirrus THATCHER, 1996: Boulengerella lucia: Jamarí River, Rondonia
State, Brazil. (Figs. 4-70 & 4-72). B = 0.57-1.4 x 0.17-0.36; OS = 0.07-0.12
x 0.07-0.14; Ph = 0.06-0.09 x 0.05-0.09; A = 0.78-0.12 in diameter; CS =

0.30-0.56 x 0.05-0.15; Eggs = 16-23 x 8-10 µm.
Paraproctotrema YAMAGUTI, 1934
Body spindle-shaped, spinous. Oral sucker subterminal; pharynx large;
esophagus longer than pharynx; ceca not reaching posterior extremity.
Acetabulum larger than oral sucker, pre-equatorial. Testis single, postequa-
torial. Cirrus sac extending posterior to acetabulum. Genital atrium
spined, opening anterior to acetabulum. Ovary lobed, postero-lateral to
acetabulum. Vitellaria of large compact follicles on each side in ovariotes-
ticular zone. Metraterm spinous. Eggs numerous, small. Parasitic in marine
and freshwater teleosts.
P. delicata THATCHER, 1996: Boulengerella lucia: Jamarí River, Rondonia State,
Brazil. (Figs. 4-68 & 4-71). B = 0.79-1.7 x 0.21-0.28; OS = 0.07-0.10; Ph =
0.046-0.074 x 0.04-0.069; A = 0.09-0.11 in diameter; CS = 0.18-0.30 x
0.041-0.078; Eggs numerous, small, 16-18 x 7-9 µm.
Sphericomonorchis THATCHER, 1996
Body small, ovoid, slightly flattened, spinous. Oral sucker subterminal;
prepharynx short; pharynx spherical; esophagus short; ceca long, stout.
Acetabulum pre-equatorial. Testis single, large, spherical, near posterior
extremity; cirrus sac large, elongate, containing spinous cirrus, seminal
vesicle and prostatic cells; genital pore medial, between intestinal bifurca-
tion and acetabulum. Ovary subspherical or irregularly lobate, immediately
pretesticular, lateral to midline; vitellaria of two bunches of large follicles,
lateral to acetabulum; metraterm elongate, saccular, spinous; uterus occu-
pying space between testis and acetabulum. Eggs small, numerous.
Excretory vesicle tubular, pore terminal. Intestinal parasites of freshwater
fish. Type species: Sphericomonorchis spinulosus THATCHER, 1996.
S. spinulosus THATCHER, 1996: Crenicichla johanna: Jamari River, Rondonia State,
Brazil. (Fig. 4-69). B = 0.66-1.2 x 0.17-0.46; OS = 0.08-0.11 x 0.093-0.13;
Ph = 0.033-0.066 x 0.038-0.066; A = 0.11-0.15 x 0.11-0.14; CS = 0.19-0.22
x 0.038-0.066; Eggs = 16-24 x 10-14 µm.
Opisthorchiidae BRAUN, 1901
Body small to elongate. Oral sucker small; pharynx present; ceca medium
to long. Acetabulum small, embedded, pre-equatorial. Testes variable in
form and position; cirrus sac absent; seminal vesicle tubular, genital pore
just preacetabular. Ovary submedian, usually pretesticular; seminal recep-
tacle present; vitellaria variable; uterus extensive. Excretory vesicle Y-
shaped. Gall bladder, bile ducts or intestine of vertebrates.
Cladocystis POCHE, 1926
Body flattened, elliptical. Oral sucker small; prepharynx and pharynx
present; esophagus short; ceca medium to long. Acetabulum small, pre-
equatorial. Testes large, indented, side by side near posterior extremity;
seminal vesicle tubular, sinuous, cirrus sac absent; genital pore preacetabu-
lar. Ovary median, trilobed; seminal receptacle postovarian; vitellaria
lateral; from seminal vesicle to testes; uterus extensive. Excretory branches

extend to oral sucker. Intestine of birds, rarely fishes.
C. intestinalis VAZ, 1931: Salminus maxillosus: Brazil. (Fig. 4-9). B = 1.5-2.5 x
0.4-0.6; OS = 0.13-0.14; Ph = 0.082 x 0.04; Egg = 33-37 x 16 x 18 µm.
Witenbergia VAZ, 1932
Body elongated, expanded posteriorly. Oral sucker cap-shaped; prephar-
ynx long; pharynx elongate; esophagus long; ceca short. Acetabulum
postequatorial. Testes diagonal; near posterior extremity; cirrus sac absent;
seminal vesicle long, sinuous; genital pore preacetabular. Ovary median,
pretesticular, seminal receptacle preovarian; vitelline follicles lateral, ex-
tend from acetabulum to posterior extremity; uterus extensive, eggs small.
Intestine of fish.
W.witenbergi VAZ, 1932: Pseudoplatystoma tigrium: Brazil. (Fig. 4-8). B = 2-3.3 x
0.2-0.44; OS = 0.14-0.16 x 0.12-0.13; Ph = 0.11 x 0.05; A = 0.13-0.16; Egg
= 41-45 x 17 x 18 µm.
Paramphistomata SZIDAT, 1936. (The classification of the amphistomes used here follows
that of SEY, 1991).
Cladorchiidae SOUTHWELL & KIRSHNER, 1932
Body elongate, oval, pear shaped, thick, rarely flattened. Acetabulum
terminal or ventro-terminal. Pharynx muscular, with dorsolateral appendag-
es; esophagus with or without muscular bulb; ceca of variable length. Testes
two, inter or extracecal; cirrus sac present. Ovary post-testicular; genital pore
near intestinal bifurcation; uterine coils mainly post-testicular; vitellaria
follicular or compact, lateral. Eggs large, with or without miracidia. Intestinal
parasites of fish, amphibians, reptiles and rarely mammals.
Alphamphistoma THATCHER & JÉGU, 1996
Body elongate, narrow, deeply concave ventrally, with lateral margins folded
in. Pharynx small, with external diverticula; esophagus without muscular
bulb; ceca slender, shorter than body. Acetabulum subterminal, without
posterior indentation on rim. Testes lobate, tandem, in anterior half of
body; cirrus sac in anterior one-third of body; genital pore immediately
post-bifurcal. Ovary spherical, on mid-line, in posterior one-third of body;
vitellaria follicular, anterolateral to ovary, not reaching level of posterior
testis; uterus intercecal, eggs numerous. Intestinal parasites of freshwater
fish. Type species: Alphamphistoma canoeforma THATCHER & JÉGU, 1996.
A. canoeforma THATCHER & JÉGU, 1996: Mylesinus paraschomburgkii: Uatumã,
Pitinga, Capucapu, Trombetas, Jari and Araguari Rivers, Amazonas and
Pará States, Brazil. (Figs. 4-61 A-B & 4-88). B = 3.4-5.9 x 1.5-2.9; A = 0.67-
1.2 x 0.71-1.2; Ph = 0.26-0.52 x 0.23-0.40; CS = 0.26-0.51 x 0.10-0.33;
Eggs = 103 -129 x 51-69 µm.
Anavilhanatrema THATCHER, 1992
Body large, cylindrical, slightly flattened and with a collar-like anterior
expansion. Pharynx small, terminal, with external diverticula; esophagus
long, without muscular bulb; ceca of moderate diameter, terminating
near acetabulum. Acetabulum small to medium size, subterminal. Testes

large, lobate, tandem, pre-equatorial; cirrus sac small, ovoid, containing
sinuous tubular seminal vesicle; genital pore at level of bifurcation.
Ovary lobate, lateral to midline, near acetabulum; vitallaria follicular,
dorso-lateral to ceca, extensive, from cecal ends to posterior testis;
uterus largely intercecal, uterine seminal receptacle present near ovary;
eggs numerous. Circulatory sysem present. Excretory vesicle saccular;
pore dorsal. Intestinal parasites of freshwater fish. Type species: Anavil-
hanatrema robuusta THATCHER, 1992.
A. robusta THATCHER, 1992: Pristobrycon striolatus: Anavilhanas Islands, Rio
Negro, Amazonas State, Brazil. (Figs. 4-86 & 4-97). B = 4.6-12.0 x 1.9-3.7;
A = 0.71-1.6 long and 0.82-1.5 wide; Ph = 0.65 x 0.62; CS = 0.63 x 0.45;
Eggs = 84-112 x 42-70 µm.
Annelamphistoma THATCHER, SEY & JÉGU, 1996:
Body subcylindrical, flattened dorsally, convex ventrally, tapering anteriorly,
with five pairs of muscular puckers dorsally and one lateral expansion on
either side of acetabulum; tegument smooth dorsally and papillate ventrally.
Acetabulum hemispherical, subterminal. Pharynx with large external diver-
ticula, mouth terminal; esophagus short, bulb small; ceca long, slender.
Testes two, large, weakly lobate, inter and extracecal, tandem, in middle third
of body; cirrus sac large, ovoid; genital atrium and small genital sucker
present; genital pore median, immediately postbifurcal. Ovary spherical to
ovoid, submedian, near level of cecal ends; vitellaria chain-like, anterolateral
to ovary and ventral to ceca, extending anteriorly to near posterior testis;
uterus largely intercecal, eggs large. Lymphatic system present. Excretory
vesicle saccular, pore dorsal. Intestinal parasites of freshwater fish. Type
species: Annelamphistoma elegans THATCHER, SEY & JÉGU, 1996.
A. elegans THATCHER, SEY & JÉGU, 1996: Myleus (Myloplus)rubripinnis and M. (M.)
asterias. Jatapu River, Amazonas State & Guaporé River, Rondonia State,
Brazil. (Figs. 4-67 A-C, 4-77, 4-79 & 4-89). B = 2.5-3.1 x 0.75-1.3; A =
0.36-0.53 x 0.42-0.50; Ph = 0.31-0.56 x 0.18-0.22; CS = 0.14-0.19 x 0.08-
0.17; Eggs = 116-138 x 50-60 µm.
Betamphistoma THATCHER & JÉGU, 1996
Body small, flattened, tapered anteriorly. Acetabulum terminal, with
posterior indentation on inner rim. Pharynx with external diverticula;
esophagus without bulb; ceca slender, shorter than body. Testes lobate,
parallel, equatorial ; cirrus sac with heavy muscular walls; genital pore
postbifurcal; small spherical genital atrium present. Ovary small, subspher-
ical, on mid-liine between cecal ends; vitellaria of few small follicles, lateral
to ovary; uterus intercecal, eggs numerous. Intestinal parasites of freshwa-
ter fish. Type species: Betamphistoma jariense THATCHER & JÉGU, 1996.
B. jariense THATCHER & JÉGU, 1996: Mylesinus paraschomburgkii: Jari River, Pará State,
Brazil. (Fig. 4-55). B = 2.8-4.1 x 1.1-1.6; A = 0.34-0.57 x 0.60-0.76; Ph = 0.26-
0.34 x 0.16-0.24; CS =0.22-0.37 x 0.15-0.38; Eggs = 85-103 x 30-40 µm.
Colocladorchis THATCHER, 1979

Body slightly flattened, tapered anteriorly, rounded posteriorly; tegument
unspined. Oral sucker terminal, spherical, with small internal diverticula;
esophagus long, with muscular bulb; ceca thick, short, terminate near
equator. Acetabulum large, postero-ventral. Testes weakly lobate, parallel,
ventro-lateral to ceca in midbody; cirrus sac small, median, genital pore
median, near bifurcation. Ovary spherical, median, between testes, vitelline
gland of two irregular masses, postero-dorsal to testes; uterus occupying
space between ovary and acetabulum, contains eggs proximally and
miracidia distally (ovoviviparous). Circulatory system present. Excretory
vesicle sacular, pore dorsal. Intestine of freshwater fish.
C. ventrastomis THATCHER, 1979: Prochilodus reticulatus: Colombia. (Fig. 4-40). B
= 0.71-1.53 x 0.40-0.89; OS = 0.1-0.19; A = 0.30-0.58; CS = 0.072-0.12 x
0.06-0.096; Egg = 60-66 x 36-49 µm; Miracidium = 85-142 x 53-95 µm.
Curimatrema THATCHER, 2000.
Body small, pyriform, with anterior expansion and large subterminal
acetabulum. Pharynx large, with prominent external diverticula. Esopha-
gus short; ceca reaching acetabulum. Testes spherical, diagonal, equatorial
or pre-equatorial; cirrus sac absent; large genital sucker present near
intestinal bifurcation. Ovary spherical, post-testicular, lateral to mid-line;
vitelline glands comprising few follicles on either side near cecal ends; eggs
large, few in number. Intestinal parasites of freshwater fishes. Type
species: Curimatrema microscopica THATCHER, 2000.
C. microscopica THATCHER, 2000: Curimata vittata: Porto Novo, Rondônia State,
Brazil. (Fig. 4-64). B = 0.61-0.77 x 0.35-0.39; A = 0.24-0.26 in diameter; Ph
= 0.15-0.18 x 0.11 -0.17; Eggs = 88-120 x 44-77.
Dadayius FUKUI, 1929
Body elongate, conical. Acetabulum massive, ventroterminal, with trans-
verse tegumental ridges on interior bottom surface. Pharynx with external
diverticula; mouth terminal; esophagus slender, with bulb; ceca reaching
acetabulum. Testes rounded to oval, not lobate; cirrus sac absent; genital
sucker present; genital pore at level of bifurcation. Ovary subspherical,
anterior or dorsal to acetabulum; vitelline follicles lateral, between ovary
and posterior testis; uterus intercecal; eggs large. Intestinal parasites of
South American freshwater fishes. Type species: Dadayius marenzelleri
(DADAY, 1907) FUKUI, 1929.
D. marenzelleri (DADAY, 1907): Mylossoma sp.: Brazil. (Fig. 4-28). B = 4-6 x 2-3;
OS = 0.5-0.8 x 0.4-0.60; Diverticula = 0.25-0.30; A = 1.2-2; E = 0.7; EB =
0.22; GS 0.37; Egg = 140-145 x 60-80 µm.
D. pacuensis THATCHER, SEY & JÉGU, 1996: Myleus (Myloplus) rubripinnis and M.
(M.) asterias: Jatapu River, Amazonas State & Guaporé River, Rondonia
State, Brazil. (Figs. 4-65 A-B, 4-73, 4-76 & 4-84). B = 1.5-3.0 x 1.1-1.9; A =
1.1-1.4 x 1.0-1.5; Ph = 0.30-0.60 x 0.34-0.43; GS = 0.31-0.43 x 0.34-0.69;
Eggs 69-86 x 26-43 µm.
Dadaytrema TRAVASSOS, 1931, THATCHER, 1979.

Body elongate, cylindrical. Oral sucker pyriform, with posterior diverticu-
la; mouth terminal, surrounded by small papillae and with several rings of
papillae posterior to mouth; esophagus long, with small bulb; ceca long
sinuous. Acetabulum subterminal, with papillate floor and posteromedian
notch in rim. Testes deeply lobed, pre-equatorial, intercecal, but invading
cecal areas, genital sac containing seminal vesicle; cirrus absent; genital
pore median, post-bifurcal, contained in shallow sucker. Ovary spherical,
median, near acetabulum; vitelline follicles small, few, lateral to ovary and
dorsal to cecal ends; uterus intercecal, filling space between ovary and
testes, with eggs proximally and miracidia distally (ovoviviparous). Circula-
tory system of longitudinal tubes with sinuses near suckers. Excretory
vesicle saccular, pore dorsal. Intestine of freshwater fish.
D. elongata VAZ, 1932: Myleus sp.: Brazil. B = 3-4.5 x 0.95-1.7; OS = 0.1x 0.7;
Diverticula = 0.2-0.35 x 0.1; A = 0.5-0.7; Egg = 115-153 x 49-70 µm.
D. oxycephala (DIESING, 1836) TRAVASSOS, 1931, THATCHER, 1979: Colossoma
bidens: Brazil (Fig. 4-37). B = 3.0-5.0 x 1.1-1.3; OS = 0.15-0.25 x 0.09-0.18;
Diverticula = 0.18-0.22 x 0.14-0.18; A = 0.56-0.90; GS = 0.18-0.40 x 0.11-
0.27; Egg = 90 x 60 µm. Miracidium = 140-190 x 85-96 µm.
Dadaytremoides THATCHER, 1979
Body flattened, widest at equator, tapering towards extremities, unspined.
Oral sucker large, rounded, with large diverticula; esophagus long, bulb
prominent; ceca thick, long. Acetabulum subterminal. Testes weakly
lobate, diagonal, in midbody; genital pore postbifurcal, with small sucker.
Ovary spherical, median, between cecal ends; vitelline follicles few, dorsal
to cecal terminations; uterus filling space between ovary and testes; with
eggs proximally and miracidia distally (ovoviviparous). Circulatory system
with longitudinal stems and sinuses near suckers. Excretory vesicle
saccular, pore dorsal. Intestine of freshwater fish.
D. grandistomis THATCHER, 1979: Astyanax fasciatus and Chaetostomus leucomelas:
Colombia: (Fig. 4-38). B = 1.9-4.0 x 0.75-1.3; OS = 0.22-0.64; Diverticula
= 0.14-0.22 x 0.09-0.14; A = 0.35-0.54; GS = 0.14-0.23 x 0.09-0.16; Egg =
72-90 x 36-45 µm; Miracidia = 95-110 x 41-62 µm.
Deltamphistoma THATCHER & JÉGU, 1996
Body cylindrical, slightly flattened. Acetabulum small, subterminal. Phar-
ynx with external diverticula; esophagus with muscular bulb; ceca shorter
than body, usually not reaching ovary. Testes large, overlapping ceca,
bluntly lobate, tandem, pre-equatorial; cirrus thin-walled; genital pore
post-bifurcal, small atrium present. Ovary subspherical, lateral to mid-line,
in posterior one-fourth of body; vitelline follicles on either side of body,
near ceca, extend from ovary to posterior testis; uterus intercecal, eggs
numerous. Intestinal parasites of freshwater fish. Type species: Deltamphis-
toma pitingaense THATCHER & JÉGU, 1996.
D. pitingaense THATCHER & JÉGU, 1996: Mylesinus paraschomburgkii: Uatumã,

Pitinga and Capucapu Rivers, Amazonas State, Brazil. (Fig. 4-56). B = 3.1-
4.1 x 1.0-1.5; A = 0.43-0.57 x 0.52-0.60; Ph = 0.26-0.34 x 0.21-0.26; CS =
0.22-0.39 x 0.13-0.34; Eggs = 95-120 x 34-52 µm.
Doradamphistoma THATCHER, 1999
Body elongate, flattened, sides nearly parallel, extremities bluntly rounded.
pharynx large, with prominent external diverticula; esophagus long, bulb
present; ceca reaching to near acetabulum. Acetabulum small, subterminal.
Testes diagonal, equatorial or pre-equatorial; cirrus sac small, subspherical;
genital pore postbifurcal. Ovary ovoid, postequatorial; vitellaria consisting
of few large follicles lateral to ovary and extracecal; uterus extensive, with
ascending and descending loops; eggs numerous; ovoviviparous. Excretory
vesicle saccular, pore dorsal. Intestinal parasites of Amazonian catfishes.
Type species: Doradamphistoma bacuensis THATCHER, 1999.
D. bacuensis THATCHER, 1999: Megalodoras irwini: Amazon River, near Manaus,
Amazonas State, Brazil. (Fig. 4-81 & 4-99). B = 6.0-9.6 x 1.8-3.1; Ph = 0.83-
1.3 x 0.62-0.83; CS = 0.20-0.26 x 0.20-0.26; Eggs =132-191 x 88-110 µm.
Gamamphistoma THATCHER & JÉGU, 1996
Body small, little flattened, with collar-like expansion anteriorly. Pharynx
with external diverticula; esophagus short, with muscular bulb; ceca long.
Acetabulum large, with posterior notch on inner rim. Testes large, lobate,
tandem, pre-equatorial; cirrus sac ovoid, thin-walled; genital pore at level
of intestinal bifurcation, small genital atrium present. Ovary spherical, on
mid-line near acetabulum; vitellaria follicular, lateral to ovary, of limited
extent; uterus intercecal. Eggs numerous. Intestinal parasites of freshwater
fish. Type species: Gamamphistoma collaris THATCHER & JÉGU, 1996.
G. collaris THATCHER & JÉGU, 1996: Mylesinus paraschomburgkii: Uatumã, Pitinga
and Capucapu Rivers, Amazonas State & Jari River, Pará State, Brazil. (Fig.
4-57). B = 1.2-2.2 x 0.6-0.9 ; A = 0.39-0.58 x 0.48-0.60; Ph = 0.19-0.26 x
0.14 -0.24; CS = 0.13-0.22 x 0.08-0.17; Eggs = 97-127 x 46-92 µm.
Inpamphistoma THATCHER, SEY & JÉGU, 1996
Body elongate, flattened, tapering anteriorly, concave dorsally and convex
ventrally, with one pair of muscular puckers near oral sucker and one
lateral expansion on either side of acetabulum; tegument smooth dorsally
and papillate ventrally. Acetabulum hemispherical, subterminal. Pharynx
terminal, with external diverticula; esophagus short, bulb small; ceca long,
slender. Testes two, large lobate, tandem, largely intercecal, in middle third
of body; cirrus sac ovoid; small genital atrium present; genital pore
median, immediately postbifurcal. Ovary ovoid or weakly lobate, submedi-
an, near level of cecal ends; vitellaria follicular, of limited extent, antero-
lateral to ovary and ventral to ceca; uterus largely intercecal, proximal eggs
small, distal eggs larger. Lymphatic system present. Excretory vesicle
saccular, pore dorsal. Intestinal parasites of freshwater fish. Type species:
Inpamphistoma papillatum THATCHER, SEY & JÉGU, 1996.
I. papillatum THATCHER, SEY & JÉGU, 1996: Myleus (Myloplus) asterias: Jatapu and
Uatumã Rivers, Amazonas State and Guaporé River, Rondônia State,
Brazil. (Fig. 4-66). B = 1.8-3.8 x 0.85-1.3; A = 0.43-0.81 x 0.46-0.81; Ph =
0.18-0.29 x 0.11-0.21; CS = 0.08-0.26 x 0.03-0.11; Proximal eggs = 55-94 x
33-44 µm; Distal eggs = 121-154 x 66-94.
Kalitrema TRAVASSOS, 1933
Body flattened, unspined, with circular groove at level of esophagus, and
deep indentation at posterior extremity. Oral sucker large, with internal
diverticula; esophagus long, without bulb; ceca medium long. Acetabulum
small, in posterior indentation. Testes small, weakly lobate, parallel, extrace-
cal, in anterior third; cirrus sac absent; small genital sucker present, posterior
to tegumental groove. Ovary small, spherical, median, posterior to cecal
ends; vitelline follicles few, near cecal termination; uterus intercecal, from
genital sucker to beyond ovary; with eggs proximally and miracidia distally
(ovoviviparous). Excretory vesicle spherical, pore dorsal. Intestine of fish.
K. kalitrema TRAVASSOS, 1933: Hypostomus punctatus: Brazil. (Fig. 4-39). B = 7-
8.7 x 2.6-3; OS = 0.36-0.39; Diverticula = 0.13; A = 0.76-0.78; Miracidia =
160-168 x 96-112 µm.
Micramphistoma THATCHER, 1992
Body medium sized, somewhat flattened, with parallel sides, dorsally concave.
Pharynx small, subspherical, without diverticula; esophagus long, without
muscular bulb; ceca slender, medium long. Acetabulum, small, subspherical,
subterminal. Testes spherical to ovoid, nearly parallel, near intestinal bifurca-
tion; male genital sac subovoid, between testes; genital pore immediately
postbifurcal. Ovary spherical, near acetabulum, lateral to midline; vitellaria
consisting of few large follicles, dorso-lateral to ceca, in middle third of body;
uterus largely intercecal; eggs small numerous. Circulatory system present.
Excretory vesicle saccular, pore dorsal. Intestinal parasites of freshwater fish.
Type species: Micramphistoma ministoma THATCHER, 1992.
M.ministoma THATCHER, 1992: Hypopomus sp.: Guaporé River, Rondônia State,
Brazil. (Figs. 4-80 & 4-95). B = 4.5-5.6 x 2.0-2..4; A = 0.50-0.59 x 0.52-
0.60; Ph = 0.43-0.56 x 0.42-0.49; Male genital sac = 0.21-0.38 x 0.27-0.38;
Eggs = 88-110 x 50-60 µm.
Myleustrema THATCHER & JÉGU, 1998
Body elongate, flattened, concave dorsally and convex ventrally, with
circlets of papillae around anterior extremity. Pharynx small, terminal,
with external diverticula; esophagus long, with small bulb; ceca slender,
not reaching acetabulum. Acetabulum small, terminal, with circle of
papillae around aperture. Testes weakly lobate, tandem, equatorial; cirrus
sac ovoid, with heavy muscular walls, contains internal seminal vesicle;
genital pore postbifurcal; small genital atrium present. Ovary subspherical,
near cecal ends in posterior quarter of body; vitellaria of limited extent,
anterolateral to ovary; uterus intercecal; eggs few. Intestinal parasites of
fish. Type species: Myleusnema concavatum THATCHER & JÉGU, 1998.
M.concavatum THATCHER & JÉGU, 1998; Myleus ternetzi: French Guiana (Fig. 4-
59). B = 2.8-3.2 x 1.0-1.2; Ph = 241-258 x 146-181; A = 172-301 long and
344-430 wide; Eggs = 115-138 x 51-64 µm.
Pacudistoma THATCHER,1992
Body large, cylindrical, little flattened. Pharynx large, terminal, with
prominent internal diverticula; esophagus long, muscular, without bulb,
cec of medium diameter, long. Acetabulum medium sized, subtermi-
nal. Testes large, lobate, tandem, pre-equatorial; cirrus sac small, ovoid,
containing saccular seminal vesicle; genital pore immediately postbifur-
cal, provided with large, muscular genital sucker. Ovary small, spheri-
cal, near acetabulum; vitellaria of small follicles, ventral to ceca,
between ovary and posterior testis; uterus largely intercecal, uterine
seminal receptacle present. Exccretory vesicle saccular, pore dorsal.
Intestinal parasites of freshwater fishes. Type species: Pacudistoma
turgida THATCHER, 1992.
P. turgida THATCHER, 1992; Myleus pacu: Jamari River, Rondônia State, Brazil
(Fig. 4-82). B = 7.1-10.1 x 3.1-3.9; Ph = 1.1-1.4 wide; A = 1.5-1.9 long x
1.4-1.7 wide; Eggs = 98-112 x 56-70 µm.
P. guianensis THATCHER & JÉGU, 1998; Myleus ternetzi: Sinnamary River, French
Guiana (Fig. 4-60). B = 2.8-3.2 x 1.0-1.1; Ph = 241-258 long x 146-181
wide; A = 172-301 long x 344-430 wide; Eggs = 115-138 x 51-64 µm.
Pronamphistoma THATCHER, 1992
Body small, stout, with anterior collar-like expansion. Acetabulum large,
round, subterminal. Pharynx large, terminal, subspherical, with internal
diverticula; esophagus short, bulb ovoid; ceca thick, long, reaching acetab-
ulum. Testes subspherical, entire, in middle third of body; cirrus sac ovoid,
containing saccular seminal vesicle; genital pore postbifurcal. Ovary
spherical, submedian, near acetabulum; vitellaria consisting of follicles in
linear rows that form fan-like configurations on each side of ovary; uterus
short, with few large eggs. Circulatory system present. Excretory vesicle
saccular, pore dorsal. Intestinal parasites of freshwater fishes. Type
species: Pronamphistoma cichlasomae THATCHER, 1992.
P. cichlasomae THATCHER, 1992: Cichlasoma severum: Guaporé River, Rondônia
State, Brazil. (Figs. 4-85 & 4-92). B = 0.69-1.12 x 0.42-0.76; A = 0.26-0.45
x 0.28-0.49; Ph = 55-82 µm in diameter; CS = 55-112 µm in diameter;
Eggs = 82-110 x 45-65 µm.
Pseudocladorchis DADAY, 1907
Body elongate, elliptical. Oral sucker terminal, with papillae around
mouth; esophagus without bulb; ceca long. Acetabulum large, ventro-
terminal, with median notch on posterior inner margin of rim. Testes
nearly parallel, in midbody; cirrus sac present; genital pore near bifurca-
tion. Ovary between cecal ends; vitelline follicles extending along ceca
from testes; uterus intercecal. Excretory vesicle between ovary and
acetabulum, pore dorsal. Intestine of freshwater fish.
P. cylindricus (DIESING, 1836) DADAY, 1907: Mylossoma aureum. Pterodoras granulo-

sus, Colossoma bidens and Pimelodus ornatus: Brazil. (Fig. 4-30). B = 4-10 x 1.5-
3.2; OS = 0.6-1.2 x 0.6-1.3; A = 0.5-2.5; Egg = 90-100 x 60-80 µm.
P. ferrumequinum (DIESING, 1836) STUNKARD, 1925: Pterodoras granulosus and
Doras dorsalis: Brazil. B = 5.5-11.5 x 3.2-5.4; OS = 1.6-2 x 1-1.6; Diverticula
= 0.35-0.45; Egg = 80 x 50 µm.
Pseudodiplodiscus MANTER, 1962
Body small, conical. Oral sucker terminal, with small diverticula; esopha-
gus with bulb; ceca short. Acetabulum large, ventro-terminal. Testis single,
in midbody; cirrus sac small; genital sucker small, near bifurcation. Ovary
irregular, submedian, preacetabular; vitellaria lateral, from cecal ends to
ovary; uterus winding between testis and acetabulum. Excretory vesicle
saccular, pore dorsal. Intestine of freshwater fish.
P. cornu (DIESING, 1939) MANTER, 1962: Doras dorsalis: Brazil. (Fig. 4-29). B =
4-4.3 x 2-2.2; OS = 0.25-0.40; A = 2-2.2; Egg = 130-140 x 60-70 µm.
Travassosinia VAZ, 1932
Body cylindrical. Oral sucker terminal, with papillae around mouth;
esophageal bulb small; ceca medium long. Acetabulum large, ventro-
terminal. Testes tandem, in midbody; genital pore median, postbifurcal.
Ovary median, post-testicular; vitelline follicles lateral along post-testicular
portions of ceca; uterus inter- and extracecal. Excretory pore dorsal.
Intestine of freshwater fish.
T. dilatata (DADAY, 1907) VAZ, 1932: Colossoma bidens and Myleus sp.: Brazil.
(Fig. 4 -27). B = 8-10 x 4-5; OS = 1.2-1.3; Egg = 115-150 x 57-80 µm.
Zetamphistoma THATCHER & JÉGU, 1996
Body small, wide, flattened, convex on both surfaces, thinner peripherally.
Pharynx with external diverticula; esophagus slender, with small bulb; ceca
shorter than body. Acetabulum terminal, with indentation on posterior
inner rim. Testes lobate, tandem, pre-equatorial, intercecal; cirrus sac
ovoid, with thin walls; genital pore post-bifurcal. Ovary subspherical, on
mid-line, near acetabulum; vitellaria of few follicles on either side of
ovary; uterus intercecal, eggs numerous, small proximally, larger distally.
Intestinal parasites of freshwater fish. Type species: Zetamphistoma compacta
THATCHER & JÉGU, 1996.
Z. compacta THATCHER & JÉGU, 1996: Mylesinus paraschomburgkii: Trombetas River,
Pará State, and Araguari River, Amapá State, Brazil. (Figs. 4-63 A-B & 4-83). B
= 2.5-3.9 x 1.5-2.5; A = 0.48-0.64 x 0.61-0.94; Ph = 0.33-0.37 x 0.16-0.26; CS
= 0.13-0.34 x 0.086-0.22; Eggs = 92-115 x 46-58 µm.
Sanguinicolidae GRAFF, 1907
Body flattened, elongate or lanceolate. Oral sucker absent or rudimentary;
pharynx absent; esophagus long; gut X or H shaped. Acetabulum absent.
Testes single, double, branched or follicular; between ovary and bifurca-
tion; cirrus sac present or absent; seminal vesicle usually present; genital
pores dorsal, posterior, submarginal; male pore posterior to female. Ovary
median, lobed or two-winged; near posterior extremity; vitellaria extensive;

uterus limited; eggs not operculate. Blood-vascular system of fish.
Sanguinicola PLEHN, 1905
Body lanceolate, with marginal striations. Oral sucker absent; esophagus
long; gut X or H shaped, sometimes with five branches. Acetabulum absent.
Testes in two rows, in medium field, between ovary and intestine; cirrus sac
present; genital pores near posterior extremity. Ovary of two symmetrical
wings, in posterior half of body; vitellaria lateral; uterus with only one egg;
egg with lateral projections. Vascular system of freshwater fish.
S. argentinensis, SZIDAT, 1951; Prochilodus platensis: Argentina. B = 1.6-1.7 x 0.37;
Ph = 0.02 x 0.015; E = 0.45; Egg = 37 x 31 µm.
S. coelomicola (SZIDAT, 1951): Iheringichthys labrosus: Argentina. (Fig. 4-32). B =
1.6 x 0. 55; E = 0.5-0.6; Egg = 11-12 x 6-7 µm.
Zonocotylidae PADILHA, 1978
Body elongate, widened posteriorly. Oral sucker terminal; esophagus long;
ceca short, saccular. Acetabulum subterminal, with thin walls and trans-
verse corrugations on floor. Testis single, median, postequatorial; cirrus
sac absent; genital pore median, midway between oral sucker and bifurca-
tion. Ovary spherical or oval, submedian, between testis and acetabulum;
vitellaria of two compact masses, one on either side of testis or ovary;
uterine coils extensive posteriorly, with eggs proximally and miracidia
distally (ovoviviparous). Circulatory system well developed. Excretory
vesicle saccular, pore dorsal. Intestine of freshwater fish.
Zonocotyle TRAVASSOS, 1948, PADILHA, 1978
Body elongate, flattened, widened posteriorly; tegument smooth, thick.
Oral sucker terminal; esophagus long, dilated posteriorly, without bulb;
ceca short, thick, with microvilli on interior walls. Testis irregularly
rounded, median, between cecal ends. Ovary spherical submedian, post-
testicular; vitelline glands of two compact masses, one on either side of
ovary; uterus extensive, with eggs proximally and miracidia distally
(ovoviviparous). Circulatory system of one continuous tube which loops
laterally on both sides, between suckers. Excretory vesicle small, spherical,
pore and vesicle both dorsal to acetabulum. Intestine of freshwater fish.
Z. bicaecata TRAVASSOS, 1948, PADILHA, 1978: Curimata elegans and C. gilberti:
Brazil. (Fig. 4-41). B = 2.65-5.23 x 0.73-1.38; OS = 0.19-0.48 x 0.31-0.55; E
= 0.70-1.56; A = 0.99-1.8 x 0.8-1.7 (with 11-14 ridges on floor);
Miracidium = 140-210 x 50-120 µm.
Zonocotyloides PADILHA, 1978
Similar to Zonocotyle except: body larger; acetabulum relatively smaller;
vitellaria of two compact V-shaped masses, one on either side of testis and
united anterior to that organ. Ovoviviparous. Intestine of freshwater fish.
Z. haroltravassossi PADILHA, 1978: Curimata gilberti: Brazil. (Fig. 4-42). B = 4.75-
7.4 x 1.6-2.1; OS = 0.36-0.50 x 0.47-0.60; E = 1.4-2.3; A = 1.2- 1.87 x 1.3-
1.56 (with 9-10 ridges on floor).
VIII. Plates of Trematoda (Digenea) (Figs. 4-1 to 4-99)
MO
CS
OS
PP
0.5 mm
PH
IC
GP
VS
UE
SV
VG
OV
SR
TE
4-1 AP
EP
4-1. Acanthostomum gnerii: entire (ventral; AP = anal pore; CS = circumoral spine; EP = excretory
pore; GP = genital pore; IC = intestinal crus; MO = mouth; OS = oral sucker; OV = ovary; PH =
pharynx; PP = prepharynx; SR = seminal receptacle; SV = seminal vesicle; TE = testis; UE = uterus
with eggs; VG = vitelline gland; VS = ventral sucker (= acetabulum).
OS
GS
SV
1.0 mm
VS
MG
OV
VG
TE
UE
IC
TR
EB
4-2
EP
4-2. Amazonadistoma negrensis: entire (ventral); EB = excretory bladder; EP = excretory pore; GS =

genital sucker; IC = intestinal crus with diverticula and microvilli; MG = MEHLIS’ gland; OS = oral
sucker; OV = ovary; SV = seminal vesicle; TE = testis; TR = tegumental ridge; UE = uterus with
eggs; VG = vitelline gland; VS = ventral sucker (= acetabulum).
SR
OV
1.5 mm
TE
UE
VG
4-3
4-3. Brasicystis bennetti: entire (ventral); OV = ovary; SR = seminal receptacle; TE = testis; UE =

uterus with eggs; VG = vitelline gland.
4-4
4-5
4-6
4-4. Bellumcorpus major; 4-5. Paurorhynchus schubarti; 4-6. Prosorhynchus costai.

4-7
4-8
4-7. Prosthenhystera obesa; 4-8. Witenbergia witenbergia.

4-9
4-10
4-12
4-11
4-9. Cladocystis intestinalis; 4-10. Crepidostomum macrorchis; 4-11. Creptotrema creptotrema; 4-12. Eocreadium
internedium.
4-13
4-14
4-13. Antorchis lintoni; 4-14. Procaudotestis uruguayensis.

4-15
4-16
4-17
4-15. Caballerotrema arapaimense; 4-16. Kalipharynx piramboae; 4-17. Magnivitellium simplex.

4-18
4-18. Crassicutis cichlasomae.

4-19
4-20
4-21
4-19. Dendrorchis neivai; 4-20. Parspina argentinensis; 4-21. Halipegus dubius.

4-22
4-23
4-22. Iheringtrema iheringi; 4-23. Paleocryptogonimus claviformis.

4-24 A 4-24 B
4-24. Megacoelium spinicavum: A. ventral; B. lateral.

4-25
4-25. Phyllodistomum duncani.

4-26
4-26. Chalcinotrema lucieni.

4-27 4-28
4-29 4-30
4-27. Travassosinia dilatata; 4-28. Dadayius marenzelleri; 4-29. Pseudodiplodiscus cornu; 4-30. Pseudocladorchis
cylindricus.
4-31 4-32
4-31. Lecithobotrioides mediacanoensis; 4-32. Sanguinicola coelomicola.

4-33
4-33. Chalcinotrema ruedasueltensis.

4-34
4-34. Saccocoelioides leporinodus.

4-35
4-35. Saccocoelioides saccodontis.

4-36
4-36. Paralecithobotrys brasiliensis.

4-37 4-38
4-37. Dadaytrema oxycephala; 4-38. Dadaytremoides grandistomis.

4-39
4-39. Kalitrema kalitrema (scale = 3 mm).

4-40
4-40. Colocladorchis ventrastomis.

4-41
4-42
4-41. Zonocotyle bicaecata; 4-42. Zonocotyloides haroltravassosi.

4-43
4-43. Saccocoelioides magnorchis.

4-44
4-45
4-46
4-44. Curumai curumai; 4-45. Denticauda quadrangulata; 4-46. Pseudoparabaris parabaris.

4-47 4-48
4-49 4-50
4-47. Brasicystis bennetti: photo of entire specimen; 4-48. Diplostomula of Strigeoidea; 4-49.
Diplostomula: close up view; 4-50. Encysted metacercaria of Ascocotyle sp. (“black spot”).
4-51 4-52
4-53 4-54
4-51 to 4-54. Tumors in Chaetobranchus semifasciatus provoked by trematode metacercariae: 4-51. large
tumor and satellite tumor in gills (operculum removed); 4-52. simple tumor in gill filament with
metacercaria in center; 4-53. swollen gill filament containing metacercaria; 4-54. multiple branching
of gill filament.
4-55
4-56
4-57
4-58
4-55. Betamphistoma jariense (scale = 1000 µm); 4-56. Deltamphistoma pitangaense (scale = 1000 µm); 4-
57. Gamadistoma collaris (scale = 1000 µm); 4-58. Saccocoelioides rotundus (scale = 250 µm).
4-59 A
4-59. Myleustrema concavatum: A. ventral (scale = 1000 µm); B. genital complex (scale = 100 µm).
4-60 A
4-60. Pacudistoma guianensis from Myleus ternetzi. A-C growth series all to same scale = 1000 µm.
4-61 A
B
4-63 A
4-62
4-61. A. Alphamphistoma canoeforma (scale = 1000 µm); B. same - terminal genitalia (scale = 250 µm);
4-62. A. Pseudocladorchis cylindricus (scale 1000 µm); B. same - genitalia lateral (scale = 500 µm); 4-63.
Zetamphistoma compacta (scale = 1000 µm).
4-64
4-64. Curimatrema microscopica (scale = 500 µm).

4-65 A
4-65. Dadayius pacuensis: A. ventral (scale = 1000 µm); B. lateral (scale = 1000 µm).
4-66
4-66. Inpamphistoma papillatum (scale = 500 µm).

4-67 A
4-67. Annelamphistoma elegans: A. ventral (scale = 500 µm); B. dorsal (scale = 1000 µm); C. lateral
(scale = 1000 µm).
4-68
4-69
4-68. Paraproctotrema delicata; 4-69. Sphericomonorchis spinulosus; (scales = 200 µm).

4-70
4-71
4-72
4-70
4-70. Genolopa magnacirrus; 4-71. Paraproctotrema delicata; 4-72. Genolopa magnacirrus; (scales = 200 µm).
4-73
4-74
4-75 4-76
Dadayius pacuensis: 4-73. ventral (scale = 1000 µm); 4-74. mouth (scale = 100 µm); 4-75. interior of
acetabulum (scale = 200 µm); 4-76. genital pore (scale = 50 µm); scanning electron micrographs.
4-78
4-77
4-79
Annelamphistoma elegans: 4-77. dorsal (scale = 500 µm); 4-78. papilla around acetabulum (scale = 3
µm); 4-79. papillae around mouth (scale = 3 µm); scanning electron micrographs.
4-80 4-81 4-82
4-83 4-84
4-80. Micramphistoma ministoma (scale = 500 µm); 4-81. Doradamphistoma bacuensis (scale = 1000 µm); 4-
82. Pacudistoma turgida (scale = 1000 µm); 4-83. Zetamphistoma compacta (scale = 1000 µm); 4-84. Dadayius
pacuensis (scale = 500 µm).
4-85 4-87
4-86
4-88 4-89
4-85. Pronamphistoma cichlasomae (scale = 1000 µm); 4-86. Anavilhanatrema robusta (scale = 500 µm); 4-
87. Dadaytrema oxycephala (scale = 1000 µm); 4- 88. Alphamphistoma canoeforma (scale = 1000 µm); 4-89.
Annelamphistoma elegans (scale = 1000 µm).
4-90
4-90. Prosorhynchus piranhas (scale = 250 µm).

4-91 4-92
4-93
4-94
Bucephalidae: 4-91 to 4-92. Glandulorhynchus turgidus (scale = 1000 µm); Haploporidae: 4-93. Rondotrema
microvitellarum (scale = 250 µm); 4-94. Lecithobotrioides elongatus (scale = 500 µm).
4-95
4-95. Micramphistoma ministoma (scale = 1000 µm).

4-96
4-96 Pronampistoma cichlasomae: ventral (scale = 250 µm).

4-97
4-97. Annavilhanatrema robusta (scale = 2000 µm).

4-98
4-98. Bacciger pellonae (scale = 200 µm).

4-99
4-99. Doradamphistoma bacuensis; intestine of Megalodoras irwini (scale = 1 mm).

ASHTON, N., BROWN, N. & D. EASTY (1969): Trematode cataract in freshwater fish. - J. Small Anim.
Pract. 10: 471-478.
BOEGER, W.A. & V.E. THATCHER (1983): Kalipharynx piramboae gen. et sp. n. (Trematoda:
Fellodistomidae) parasita do peixe pulmonado amazônico, Lepidosiren paradoxa FITZINGER. -
Acta Amazonica 13(1): 171-175.
BROOKS, D.R. (1977): A new genus and two new species of trematodes from characid fishes in
Colombia. - Trans. Am. Mic. Soc. 96(2): 267-270.
DAWES, B. (1946): The Trematoda with special reference to British and other European forms. -
Cambridge Univ. Press, London: 644 pp.
FREEMAN, R.S. (1964): Flatworm problems in fish. - Can. Fish Cult. 32: 11-18.
FREEMAN, R.S., STUART, P.F., CULLEN, J.B., RITCHIE, A.C., MILDON, A., FERNANDES, B.J. & R. BONIN
(1976): Fatal human infection with mesocercariae of the trematode Alaria americana. - Am. J.
Trop. Med. Hyg. 25: 803-807.
HALTON, D.W. (1966): Occurrence of microvilli-like structures in the gut of digenetic trematodes. -
Experientia 22: 828-829.
HALTON, D.W. (1967): Observations on the nutrition of digenetic trematodes. - Parasit. 57: 639-660.
HYMAN, L.R. (1951): The Invertebrates: Platyhelminthes and Rhynchocoela. The Acoelomata
Bilateria. Vol 2. - McGraw-Hill, New York: 550 pp.
KOHN, A. (1962): Sôbre um novo gênero de trematódeo bucefaliforme parasito de peixe de água
doce. - Rev. Brasil. Biol. 22(4): 351-355.
KOHN, A. (1963): Paurorhynchus schubarti sp. n., metacercária parasita de dourado (Trematoda,
Bucephaliformes). - Rev. Brasil. Biol. 23(3): 259-261.
KOHN, A. (1970a): Pararhipidocotyle jeffersoni gen. n., sp. n., trematódeo bucefaliforme parasito de
dourado. - Atas Soc. Biol. Rio de Janeiro 13(5-6): 181-183.
KOHN, A. (1970b): Bellumcorpus schubarti (KOHN, 1963) comb. nov. para Paurorhynchus schubarti
(KOHN, 1963) (Trematoda, Prosorhynchidae). - Atas Soc. Biol. Rio de Janeiro 13 (5-6):
185-186.
PADILHA, T.N. (1978): Caracterização da familia Zonocotylidae com redescrição de Zonocotyle
bicaecata TRAVASSOS, 1948, e descrição de um novo gênero (Trematoda, Digenea). - Rev. Brasil.
Biol. 38(2): 415-429.
SCHELL, S.C. (1970): How to know the trematodes. - Wm. C. Brown Co. Publ., Dubuque: 355 pp.
SKRJABIN, K.I (ed.) (1947-1974): Trematodi zhivotnikh I cheloveka (Trematodes of animals and
man) (In Russian). - Akad. Nauk. U.S.S.R., Moscow. Vols. 1-15.
SZIDAT, L. (1951): Neue Arten der Trematoden-Familie Aporocotylidae aus dem Blut und der
Leibeshöhle von Süßwasserfischen des Rio de la Plata. - Z. Parasitenkde. 15: 70-86.
SZIDAT, L. (1954): Tremátodes nuevos de peces de agua dulce de la República Argentina e intento
para aclarar su carácter marino. - Rev. Inst. Nac. Inves. Cienc. Nat. Argen. Bernadino
Rivadavia Zool. 3(1): 1-85.
TEIXEIRA DE FREITAS, J.F. (1941a): Nôvo trematódeo parasito de peixe de água doce. - Mem. Inst.
Oswaldo Cruz 35(3): 569-570.
TEIXEIRA DE FREITAS, J.F. (1941b): Nôvo trematódeo parasito de peixe do rio Miranda. - Rev. Brasil.
Biol. 1(3): 249-251.
TEIXEIRA DE FREITAS, J.F. (1947a): Chalcinotrema salobrensis n. g., n. sp., (Trematoda, Waretrematidae). -
Rev. Brasil. Biol. 7(4): 461-464.
TEIXEIRA DE FREITAS, J.F. (1947b): Nôvo gênero de Haploporinae (Trematoda, Haploporoidea). -
Mem. Inst. Oswaldo Cruz 45(3): 587-589.
THATCHER, V.E. (1978): Quatro novas espécies de Haploporidae (Trematoda: Digenea) de peixes de
água doce do Colômbia com uma revisão do gênero Saccocoelioides SZIDAT, 1954. - Acta
Amazonica 8(3): 477-484.
THATCHER, V.E. (1979a): Paramphistomidae (Trematoda: Digenea) de peixes de água doce: dois
novos gêneros da Colômbia e uma redescricão de Dadaytrema oxycephala (DIESING, 1836)
TRAVASSOS, 1934, da Amazônia. - Acta Amazonica 9(1): 203-208.
THATCHER, V.E. (1979b): O primeiro trematódeo negro, Amazonadistoma negrensis n. gen., n. sp.
(Digenea: Gorgoderidae), parasita de um peixe amazônico. - Acta Amazonica 9(2): 389-392.
THATCHER, V.E. (1979c): Brasicystis bennetti n.gen., n.sp. (Trematoda: Didymozoidae) parasita da
pescada (Sciaenidae) da Amazônia, Brasil. - Acta Amazonica 9(4): 747-749.
THATCHER, V.E. (1980): Duas novas espécies de Caballerotrema (Trematoda: Echinostomatidae) do
pirarucu e da aruanã (Osteoglossidae), com uma redefinição do gênero e una redescrição de
C. brasiliense PRUDHOE, 1960. - Acta Amazonica 10(2): 419-423.
THATCHER, V.E. (1981): Patologia de peixes da Amazônia brasileira, 1. Aspectos gerais. - Acta
Amazonica 11(1): 125-140.
THATCHER, V.E. (1992a): Bacciger pellonae n. sp. (Trematoda, Fellodistomidae) from a freshwater
fish, Pellona castelnaeana VALENCIENNES of Rondônia State, Brazil. - Acta Amazonica 22(4):
605-608.
THATCHER, V.E. (1992b): Two unusual new genera of Paramphistomidae (Trematoda, Digenea)
from freshwater fish of the Brazilian Amazon. - Acta Amazonica 22(4): 609-613.
THATCHER, V.E. (1996): A new genus and three new species of Monorchiidae (Trematoda) from
freshwater fish of Rondônia State, Brazil. - Amazoniana 14(1/2): 131-136.
THATCHER, V.E. (1999a): Two new Bucephalidae (Trematoda) from fishes of the Brazilian Amazon.
- Acta Amazonica 29(2): 331-335.
THATCHER, V.E. (1999b): Two new Haploporidae (Trematoda) of fishes from the Brazilian State of
Rondônia. - Acta Amazonica 29(4): 601-605.
THATCHER, V.E. (1999c): Surface morphology of some amphistomes (Trematoda) of Amazonian
fishes and the description of a new genus and species. - Acta Amazonica 29(4): 607-614.
THATCHER, V.E. (2000): A new genus and species of amphistome parasite (Trematoda: Cladorchi-
idae) from a fish of Rondônia State, Brazil. - Amazoniana 16(1/2): 237-240.
THATCHER, V.E. & M.D. DOSSMAN (1974): Lecithobotrioides mediacanoensis n. g., n. sp. (Trematoda:
Haploporidae) from a freshwater fish (Prochilodus reticulatus) in Colombia. - Trans. Am. Mic.
Soc. 93(2): 261-264.
THATCHER, V.E. & M.D. DOSSMAN (1975): Unicoelium prochilodorum gen. et sp. n. (Trematoda:
Haploporidae) from a freshwater fish (Prochilodus reticulatus) in Colombia. - Proc. Helminthol.
Soc. Wash. 42(1): 28-30.
THATCHER, V.E. & M. JÉGU (1996): Intestinal helminths as population markers of the Amazonian
fish Mylesinus paraschomburghii, with descriptions of five new genera and seven new species of
trematodes. - Amazoniana 14(1/2): 143-155.
THATCHER, V.E. & M. JÉGU (1998): Amphistomes as species markers of the serrasalmid fish, Myleus
ternetzi (NORMAN), from French Guiana, with descriptions of two new species and one new
genus. - Amazoniana 15(1/2): 103-112.
THATCHER, V.E. & A.B. VARELLA (1980): Patologia de peixes da Amazônia brasileira. 2. Um tumor
maligno das brânquias relacionado com as metacercárias de um trematódeo. - Acta Amazonica
10(3): 651-656.
THATCHER, V.E. & A.B. VARELLA (1981): Duas novas espécies de Megacoelium SZIDAT, 1954
(Trematoda: Haploporidae) parasitos estomacais de peixes da Amazônia brasileira, com uma
redefinição do gênero. - Acta Amazonica 11(2): 285-289.
TRAVASSOS, L. (1933): Sur un nouveau trématode de poisons de la vallée do fleuve Parahyba. - C. R.
Soc. Bio., Paris 114: 839-840.
TRAVASSOS, L. (1934): Sinopse dos Paramphistomoidea. - Mem. Inst. Oswaldo Cruz 29(1): 19-178.
TRAVASSOS, L. (1948a): Contribuição ao conhecimento dos helmintos dos peixes de água doce do
Brasil. I. Zonocotyle bicaecata n. g., n. sp. (Trematoda, Aspidogastridae). - Mem. Inst. Oswaldo
Cruz 45(2): 513-516.
TRAVASSOS, L. (1948b): Contribuição ao conhecimento dos helmintos dos peixes de água doce do
Brasil. II. Iheringtrema ilhringi n. g., n. sp. (Trematoda, Heterophyidae). - Mem. Inst. Oswaldo
Cruz 45(2): 517-520.
TRAVASSOS, L. (1961): Sôbre um novo trematódeo parasito de intestino de peixes de água doce do
Brasil. Curumai curumai gen. nov., sp. nov. (Trematoda, Angiodictyidae).- Atas Soc. Biol. Rio de
Janeiro 5(1-2): 1-4.
TRAVASSOS, L. & A. KOHN (1965): Lista dos helmintos parasitos de peixes encontrados na Estação
Experimental de Biologia e Piscicultura de Emas, Pirassununga, Estado de São Paulo. - Pap.
Avuls. Dept. Zool. S. Paulo 17(5): 35-52.
TRAVASSOS, L., TEIXEIRA DE FREITAS, J.F. & A. KOHN (1969): Trematódeos do Brasil. - Mem. Inst.
WILLEY, C.H. (1930): Studies on the lymph system of digenetic trematodes. - J. Morph. 50: 1-30.
WOTTON, R.M. & F. SOGANDARES-BERNAL (1963): A report on the occurrence of microvillus-like
structures in the caeca of certain trematodes (Paramphistomidae). - Parasit. 53: 157-161.
YAMAGUTI, S. (1953): Systema Helminthum. Part 1. Digenetic trematodes of fishes. - Publ. by
author, Tokyo: 405 pp.
YAMAGUTI, S. (1958): Systema Helminthum. Vol.1, parts 1 & 2. The digenetic trematodes of
Vertebrates.- Intersci. Publ. Co., New York: 1575 pp.
YAMAGUTI, S. (1971): Synopsis of the digenetic trematodes of Vertebrates. - Vol. 1., Keigaku Co.,
Tokyo: 1074 pp.
5
CESTODARIA AND CESTODA
The cestodarians are large unsegmented worms that are endoparasites in the body cavity
or gut of ancient fishes. The Subclass Cestodaria contains the Order Gyrocotylidea with 2
genera and 9 species, and Amphilinidea with 9 genera and 9 species. Gyrocotylideans
inhabit the intestinal tracts of certain marine fishes and are therefore beyond the scope of
this book, Amphilinideans are found in the body cavities of fishes, except for a single
species reported from an Australian turtle. The giant arapaima (Arapaima gigas), which is
the world’s largest strictly freshwater fish and occurs in the Amazon basin, is host to two
genera and species of this group. The origin and relationships of the Cestodaria are
obscure. They had been thought of as primitive tapeworms, but some zoologists now
insist that they are highly evolved and far from primitive. They were often regarded as
animals intermediate between trematodes and cestodes, but it is now believed that they
are more closely related to monogenoideans.
The two cestodarians from the arapaima are: Schizochoerus liguloideus (Fig. 5-38 A-B)
and Nesolecithus janicki (Figs. 5-43 A-B & 5-44). Both of these species reach a length of
15-16 cm but the former is only about 1.5 cm wide while the latter reaches 4 cm in
width. The internal anatomy is similar in both genera. The female system has an ovary,
oviduct, ootype, vagina with two posterior vaginal pores, seminal receptacle and uterus,
with an anterior pore. The uterus traverses the length of the body three times, and the
vitelline follicles extend throughout most of the body length in lateral strings. The male
system consists of numerous testes situated in elongate lateral fields, vasa efferentia, a
vas deferens and a copulatory bursa with a pore at the posterior extremity (posterior to
the vaginal openings).
The Class Cestoda (= Eucestoda) contains the true tapeworms most of which have a
flattened segmented body (strobila) and an attachment organ (scolex) provided with
suckers or hooks. The scolex has usually been regarded as the head end, but some
zoologists affirm that it really represents the posterior extremity and that it was derived
from the monogenoidean opisthaptor. Be that as it may, the scolex is followed by a “neck”
region from which new segments are budded. Beyond the neck, there are usually a
number of young immature segments (proglottids) within which the “anlagen” of
reproductive structures can be seen. Following the young proglottids, there are mature
ones showing fully developed sexual organs. Nearly all tapeworms are hermaphroditic and
each mature proglottid is provided with one or two complete sets of male and female
organs. Posterior to the mature proglottids there are normally several gravid segments in
which the reproductive organs are reduced but the uterus is large and engorged with eggs.
The reproductive systems of each mature proglottid are similar to those of
trematodes (Fig. 5-17 B). There is a lobate ovary, an oötype, a saccular uterus and a vagina
opening laterally. There are numerous vitelline follicles laterally. On the male side there are
several spherical testes, vasa efferentia, a vas deferens, a cirrus and cirrus sac. The
common genital pores may be ventral or lateral in the strobila and they may alternate
regularly or irregularly (when lateral).
Although no life-cycle has been experimentally demonstrated for any Neotropical fish
tapeworm, we can presume the following sequence based on indirect evidence. Eggs from
the gravid proglottid are shed in the feces of the host fish. Contractions of the proglottid
force the eggs out of the uterus. A six-hooked (hexacanth) ciliated larva hatches from
each egg and swims actively for some hours. This larva is ingested by a crustacean
(probably a copepod) where it penetrates to the hemocoel and develops into a more
advanced larval form with a scolex (procercoid larva). The crustacean is in turn eaten by
a small fish where the procercoid develops further into a plerocercoid. The plerocercoid
larva normally becomes encysted in the organs or muscles of the second intermediary
host. The latter is eventually devoured by a larger predatory fish where the plerocercoid
escapes the cyst and develops into the adult stage in the intestinal tract.
III. Pathology
Adult tapeworms in the intestinal lumen of the definitive host cause little or no damage.
There may be some irritation at the site of attachment of the scolex with a concomitant
increase in mucus production. Plerocercoid larval invasion in the intestinal wall of the
fish intermediate host, on the other hand, provokes temporary hemorrhaging followed
by an inflammatory reaction and eventual encapsulation of the invaders. Some fish
react strongly to the presence of these larvae and are able to encapsulate many of them
in the intestinal wall itself (Fig. 11-2). Other host species are seemingly unable to
encapsulate the larvae in the intestinal wall and they therefore pass through to the
mesenteries or the surfaces of internal organs where they finally become encysted.
Encapsulated plerocercoids are frequently seen in large numbers and when present in
the intestinal wall they may decrease the fish’s capacity to absorb nutrients. When
numerous in the mesenteries, these larvae can form tissue-like masses that resemble
adipose or testicular formations. They can be distinguished from the latter, however, by
the inverted muscular protoscolex in each capsule.
Wild caught crustaceans and minnows should not be fed to captive fish. If it is
necessary to use live animals as food, they should be grown in a separate culture for this
purpose. Freshly killed animal foods should also be avoided. Such material can be
utilized, however, if it has been deep frozen for 48 hours to kill any larval cestodes that
may be present. The recommended treatment for adult tapeworms in the definitive fish
host is: Di-N-Butyl Tin Oxide which can be mixed with the ration at 0.3 % of body
weight and given in a one to five day period.
Adult tapeworms can be collected and prepared in the same way that was described for
trematodes in Chapter 4. Special care should be taken when fixing the strobila in order to
get it straight and flat. Fixation between glass slides or between slide and cover glass is
recommended. Tapeworms can be killed by placing them in a Petri dish of water or
physiological saline solution and leaving the dish in an embedding oven for 5-10 minutes
or in the refrigerator for 10-30 minutes.
The only cestodarians known from Neotropical freshwater fishes are the two species
mentioned above from the giant arapaima. The Class Eucestoda is represented by the
Orders Proteocephalidea (with 19 genera and 45 species). Tetraphyllidea (with 5 genera
and 15 species) and Trypanorhyncha (with 1 genus and species). The latter two orders are
known only from rays except for one species of Tetraphyllidea reported from a sawfish.
The proteocephalideans parasitize mainly catfishes.
Genera and species of Neotropical fish tapeworms are not easy to characterize or
identify. Such characters as forms and size of the scolex, number and position of the
testes, position of the genital pores and number of segments in the strobila have been
used. In the proteocephalids, there are longitudinal muscle fibers running down the
center of mature proglottids that mark off a central (medullary) region from an outer
(cortical) area. Genera differ as to which of the reproductive organs occur in the
medulla and which in the cortex. To study this aspect it is necessary to make transverse
sections of mature proglottids.
Relatively little work has been done on freshwater fish parasites in the Neotropics.
The principal references to the group are: BROOKS & THORSON 1976; BROOKS et al. 1981;
MAYES et al. 1978; REGO 1979, 1984; REGO & LUNA DIAS 1976; REGO et al. 1974;
WOODLAND 1925, 1933a-c, 1934a-c and 1935a-c.
Key to Cestodaria and Cestoda of Neotropical freshwater fishes
I. Body unsegmented, rounded or tapered at extremities; scolex absent; from peritoneal cavity of host .......................................................
........................................................................................................................... Cestodaria ..................................................................... Amphilinidae
A. Body about 4 times longer than wide ................................................................................................................... Nesolecithus (Fig. 5-43 A-B)
B. Body about 10 times longer than wide ............................................................................................................... Schizochoerus (Fig. 5-42 A-B)
II. Body segmented, usually tapered towards anterior extremity; scolex present; from digestive tract of host ................................ Cestoda
A. Scolex provided with 2 bothridia and 4 invaginable tentacles which are covered with hooks ..................................................................
Trypanorhyncha ....................................................................................... Eutetrarhynchidae .................................... Eutetrarhynchus (Fig. 5-1)
B. Scolex provided with 4 bothridia; tentacles absent; hooks present or absent .................................................................... Tetraphyllidea
1. Hooks absent .......................................................................................................................................................................... Phyllobothriidae
a. Bothridia sessile, without luculi .................................................................................................................... Anthobothrium (Fig. 5 -37)
b. Bothridia stalked, locular.
1) Bothridia elongate, rounded ......................................................................................................................................... Rhinebothrium

2) Bothridia short, quadrate ........................................................................................................... Rhinebothroides (Fig. 5 -2 A-B)
2. Four pairs of hooks present ................................................................................................................................................. Oncobothriidae
a. Bothridia triloculate ........................................................................................................................................ Acanthobothrium (Fig. 5-5)
b. Bothridia nonloculate ...................................................................................................................... Potamotrygonocestus (Fig. 5-33 A-B)
C. Scolex usually provided with 4 suckers; bothridia and tentacles absent ....................................................................... Proteocephallidae
1. Suckers absent or concealed by metascolex.
a. Suckers absent ...................................................................................................................................................... Ornithoscolex (Fig. 5-18)
b. Suckers concealed by metascolex .......................................................................................................................... Manaosia (Fig. 5-12)
2. Suckers replaced by 4 sac-like structures ............................................................................................................... Megathylacus (Fig. 5-24)
3. Each sucker composed of two compartments, only one of which opens exteriorly ........................................... Brayela (Fig. 5-36)
4. Suckers bilocular
a. Sucker margins contiguous ................................................................................................................................... Peltidocotyle (Fig. 5-22)
b. Sucker margins separated ........................................................................................................................... Amphoteromorphus (Fig. 5-8)
209
5. Suckers trilocular Gibsoniela (Fig. 5-20).
210
6. Suckers with triangular spinos apertures ....................................................................................................................... Endorchis (Fig. 5-3)
7. Four simple suckers and metascolex present.

a. Scolex pointed anteriorly; metascolex half covers suckers ........................................................................ Choanoscolex (Fig. 5-11)
b. Scolex pointed or not; metascolex covers suckers in lateral view.
1) Fewer than 15 segments in strobila ........................................................................................................ Sciadocephalus (Fig. 5-31)
2) More than 15 segments in strobila ............................................................................................... Rudolphiela (Figs. 5-34 & 5-35)
c. Scolex pointed or not; metascolex distant from suckers.
1) Metascolex forming discrete horizontal band ................................................................................... Ephedrocephalus (Fig. 5-10)
2) Metascolex forming longitudinal ridges ....................................................................................................... Monticellia (Fig. 5-23)
3) Metascolex forming irregular mass .......................................................................................................... Woodlandiella (Fig. 5-19)
8. Four simple suckers present; metascolex absent.
a. Suckers very large, spherical, fused to each other internally ..................................................................... Zygobothrium (Fig. 5-15)
b. Suckers adorned.
1) Suckers surrounded by tegumental flaps ................................................................................................... Nominoscolex (Fig. 5-4)
2) Suckers spinous ................................................................................................................................................................. Spasskyellina
c. Suckers unadorned.
1) Reproductive organs including vitellaria in medulla ................................................. Proteocephalus (Figs. 5-26, 5-30 & 5-32)
2) Reproductive organs in medulla except for vitellaria which are in lateral cortex .......................... Nominoscolex (Fig. 5-14)
Note: The genus Myzophorus WOODLAND, 1934 (Fig. 5 – 9) is of doubtful validity and was considered to be a phyllobothriid by WARDLE &
MCLEOD (1952).
Clave para Cestodaria y Cestoda de peces Neotropicales de agua dulce
I. Cuerpo sin segmentación, arredondeado o alongado en las extremidades; escólex ausente; encontrado en la cavidad peritoneal del
hospedero ........................................................................................................ Cestodaria ..................................................................... Amphilinidae
A. Cuerpo ± 4 veces más largo que ancho ............................................................................................................... Nesolecithus (Fig. 5-39 A-B)
B. Cuerpo ± 10 veces más largo que ancho ........................................................................................................... Schizochoerus (Fig. 5-38 A-B)
II. Cuerpo segmentado, generalmente alongado hacia la extremidad anterior; escólex presente; en el tracto digestivo del hospedero .....
............................................................................................................................................................................................................................... Cestoda
A. Escólex provisto de 2 botridias y 4 tentáculos invaginables cubiertos con ganchos ...................................................................................
Trypanorhyncha ....................................................................................... Eutetrarhynchidae .................................... Eutetrarhynchus (Fig. 5-1)
B. Escólex provisto con 4 botridias; tentáculos ausentes; ganchos presentes o ausentes ..................................................... Tetraphyllidea
1. Ganchos ausentes .................................................................................................................................................................. Phyllobothriidae
a. Botridias sésiles, con lóculos .......................................................................................................................... Anthobothrium (Fig. 5-37)
b. Botridias pedunculadas, loculares.

1) Botridias alargadas, arredondeadas ............................................................................................................................. Rhinebothrium
2) Botridias cortas, cuadradas .......................................................................................................... Rhinebothroides (Fig. 5-2 A-B)
2. Cuatro pares de gancho presentes ...................................................................................................................................... Oncobothriidae
a. Botridias triloculadas ...................................................................................................................................... Acanthobothrium (Fig. 5-5)
b. Botridias sin lóculos ......................................................................................................................... Potamotrygonocestus (Fig. 5-33 A-B)
C. Escólex generalmente provisto de 4 ventosas; botridias y tentáculos ausentes ........................................................... Proteocephallidae
1. Ventosas ausentes o cubiertas por el metaescólex.
a. Ventosas ausentes ................................................................................................................................................ Ornithoscolex (Fig. 5-18)
b. Ventosas cubiertas por el metaescólex ................................................................................................................. Manaosia (Fig. 5-12)
2. Ventosas substituidas por 4 estructuras en forma de saco ................................................................................ Megathylacus (Fig. 5-24)
3. Cada ventosa compuesta por 2 compartimentos, uno de los cuales abre exteriormente .................................... Brayela (Fig. 5-36)
4. Ventosas biloculares
a. Márgenes de la ventosa contiguos ...................................................................................................................... Peltidocotyle (Fig. 5-22)
b. Márgenes de la ventosa separados .......................................................................................................... Amphoteromorphus (Fig. 5 -8)
211
5. Ventosas triloculares ....................................................................................................................................................... Gibsoniela (Fig. 5-20)
212
6. Ventosas con aberturas triangulares y espinosas ......................................................................................................... Endorchis (Fig. 5-3)
7. Cuatro ventosas simples y metaescólex presentes

a. Escólex puntiagudo anteriormente; metaescólex cubre la mitad de las ventosas ................................. Choanoscolex (Fig. 5-11)
b. Escólex puntiagudo o no; metaescólex cubre las ventosas en vista lateral.
1) Menos de 15 segmentos en estróbilo ..................................................................................................... Sciadocephalus (Fig. 5-31)
2) Más de 15 segmentos en estróbilo ............................................................................................... Rudolphiela (Figs. 5-34 & 5-35)
c. Escólex puntiagudo o no; metaescólex distante de las ventosas.
1) Metaecoolex formando una banda horizontal discreta ................................................................... Ephedrocephalus (Fig. 5-10)
2) Metaescólex formado arrugas longitudinales ............................................................................................. Monticellia (Fig. 5-23)
3) Metaescólex formando masa irregular .................................................................................................... Woodlandiella (Fig. 5-19)
8. Cuatro ventosas simples presentes; metaescólex ausente.
a. Ventosas muy grandes, esféricas, fundidas unas con otras internamente ............................................... Zygobothrium (Fig. 5-15)
b. Ventosas adornadas.
1) Ventosas rodeadas de ondulaciones tegumentarias ................................................................................. Nominoscolex (Fig. 5-4)
2) Ventosas espinosas ........................................................................................................................................................... Spasskyellina
c. Ventosas sin adornos.
1) Organos reproductores incluyendo vitelaria en la médula ...................................... Proteocephalus (Figs. 5-26, 5-30 & 5-32)
2) Organos reproductores en médula excepto por vitellaria que está en la corteza lateral ............. Nominoscolex (Fig. 5-14)
Nota: El género Myzophorus WOODLAND, 1934 (Fig. 5 – 9) es de validez dudosa y estaba considerado como un phyllobothriid por WARDLE &
MCLEOD (1952).
VII. Checklist of Cotyloda (= Cestodaria) and Eucestoda (= Cestoda) from

Neotropical freshwater fishes
Measurements are given normally in millimeters and in microns (µm) for hooks and eggs,
unless otherwise indicated.
Class Cotyloda WARDLE, MCLEOD & RADINOVSKY, 1974 ( = Cestodaria MONTICELLI, 1892)
Order Amphilinidea POCHE, 1922
Amphilinidae CLAUS, 1879
Body flat. linguiform or tape-like. with rounded or pointed extremities;
without segmentation. Anterior end provided with small protrusible
proboscis, or sucker-like structure. Testes numerous, in two lateral,
longitudinal rows. Ovary near posterior extremity. Genital pores (male and
female) near posterior end; separate or opening into common atrium.
Uterus N-shaped with external opening near anterior extremity. Excretory
system of two longitudinal ducts opening posteriorly through a common
pore. Body cavity of fish.
Nesolecithus POCHE, 1922
Amphilinidae. Body width about one fourth of length. Accessory seminal
receptacle about one sixth of body length.
N. janicki POCHE, 1922: Arapaima gigas: Amazonia (Brazil, Peru, Colombia &
Venezuela). (Fig. 5-43 A-B). Body = 16 x 4 cm. (maximum).
Schizochoerus POCHE, 1922
Amphilinidae. Body width about one tenth of length. Accessory seminal
receptacle about one third of body length.
S. liguloideus (DIESING, 1850): Arapaima gigas: Amazonia (Brazil, Peru, Colom-
bia & Venezuela). (Fig. 5-42 A-B). Body = 15 x 1.5 cm (maximum).
Class Eucestoda SOUTHWELL, 1930 (= Cestoda MONTICELLI, 1892)
Order Proteocephalidea MOLA, 1928
Proteocephalidae LA RUE, 1914, WOODLAND, 1933
Scolex with suckers (usually 4) not bothridia. Body parenchyma divided by
muscle layers into external (cortical) and internal (medullary) regions.
Ovary, testes, vitellaria and uterus usually medullary, but may be cortical.
Vagina dorsal to cirrus sac. Gravid proglottids not separating from
strobila. Intestine of fishes, amphibians and reptiles.
Amphoteromorphus DIESING, 1850
Scolex large; suckers bilocular, largely concealed by anteriorly projecting
prepuce-like tegument; apical organ absent. Genital apertures unilateral.
Vitellaria cortical, lateral, in dorsal and ventral groups. Intestine of
Amazonian catfish.
A. parkarmoo WOODLAND, 1935: Pseudopimelodus zungaro: Brazil. (Fig. 5-8). Body
= 33 x 1 (maximum); Scolex = 0.47-0.88 x 0.53-1.12 (wider than long);
Testes = fewer than 40; Egg = 26 x 16 µm.
A. peniculus DIESING, 1850: Brachyplatystoma rousseauii: Brazil. (Fig. 5-17 A-B).

Body and scolex similar to those of A. parkarmoo, but suckers are more
exposed and transverse excretory canals are absent.
A. piraeeba WOODLAND, 1934: “piraeeba”: Brazil. (Fig. 5-16). Body = 50-60 x
1.3 (maximum); Proglottids wider than long; Testes numerous (about 200);
Egg = 30 µm in diameter.
Brayela REGO, 1985
Scolex with 4 suckers, each having 2 compartments only one of which
opens exteriorly. Reproductive organs medullary. Uterus tubular, without
lateral outgrowths. Intestine of Amazonian fishes.
B. karuatayi (WOODLAND, 1934); Glanidium sp.: Brazil. (Fig. 5-36). Body = 30 x
1; Genital pores alternating irregularly; Scolex = 0.24 x 0.40; Testes =
fewer than 50 (about 24 on each side).
Choanoscolex LA RUE, 1911
Scolex tapering anteriorly, enlarged at base, with body folds covering bases of
suckers. Neck unsegmented. Last proglottids longer than wide. Cirrus sac large;
genital pores lateral in anterior part of proglottid. Intestine of siluriform fishes.
C. abscisus (RIGGENBACH, 1895) LA RUE, 1911: Silurus sp.: Paraguay. (Fig. 5-
11). Body = 25 x 30 x 0.77 mm; Strobila = maximum of 70 proglottids;
Testes = about 100; Egg = 21 x 16 µm.
Endorchis WOODLAND, 1934
Scolex with 4 suckers having triangular openings and armed with minute
spines. Apical organ prominent. Genital apertures alternating irregularly.
Vitellaria and part of ovary cortical. Testes medullary. Mature proglottid
longer than wide. Intestine of Amazonian siluroid fishes.
E. piraeeba WOODLAND, 1934: Brachyplatystoma filamentosum: Brazil. (Fig. 5-3).
Body = 17-45 x 1.3; Mature proglottids nearly square, marked with
transverse creases; Scolex = 0.6 x 0.54; Testes = 100-150 in number.
E. mandube WOODLAND, 1935: Ageneiosus brevifilis: Brazil. (Fig. 5-6). Body = 40
x 0.79; Mature proglottids wider than long; Scolex = 0.7 x 0.6; Testes =
about 150 in number (note: The suckers on the scolex of this species
appear to be triloculate, and if so, it might be a species of Gibsoniela or a
synonym of G. mandube, as was suggested by REGO 1985).
Ephedrocephalus DIESING, 1850
Scolex large, wrinkled, longitudinally grooved, with 4 suckers on an
anterior projection. Testes in dorsal cortex. Vitellaria in ventral cortex.
Ovary medullary. Intestine of Amazonian siluriform fishes.
E. microcephalus DIESING, 1850: Phractocephalus hemiliopterus: Brazil. (Fig. 5-10).
Body = 100-150 x 4-8; Unsegmented neck region short, or absent;
Proglottids wider than long; Testes = 20-30 in number; Egg = 44 x 22 µm.
Gibsoniela REGO, 1985
Scolex with 4 elongate, triloculate suckers. Apical organ absent. Proglottids wider
than long, except for terminal ones. Reproductive organs medullary, except for
vitellaria which extend into cortex. Intestine of Amazonian siluroid fishes.
G. mandube (WOODLAND, 1935): Ageneiosus brevifilis: Brazil. (Fig. 5-20). Body =

45 x 2; Genital pores alternating irregularly; Scolex = 0.18-0.21 x 0.18-
0.24; Testes = 100-150 in number; Egg = 19 µm in diameter.
Manaosia WOODLAND, 1935
Scolex large, with wrinkled surface which conceals 4 large suckers. Neck
region un-segmented. Ovary medullary, but extends into cortex. Intestine
of Amazonian siluroid fish.
M. bracodemoca WOODLAND, 1935: Platystoma sp.: Brazil. (Fig. 5-12). Body =
6-7 x 0.7; Scolex = 0.94 in diameter; Suckers = 0.33 in diameter.
Megathylacus WOODLAND, 1934
Scolex large, with 4 voluminous sac-like structures instead of suckers;
apical organ absent. Neck segmented. Proglottids wider than long, except
for last few which are nearly square. Medullary region large, contains all
reproductive organs. Intestine of Amazonian siluroid fishes.
M.jandia WOODLAND, 1934: Rhamdia sp.: Brazil. (Fig. 5-24). Body = 25 x 1.65;
Scolex = 2.5 x 1.6; Genital pores alternating irregularly.
Monticellia LA RUE, 1911
Scolex with 4 simple suckers. Apical organ absent. Genital pores alternat-
ing irregularly. Testes, vitellaria, uterus and most of ovary in cortex;
vitellaria ventrolateral. Intestine of Amazonian siluroid fishes.
M.coryphicephala (MONTICELLI, 1892) LA RUE, 1914: Silurus sp.: locality un-
known, but probably Brazil. (Fig. 5-13). Body = length unknown, width =
1.8-2.0; About 100 testes present.
M.megacephala WOODLAND, 1933: Platystomatichthys sturio: Brazil. Body = 24 x
1.1; Scolex = 0.41-0.76 x 0.53-0.79; Mature proglottids nearly square;
gravid proglottids longer than wide.
M.piracatinga WOODLAND, 1935: Pimelodus pati: Brazil. Body = 6 x 0.5; Scolex =
0.59-0.74 x 0.34-0.36; Suckers = 0.16-0.22 in diameter.
M.piramutab (WOODLAND, 1933): Brachyplatystoma vaillanti: Brazil. Body = 35 x
1.5; Scolex = 0.88 x 1.4; Suckers = 0.39; Egg = 18 x 13 µm.
M.rugosa WOODLAND, 1935: Pseudoplatystoma fasciatum: Brazil. (Fig. 5-23).
Body = 40 x 1.0; Scolex = 0.88-1.3 x 0.6-1.0; Suckers = 0.29 x 0.19;
Egg = 14-15 µm.
M.siluri (FUHRMANN, 1916): Cetopsis caecutiens: Brazil. (Fig. 5-21). Body = 80 x
1.6-2.0; Scolex = 0.7 wide; Suckers = 0.28 in diameter.
M.surubim (WOODLAND, 1933); Pseudoplatystoma tigrinum: Brazil. Body = 5.8 x
1.4; Testes = 200-300 in number.
Myzophorus WOODLAND, 1934
Scolex with 4 simple suckers. Apical organ absent. Genital pores alternat-
ing irregularly. Testes medullary; uterus vitellaria and part of ovary
cortical. Intestine of Amazonian siluroid fishes.
M.admonticellia WOODLAND, 1934: Pinirampus sp.: Brazil (Manaus). (Fig. 5-9).
Body = 100 x 1; Scolex = 0.17-0.36 x 0.25-0.48; Strobila = segments in
anterior one-fifth wider than long; mature proglottids nearly square;
posterior two-thirds of strobila with gravid proglottids which are longer

than wide; Egg = 30 µm. in diameter; Testes = about 200 in number.
M.pirarara WOODLAND, 1935: Phractocephalus hemilopterus: Brazil. Body = 33 x
0.94; Scolex = 0.7 x 0.8; Suckers = 0.2; Testes = 50-60 in number; Egg =
14-21 µm.
Nominoscolex WOODLAND, 1934
Scolex with 4 simple suckers. Apical organ present or absent. Testes, ovary
and uterus medullary. Vitellaria lateral, in cortex. Genital pores alternating
irregularly. Intestine of Amazonian siluroid fishes.
N. dorad (WOODLAND, 1934): Brachyplatystoma rousseauxii: Brazil. (Fig. 5-14).
Body = 134 x 1.7; Scolex = 0.16-0.26 x 0.24-0.38; Suckers = 0.19; Testes =
100 in number; Egg = 30 µm.
N. kaparari WOODLAND, 1935: Pseudoplatystoma tigrinum: Brazil. Body = 30 x 0.99;
Scolex = 0.15-0.26 x 0.28-0.33; Testes = fewer than 100; Egg = 14µm.
N. lenha (WOODLAND, 1933): Platystomatichthys sturio: Brazil. Body = 130 x 2.6;
Scolex = 0.16-0.28 x 0.19-0.28; Suckers = 0.09-0.16; Testes = more than
200; Egg = 23 µm.
N. piracatinga WOODLAND, 1935: Pimelodus pati: Brazil. (Fig. 5-7). Body = 20 x
0.7; Scolex = 0.19 x 0.22; Suckers = 0.10-0.11; Testes = about 150; Egg =
10-11 µm.
N. piraeeba WOODLAND, 1934: Brachyplatystoma filamentosum: Brazil. Body = 40 x
0.92; Scolex = 0.18-0.23 x 0.25-0.33; Testes = 100-150 in number; Egg =
25 µm.
N. sudobim WOODLAND, 1934: Pseudoplatystoma fasciatus and Brachyplatystoma
rousseauxii: Brazil. Body = 53 x 1.2; Scolex = 0.20-0.22 x 0.12-0.15; Testes
= 200-250 in number; Egg = 24-26 µm.
N. touzeti CHAMBRIER & VAUCHER, 1992: Ceratophrys cornuta: Rio Aguarico,
Napo Province, Amazonian Ecuador. Body = about 73 proglottids; 64 mm
long; Scolex = 0.48-0.53 x 0.36-0.38; Testes = 97-137 in number; Egg =
70-120 µm.
N. woodlandi FREZE, 1965: Pseudoplatystoma fasciatum: Brazil (Fig. 5-4). Body =
85 x 2.0; Scolex with four suckers having 4 pointed flaps on each one; size
= 0.39 x 0.26; Testes about 150.
Ornithoscolex WOODLAND, 1933
Scolex covered externally by convoluted longitudinal masses of cuticular
and subcuticular tissue; suckers absent. Neck absent. Mature proglottids
wider than long; with psedo-segmenation. Vitellaria lateral, in dorsal and
ventral fields. Intestine of Amazonian siluroid fishes.
O. lenha WOODLAND, 1933: Platystomatichthys sturio: Brazil. (Fig. 5-18). Body =
40 x 2.2; Scolex = 0.8-1.4 x 1.0-1.5; Testes = about 100; Egg = 21 µm.
Peltidocotyle DIESING, 1850
Scolex dorso-ventrally flattened, covered by convoluted tegument and
provided with 4 bilocular suckers. Internal anatomy unknown. Intestine of
Amazonian siluroid fishes.
P. rugosa DIESING, 1850: Pseudoplatystoma tigrinum: Brazil. (Fig. 5-22). Scolex

about twice as wide as strobila; measurements unavailable.
Proteocephalus WEINLAND, 1858
Scolex with 4 simple suckers which are without tegumental adornment.
Apical sucker present or absent. Reproductive organs medullary. Vitellaria
in lateral bands. Testes continuous across proglottid. Intestine of freshwa-
ter fish, rarely amphibians.
P. fossatus (RIGGENBACH, 1895): Pimelodus pati: Paraguay. (Fig. 5-27). Body =
35-40 long; Scolex = 0.71 in diameter; Suckers = 0.34; Testes = 120-150
in number.
P. jandia WOODLAND, 1934: Ramdia sp.: Brazil. (Fig. 5-26). Body = 49 x 0.94;
Scolex = 0.16 wide; Testes = about 100; Egg = 30 µm.
P. kuyukuyu WOODLAND, 1935: Oxydorus niger: Brazil. Body = 35 x 0.94;
Scolex = 0.16-0.25 x 0.13-0.25; The description was based on immature
specimens.
P. macdonaghi (SZIDAT & NANI, 1951): Basilichthys microlepidotus: Argentina. (Fig.
5-29). Body = 15 x 0.24; Scolex = 0.4 wide; Suckers = 0.16; Testes = 4-60
in number; Egg = 50 x 20 µm.
P. macrophallus (DIESING, 185 0): Cichla ocellaris: Brazil. (Fig. 5-30). Body = 6 x
0.94; Suckers = 0.2; Strobila with fewer than 20 proglottids; Testes = 30-40
in number.
P. manjuariphilus VIGUERAS, 1936: Atractosteus tristoechus: Cuba. (Fig. 5-28).
Body = 110-115 x 1.5-2.0; Scolex = 0.13 x 0.11; Testes = 30-40 in number;
Egg = 32 µm.
P. microscopicus WOODLAND, 1935: Cichla ocellaris: Brazil. (Fig. 5-32). Body =
2.06 x 0.26-0.41; Suckers = 0.14-0.18; Testes = about 20; Egg = 8 x 1 µm.
P. platystomi LYNSDALE, 1959: Pseudoplatystoma sp.: Brazil. Body = 45 x 1.2
(incomplete fragment); Scolex = 1.3 x 0.87; Testes = 35-50 in number;
Egg = 50-60 µm.
P. renaudi CHAMBRIER & VAUCHER, 1994: Platydoras costatus: Neembucu Prov-
ince, Paraguay (Fig. 5-40). Body = 120-480 mm long. Testes = 124-276 in
number.
P. soniae CHAMBRIER & VAUCHER, 1994: Platydoras costatus: Neembucu Prov-
ince, Paraguay. (Fig. 5-41). Body = 84-145 mm long. Testes = 139-371 in
number.
P. sophiae CHAMBRIER & REGO, 1994: Paulicea luetkeni: Brazilian Amazon. (Figs.
5-38 & 5-39). Body = 60-130 mm long. Testes = 83-141 in number.
Vitellaria lateral, postporal only.
Rudolphiella FÜHRMANN, 1916
Scolex large, wrinkled, longitudinally furrowed, apex projected and with 4
suckers at its margin. Testes cortical, in single layer over entire proglotiid.
Vitellaria cortical and ventral. Uterus medullary. Ovary dorsal, part
medullary and part cortical. Intestine of South American siluroid fishes.
R. labosa (RIGGENBACH, 1895): FÜHRMANN, 1916: Pimelodus pati: Paraguay.

Body = 20-30 x 1.3 (40-50 proglottids); Testes = 150-200 in number;
Egg = 18-19 µm.
P. myoides (WOODLAND, 1934); Pirinampus pirinampus: Brazil. (Fig. 5-34). Body =
11 x 0.6; Scolex = 0.47 x 0.59; Strobila with about 50 proglottids; Testes =
about 100; Egg = 62-77 x 16 µm.
P. piranabu (WOODLAND, 1934); “piranabu”: Brazil. (Fig. 5-35). Body = 10 x
0.88; Scolex = 0.29-0.49 x 0.58-0.68; Strobila with fewer than 50 proglotti-
ds; Testes = about 100; Egg = 20 -26 x 10-11 µm.
Sciadocephalus DIESING, 1850.
Scolex cup-shaped, with 4 simple suckers; small apical sucker present.
Strobila short and wide, with fewer than 15 proglottids. Parenchymal
muscles lacking. Intestine of Amazonian fishes.
S. megalodiscus DIESING, 1850: Cichla ocellaris: Brazil. (Fig. 5-31). Body = 3-9 x 1;
Suckers = 0.18 in diameter; Testes numerous.
Spasskyellina FREZE, 1965
Scolex simple, with 4 spinous suckers. Testes, ovary and uterus in dorsal
cortex. Vitellaria lateral or ventral, in cortex. Intestine of Amazonian
siluroid fishes.
S. lenha (WOODLAND, 1933): Platystomatichthys sturio: Brazil. Body = 30 x 1;
Scolex = 0.24-0.29 x 0.23-0.36; Testes = more than 200; Egg = 21µm.
S. spinulifera (WOODLAND, 1934): Pseudoplatystoma fasciatum: Brazil. Body =
25 x 0.59; Scolex = 0.21-0.28 x 0.26-0.49; Suckers = 0.18-0.24; Testes =
about 60.
Woodlandiella FREZE, 1965
Scolex with 4 suckers and a prominent, wrinkled tegumental enlargement
(metascolex). Neck absent. Vitellaria lateral, in dorsal and ventral bands.
Intestine of Amazonian siluroid fishes.
W.myzofera (WOODLAND, 1933); Platystomatichthys sturio: Brazil. (Fig. 5-19).
Body = 30 x 1.8; Scolex = 1.1 x 2.3; Testes = about 150.
Zygobothrium DIESING, 1850
Scolex with 4 large, globular, hollow suckers which are fused to each other
laterally. Apical organ absent. Genital pores unilateral. Testes numerous, in
dorsal medulla. Vitellaria cortical, in two lateral crescents, with few follicles
in mid-dorsal line. Ovary and uterus medullary. Intestine of Amazonian
siluroid fishes.
Z. megacephalum DIESING, 1850: Phractocephalus hemiliopterus and Pirarara bicolor:
Brazil. (Fig. 5-15). Body = 76 x 5; Scolex = 2 x 3; Testes = 150-200 in
number; Egg = 45 µm.
Order Tetraphyllidae CARUS, 1863
Onchobothriidae BRAUN, 1900
Scolex with 4 sessile or slightly stalked bothridia and 4 pairs of hooks.
Accessory suckers present or absent. Genital pores alternating regularly or
irregularly. Intestinal spiral valve of elasmobranch fishes.
Acanthobothrium BENEDEN, 1850

Scolex with 4 trilocular bothridia and 4 pairs of forked hooks. Spiral valve
of elasmobranch fishes.
A. amazonensis MAYES, BROOKS & THORSON, 1978: Potamotrygon circularis:
Brazil (Amazonia). Body = 35 (maximum), with 75-100 proglottids;
Scolex = 0.061-0.79 wide: Bothridia = 0.51-0.62 x 0.30-0.42; Hooks =
145-184 µm.
A. quinonesi MAYES, BROOKS & THORSON, 1978: Potamotrygon magdalenae: Colom-
bia (Magdalena River). Body = 25 (maximum), with 55-75 proglottids;
Scolex = 0.51-0.62 wide; Bothridia = 0.38-0.48 x 0.28-0.31; Hooks = 100-
142 µm; Testes = 43-60 in number.
A. regoi BROOKS, MAYES & THORSON, 1981: Potamotrygon hystrix: Venezuela
(Orinoco River). Body = 45 (maximum), with 87-120 proglottids; Scolex =
0.70-0.90 x 0.80-1.1; Bothridia = 0.50-0.60 x 0.30-0.35; Hooks = 122-163
µm. Testes = 47-70 in number.
A. terezae REGO & DIAS, 1976: Potamotrygon motoro: Brazil. (Mato Grosso State).
Body 88 = 110, with 200-260 proglottids; Scolex = 2-3 wide; Hooks =
180-326 µm. Long; Testes = 120-140 in number.
Potamotrygonocestus BROOKS & THORSON, 1976
Scolex with 4 sessile nonloculate bothridia, each with accessory sucker and
pair of simple hooks. Spiral valve of elasmobranch fishes.
P. amazonensis MAYES , B ROOKS & T HORSON , 1981: Potomotrygon circularis:
Brazil (Amazonia). Body = 1.2-3.5 long, with 10-13 proglottids;
Scolex = 0.24-0.30 x 0.24-0.30; Bothridia = 0.33-0.41 x 0.13-0.21;
Hooks = 58-78 µm long (prong); 26-35 µm (base); Testes = 21-24 in
number.
P. magdalenensis BROOKS & THORSON, 1976: Potamotrygon magdalenae: Colombia
(Magdalena River). Body = 1.0-2.1 long, with 7-10 proglottids; Scolex =
0.14-0.16 x 0.24-0.29; Bothridia = 0.17-0.24 x 0.11-0.16; Hooks = 43-55
µm long; 19-29 µm base; Testes 20-29 in number.
P. orinocoensis BROOKS , M AYES & THORSON , 1981: Potamotrygon reticulates:
Venezuela (Orinoco River). (Fig. 5-33 A-B). Body = 1.3-1.6 long,
with 5-10 proglottids; Scolex = 0.15-0.23 x 0.20-0.22; Bothridia =
0.24-0.27 x 0.07-0.11; Hooks = outer 73-105 µm long (prong); 30-50
µm (base); inner: 80-125 µm (prong); 25-48 µm (base); Testes = 17-25
in number.
Phyllobothriidae
Scolex unarmed, but with 4 stalked or sessile bothridia which may be
simple, complexly folded or loculate. Accessory suckers sometimes
present. Genital pores unilateral or alternating regularly or irregularly.
Mature segments may detach from strobila and become gravid after
separation. Eggs spindle-shaped. Spiral valve of elasmobranch fishes.
Anthobothrium BENEDEN, 1850, emend. SOUTHWELL, 1925.
Bothridial surfaces nonloculate. Accessory suckers absent.
A. pristis WOODLAND, 1934: Pristis perottetti: Brazil (Amazonia). (Fig. 5-37).

Body = 110 x 2-3; Scolex = 0.40 x 0.36; Genital pores alternating
irregularly; Testes = more than 200.
Rhinebothrium LINTON, 1889
Scolex with 4 elongate, stalked bothridia having loculate surfaces. Genital
apertures alternating irregularly; female pore anterior to male; open into
genital atrium. Testes medial; vitellaria lateral. Uterus tubular. Spiral valve
of elasmobranch fishes.
R. paratrygoni REGO & DIAS, 1976: Potamotrygon hystrix, P. falkneri and P.
reticulatus: Brazil (Mato Grosso State), Venezuela (Orinoco River) &
Paraguay (Paraná River). Body = 23 x 0.3; Scolex = 0.87 (expanded);
Bothridia = 72-76 loculi; Testes = 4-8 in number.
Rhinebothroides MAYES, BROOKS & THORSON, 1981
Scolex with 4 pedicellated, quadrate bothridia which have shallow horizon-
tal loculi. Testes preovarian; cirrus sac at level of ovary, contains spinous
cirrus and internal seminal vesicle. Ovary bilobed, with shorter lobe on
poral side. Uterus with lateral diverticula. Spiral valve of South American
freshwater stingrays.
R. circularisi MAYES, BROOKS & THORSON, 1981: Potamotrygon circularis.: Brazil
(Amazonia). Body = 27 long, with 18-24 proglottids; Scolex = 0.9-1.5
wide; Bothridia = 0.67-1.0 x 0.40-0.53; number of locul = 70-80; Testes =
66-88 in number; Egg = 18 29 µm.
R. freitasi (REGO, 1979): Potamotrygon hystrix: Brazil (Amazonia). Body =
12-15 proglottids; Bothridia with about 60 loculi; Testes = 48-64 in
number.
R. glandularis BROOKS, MAYES & THORSON, 1981: Potamotrygon hystrix: Venezu-
ela (Orinoco River). Body = 50 long, with 25-30 proglottids; Bothridia =
0.61-0.71 x 0.46-0.71; medial loculi 51-59 in number; marginal loculi =
51-59 in number; Testes = 41-51 in number; Egg = 29-37 µm in
diameter.
R. moralarai (BROOKS & THORSON, 1976): Potamotrygon magdalenae: Colombia
(Magdalena River). Body = 18-24 proglottids; Bothridial loculi = 69-79 in
number; Testes = 66-88 in number. Uterine diverticula = 49-80 in number.
R. scorzai (LOPES-NEYRA & DÍAS-UNGRÍA, 1958): Potamotrygon hystrix, P. motoro,
P. reticulatus and Elipesurus spinicauda: Venezuela (Orinoco River) & Brazil
(Mato Grosso State). Body = 60-100 proglottids; Bothridial loculi = 69-79
in number; Testes = 60-99 in number; Uterine diverticula = 54-80 in
number.
R. venezuelensis BROOKS, MAYES & THORSON, 1981: Potamotrygon hystrix and P.
yepsi: Venezuela (Orinoco & Cachiri Rivers). Body = 60 long, with 20-30
proglottids; Bothridia = 0.36-0.72 x 0.26-0.69; medial loculi = 51-53 in
number; marginal loculi = 51-53 in number; Testes = 45-64 in number;
Egg = 19-24 µm in diameter.
Order Trypanorhyncha DIESING, 1863

Eutetrarhynchidae GUIART, 1927
Scolex with 4 invaginable tentacles which are covered with hooks of
several shapes, and 2 bothridia with free and notched posterior margins.
Tentacular bulbs at least three times longer than wide. Testes preovarian.
Spiral valve of elasmobranch fishes.
Eutetrarhynchus PINTNER, 1913
Eutetrarhynchidae. Scolex surface covered by hair-like spinules. Testes in
two layers. Vitellaria surrounding testicular area. Uterus extending entire
length of gravid proglottis. Spiral valve of elasmobranch fishes.
E. araya (WOODLAND, 1934): Potamotrygon hystrix, P. motoro, P. falkneri, P. reticulatus
and Elipesurus sp.: Brazil, Venezuela & Paraguay. (Fig. 5-1). Body = 27 x 0.68,
with about 40 proglottids; Scolex = 3.8-5.5 long; Bothridia = 0.41 x 0.15;
Testes = 300-400 in number. Genital pores alternating irregularly.
VIII. Plates of Cestoda and Cestodaria (Figs. 5-1 to 5-44)
LO
BO
5-1
5-2 A
TE
VG
SV
VA
GP 5-3
CS
OV
5-2 B
5-4
5-5
5-1. Eutetrarhynchus araya (scolex); 5-2. Rhinebothroides moralarai: A. scolex; BO = bothridium; LO =

loculus; B. mature proglottid; CS = cirrus sac; GP = genital pore; OV = ovary; SV = seminal vesicle;
TE = testes; VA = vagina; VG = vitelline glands; 5-3. Endorchis piraeeba (scolex); 5-4. Nominoscolex
woodlandi (scolex); 5-5. Acanthobothrium regoi (scolex).
5-6 5-7 5-8
5-9
5-10 5-11
5-12 5-13 5-14

5-6. Endorchis mandube (scolex); 5-7. Nominoscolex piracatinga (scolex); 5-8. Amphoteromorphus parkarmoo
(scolex); 5-9. Myzophorus admonticellia (scolex); 5-10. Ephedrocephalus microcephalus (scolex); 5-11.
Choanoscolex abscissum (scolex); 5-12. Manaosia bracodemoca (scolex); 5-13. Monticellia coryphicephala (scolex);
5-14. Nominoscolex dorad (scolex).
5-15 5-16
5-17 A 5-18
VG TE UT VA
5-17 B
CI
OV
5-15. Zygobothrium megacephalum (scolex); 5-16. Amphoteromorphus piraeba (scolex); 5-17. Amphoteromorphus
peniculus: A. scolex; B. mature proglottid; CI = cirrus; OV = ovary; TE = testes; UT = uterus; VA =
vagina; VG = vitelline glands; 5-18. Ornithoscolex lenha (scolex).
5-19 5-20
5-22
5-21
5-23 5-24 5-25

5-19. Woodlandiella myzophora (scolex); 5-20. Gibsoniela mandube (scolex); 5-21. Monticellia siluri; 5-22.
Peltidocotyle rugosa (scolex); 5-23. Monticellia rugosa (scolex); 5-24. Megathylacus jandia (scolex); 5-25.
Monticellia megacephala (scolex).
5-26 5-27 5-28
5-29 5-30 5-31
5-26. Proteocephalus jandia (scolex); 5-27. Proteocephalus fossatus (scolex); 5-28. Proteocephalus manjuariphilus
(scolex); 5-29. Proteocephalus macdonaghi (scolex); 5-30. Proteocephalus macropallus (entire; original); 5-31.
Sciadocephalus megalodiscus (entire; original).
TE
CS
TE VG
VG
VA
OV 5-33 A OV
CS
UE
5-33 B
5-32
5-34
5-35 5-36 5-37
5-32. Proteocephalus microscopicus (entire; original); OV = ovary; TE = testes; UE = uterus with eggs; VG
= vitelline glands; 5-33. Potamotrygonocestus orinocoensis: A. scolex; B. mature proglottid; CS = cirrus sac;
OV = ovary; TE = testes; VA = vagina; VG = vitelline glands; 5-34. Rudolphiella myoides (scolex); 5-35.
Rudolphiella piranabu (scolex); 5-36. Brayela karuatayi (scolex); 5-37. Anthobothrium pristis (scolex).
5-38 5-39
5-40
5-41
5-38. Proteocephalus sophiae: apical view of scolex (redrawn from CHAMBRIER & REGO 1994); 5-39.
Proteocephalus sophiae: lateral view of scolex (redrawn from CHAMBRIER & REGO 1994); 5-40. Proteocephalus
renaudi: scolex (redrawn from CHAMBRIER & VAUCHER 1994); 5-41. Proteocephalus soniae: scolex (redrawn
from CHAMBRIER & VAUCHER 1994).
5-42 A
VG
5-42 B
TE
5.0 mm
AO
5-43 A
UE
SR
OV
VA OT
5.0 mm
5-43 B
5-42. Schizochoerus liguloideus: A. anterior end; B. posterior end; OT = ootype; OV = ovary; SR =

seminal receptacle; TE = testes; UE = uterus with eggs; VA = vagina; VG = vitelline glands; 5-43.
Nesolecithus janicki: A. anterior end; AO = apical organ; B. posterior end.
AO
5-44
UE
VG
TE
SR
OV
5-44. Nesolecithus janicki: photograph of an entire stained specimen; AO = apical organ; UE = uterus
with eggs; VG = vitelline glands; TE = testes; OV = ovary; SR = seminal receptacle.
BEKÉSI, L., FEITOSA, V.A. & F.A.B. CABRAL (1992): Metacestodosis caused by plerocercoids of
Proteocephalidea (Cestoda) in fish fry cultured in large scale in the Brazilian Northeast. -
Parasitol. Hung. 25: 9-13.
BROOKS, D.R. (1995): Neotropical freshwater stingrays and their parasites: a tale of an ocean and a
river long ago. - J. Aquaculture Aquat. Sci. 7: 52-61.
BROOKS, D.R. & J.F.R. AMATO (1992): Cestodes parasites in Potamotrygon motoro (Chondrichthyes:
Potamotrigonidae) from Southwestern Brazil, including Rhinebothrioides melennanae sp. n.
(Tetraphyllidea: Phyllobothriidae), and a revised host-parasite checklist for helminthes
inhabiting Neotropical freshwater stingrays. - J. Parasitol. 78: 393-398.
BROOKS, D.R. & T.L. DEARDORFF (1980): Three proteocephalid cestodes from Colombian
siluriform fishes, including Nomimoscolex alovarius sp. n. (Monticellidae: Zygobothriinae). -
Proc. Helminthol. Soc. Wash. 47: 15-21.
BROOKS, D.R. & G. RASMUSSEN (1984): Proteocephalidean cestodes from Venezuelan siluriform
fishes with a revised classification of the Monticellidae. - Proc. Biol. Soc. Wash. 97: 748-760.
BROOKS, D.R. & T.B. THORSON (1976): Two tetraphyllidean cestodes from the freshwater stingray
Potamotrygon magdalenae DUMERIL, 1852 (Chondrichthyes: Potamotrygonidae) from Colombia. - J.
Parasitol. 62(6): 943-947.
BROOKS, D.R., MAYES, M.A. & T.B. THORSON (1981): Systematic review of cestodes infecting
freshwater stingrays (Chondrichtyes: Potomotrygonidae) including four new species from
Venezuela. - Proc. Helminthol. Soc. Wash. 48(1): 43-64.
CAMPBELL, R.A., MARQUES, F. & V.A. IVANOV (1999): Paroncomegas araya (WOODLAND, 1934) n. gen.
et comb. (Cestoda: Trypanorhyncha: Eutetrarhynchidae) from the freshwater stingray Potamo-
trygon motoro in South America. - J. Parasitol. 85(2): 313-320.
CHAMBRIER, A. DE & A.A. REGO (1994): Proteocephalus sophiae n. sp. (Cestoda: Proteocephalidae), a
parasite of the siluroid fish Paulicea luetkeni (Pisces: Pimelodidae) from the Brazilian Amazon.
- Rev. Suis. Zool. 101(2): 361-368.
CHAMBRIER, A. DE & C. VAUCHER (1992): Nomimoscolex touzeti n. sp. (Cestoda), a parasite of
Ceratophrys cornuta (L.); first record of a MonticelIidae in an amphibian host. - Mem. Inst.
Oswaldo Cruz: 87, Suppl.1: 61-67.
CHAMBRIER, A. DE & C. VAUCHER (1994): Etude morpho-anatomique et génétique de deux
nouveaux Proteocephalus WEINLAND, 1858 (Cestoda: Proteocephalidae) parasites de Platydoras
costatus (L.), poisson siluriforme du Paraguay. - Syst. Parasitol. 27: 173-185.
DÍAZ-UNGRÍA, C. (1973): Helmintos endoparásitos de Venezuela. - Cien. Vet. Maracaibo 3(1-2): 37-244.
DÖNGUES, J. & W. HARDER (1966): Nesolecithus africanus n. sp. (Cestodaria, Amphilinidae) aus dem
Coelom von Gymnarchus niloticus CUVIER 1829 (Teleostei). - Z. Parasitenkde. 28: 125-141.
KHALIL, L.F., JONES, A. & R.A. BRAY (1994): Keys to the cestode parasites of vertebrates. - CAB
International, Wallingford, Oxon: 751 pp.
MAYES, M.A., BROOKS, D.R. & T.B. THORSON (1978): Two new species of Acanthobothrium VAN
BENEDEN, 1848 (Cestoidea, Tetraphyllidea) from freshwater stingrays in South America. - J.
Parasitol. 64(5): 838-841.
PERTIERRA, A.A.G. (1995): Nomimoscolex microacetabula sp. n. y N. pimelodi sp. n. (Cestoda:
Proteocephalidea), parásitos de Siluriformes del Rio de La Plata. - Neotrópica 41: 19-25.
PERTIERRA, A.A.G. (2002): Redescription of Proteocephalus bagri and P. rhamdiae (Cestoda: Proteo-
cephalidae), parasites of Rhamdia quelen (Siluriformes: Pimelodidae) from South America, with
comments on morphological variation. - Folia Parasitologica 49: 55-60.
PERTIERRA, A.A.G. & G. VIOZZI (1999): Redescription of Cangatiella macdonaghi (SZIDAT & NANI,
1951) comb. nov. (Cestoda: Proteocephalidae) a parasitic of the Atheriniform fish Odontesthes
hatcheri (EIGENMANN, 1909) from the Patagonian region of Argentina. - Neotropica 45(113-
114): 13-20.
REGO, A.A. (1979): Contribuição ao conhecimento dos helmintos de raias fluviais Paratrygonidae. -
Rev. Brasil. Biol. 39(4): 879-890.
REGO, A.A. (1980): Encontro de plerocercos de Trypanorhyncha (Cestoda) em ofídio de rio da
América do Sul. - Mem. Inst. Butantan 44/45: 239-243.
REGO, A.A. (1982): Identificação de Temnocephalida (Turbellaria) e de plerocercos de Trypanorhyn-
cha (Cestoda) encontrados em caranguejos de água doce. - Rev. Brasil. Biol. 42(2): 275-278.
REGO, A.A. (1984): Proteocefalídeos (Cestoda) de Phractocephalus hemilopterus, peixe da Amazônia. -
Mem. Inst. Oswaldo Cruz 79(2): 257-261.
REGO, A.A. (1985): Proteocephalidea from Amazonian freshwater fishes: new systematic arrange-
ment for the species described by WOODLAND as Anthobothrium (Tetraphyllidea). - Parasitol. al
Día (Santiago, Chile) 9: 4-9.
REGO, A.A. (1987a): Redescription de Pterobothrium crassicolle DIESING, 1850 (Cestoda : Trypano-
rhyncha) e revalidação da espécie. - Mem. Inst. Oswaldo Cruz 82: 51-52.
REGO, A.A. (1987b): Cestóides proteocefalídeos do Brasil. Reorganização taxonômica. - Rev. Brasil.
Biol. 47: 203-212.
REGO, A.A. (1997): Senga sp., occurrence of a pseudophyllid cestode in a Brazilian freshwater fish. -
Mem. Inst. Oswaldo Cruz 92: 607.
REGO, A.A., CHUBB, J.C. & G.C. PAVANELLI (1999): Cestodes in South American freshwater teleost
fishes: keys to genera and brief description of species. - Rev. Brasil. Zool. 16: 299-367.
REGO, A.A. & A.P. LUNA DIAS (1976): Estudos de cestóides de peixes do Brasil. 3a Nota: Cestóides
de raias fluviais Paratrygonidae. - Rev. Brasil. Biol. 36(4): 941-956.
REGO, A.A. & G.C. PAVANELLI (1990): Novas espécies de cestóides proteocefalídeos parasitas de
peixes não siluriformes. - Ver. Brasil. Biol. 50: 91-101.
REGO, A.A., SANTOS, J.C. DOS & P.P. SILVA (1974): Estudos de cestóides de peixes do Brasil. - Mem.
Inst. Oswaldo Cruz 72(3/4): 187-204.
SKRJABIN, K.I. (ed.) (1965): Osnovi Tsestodologii, Tom. V.: V.I. FREZE: Proteotsefalyati lentochni
gelminti rib, amfibii I reptilii. - Akad. Nauk. U.S.S.R., Moscow: 538 pp.
STUNKARD, H.W. (1962): The organization, ontogeny and orientation of the Cestoda. - Quart. Rev.
Biol. 37(1): 23-34.
WARDLE, R.A. & J.A. MCLEOD (1952): The zoology of tapeworms. - Univ. of Minnesota Press,
Minneapolis: 780 pp.
WARDLE, R.A., MCLEOD, J.A. & RADINOVSKY (1974): Advances in the zoology of tapeworms. -
Univ. of Minnesota Press, Minneapolis: 274 pp.
WOODLAND, W.N.F. (1925): On three new proteocephalids (Cestoda) and a revision of the genera
of the family. - Parasit. 17(4): 370-394.
WOODLAND, W.N.F. (1933a): On the anatomy of some fish cestodes described by DIESING from
the Amazon. - Quart. J. Microsc. Sci. 76: 175-208.
WOODLAND, W.N.F. (1933b): On two new cestodes from the Amazon siluroid fish Brachyplatystoma
vaillanti CUV. - Parasit. 25(4): 486-490.
WOODLAND, W.N.F. (1933c): On a new subfamily of proteocephalid cestodes - the Othinoscoleci-
nae - from the Amazon siluroid Platystomatichthys sturio (KNER). - Parasit. 25(4): 491-500.
WOODLAND, W.N.F. (1934a): On six new cestodes from Amazon fishes. - Proc. Zool. Soc. London
1: 33-44.
WOODLAND, W.N.F. (1934b): On the Amphilaphorchidinae, a new subfamily of proteocephalid
cestodes, and Myzophorus admonticellia, gen. et sp. n., parasitic in Pirinampus spp. from the
Amazon. - Parasit. 26(1): 141-149.
WOODLAND, W.N.F. (1934c): On some remarkable new cestodes from Amazon siluroid fish. -
Parasit. 27(2): 208-225.
WOODLAND, W.N.F. (1935a): Some new proteocephalids and a ptychobothriid (Cestoda) from the
Amazon. - Proc. Zool. Soc. London 3: 619-624.
WOODLAND, W.N.F. (1935b): Some more remarkable cestodes from Amazon siluroid fish. - Parasit.
28(2): 207-225.
WOODLAND, W.N.F. (1935c): Additional cestodes from the Amazon siluroids, pirarara, dorad and
sudobim. - Proc. Zool. Soc. London 4: 851-862.
YAMAGUTI, S. (1959): Systema Helminthum. Vol. 2. The Cestodes of Vertebrates. - Intersci. Publ.,
New York: 860 pp.
6.
NEMATODA
Nematodes are elongate cylindrical worms that are found as free living forms in water
and soil as well as parasitic invaders of plants and animals. The general nematode body
plan is comparable to a tube within a tube. The inner tube is a complete digestive tract
and the outer is the body wall. The reproductive organs are also long and tubular and
are situated in the body cavity between the digestive tract and the body wall. The
nematode digestive system consists of two or three lips, an oral aperture, buccal cavity,
esophagus, intestine and anus.
In nematodes, the sexes are separate and the male structures are: a testis, vas
deferens, seminal vesicle and copulatory spicules. The female system includes: one or two
ovaries with accompanying oviducts and uteri, followed by a vagina, genital pore and
vulva. The male reproductive system normally opens by way of the anus, but the female
pore can be at any level from the esophagus to the anus.
Adult nematodes in fish live either in the digestive tract or in the body cavities.
Fertilization is internal and is accomplished by the male inserting the spicules into the
vagina of the female. Male nematodes usually have papillae, a bursa or a genital sucker
on the tail to aid in grasping the female. Some also have an internal hard structure
(gubernaculum) to guide the spicules. Ova are fertilized within the female uterus and
embryonic development begins. Nematodes grow by periodically moulting the outer
cuticle, as do arthropods. There are four larval stages prior to the adult stage, In the case
of fish nematodes; the first larval stage may be temporarily free in the water and all
other stages parasitic.
In the genera Rondonia and Monhysterides (Atractidae), the females are live-bearing and
give birth to fourth stage larvae or subadults. Eggs are not even formed in these nematodes
and the embryos and larval stages obtain nourishment from the uterine walls. Transmission
of these nematodes is apparently direct and no intermediate hosts are required.
Nematodes of the families Camallanidae, Cucullanidae and Philometridae are
also viviparous, but a microcrustacean (usually a copepod) is needed as an intermedi-
ate host in the cycle. Copepods ingest the actively swimming first larval stage of these
worms and the larvae develop to the third or fourth stage in the hemocoel of the
crustaceans. Fish become infected when they ingest copepods containing third or
fourth stage larvae.
Some Ascaroidea of fish require two intermediate hosts to complete their cycles.
The first of these is a microcrustacean and the second a small fish. In such cases, the
larvae do not mature in the first fish but penetrate into its tissues where they wait for this
host to be eaten by a larger predatory fish. An example of such a cycle is that of
Porrocaecum draschei (= Terranova serrata) which is an intestinal parasite of Arapaima gigas in
the Amazon River system. Probably other fish nematodes have similar cycles but these
have not been studied. Amazon fish commonly have numerous encysted nematode larvae
in their tissues (especially in the mesenteries). Some of these represent species of
Contracaecum, Multicaecum, Terranova, Dujardinascaris and Eustrongylides that mature in
piscivorous vertebrates other than fish (especially birds and crocodilians).
III. Pathology
The damage caused to fish by nematodes varies greatly depending on the species
present, the organ invaded and the number of worms involved. The oxyuroids,
Monhysterides iheringi and Rondonia rondoni, are often found in countless thousands in the
intestinal tracts of Amazonian fish, especially catfish and “pacus” but they do not appear
to cause any histological alterations. The relationship of these nematodes to their hosts
seems to be more in the nature of commensalism. They do not attach to the intestinal
wall, but maintain their position by active swimming.
Nematodes that attach to the intestinal wall or bury their heads in it are more
pathogenic. Several species of Spirocamallanus are common in Neotropical fish, and
these draw the host’s intestinal wall into their buccal capsules and feed on blood.
There is always a localized inflammatory reaction at the attachment site and the
worms can probably cause primary anemia from blood loss. Camallanids can also
cause intestinal blockage, especially in small fish. We have found a 90 % incidence of
Procamallanus (S.) inopinatus in young specimens of Brycon erythropterus (from 4-11 cm in
length) that were caught in the Amazon River near Manaus, Brazil. In these infections,
there were only one or two worms present, but in the smaller fish, they nearly filled
the intestinal lumen. If such infections do not immediately cause the death of the
fish, they certainly must affect their growth rates.
In general, nematodes with cuticular spines invade the intestinal mucosa of the host.
Species of Spinitectus insinuate all or most of the body into the intestinal wall where they
cause severe inflammation. Species of Echinocephalus have spherical head region covered
with small spines. This head bulb is inserted into the intestinal mucosa of stingrays. This
provokes localized inflammation leading to fibrous encapsulation of the head bulb.
Another spinous nematode that penetrates into the intestinal mucosa is Goezia
spinulosa, an ascaridoid. TEXEIRA DE FREITAS & LENT (1946) reported that in Ceará State,
Brazil, this species caused the death of larvae and young of Astronotus ocellatus and
Arapaima gigas in a pisciculture station. The problem became so severe that attempts to
rear these two species at the station were suspended.
In addition to the effect of nematodes on fish health, these worms cause
considerable economic loss in the fisheries industry. Since many nematode larvae are large
enough to be seen by the unaided aye, their presence can make fish unmarketable. Also,
health authorities sometimes prohibit the sale of fish containing visible larvae. As an
example, Eustrongylides larvae are bright red in color and attain a length of 8 to 10 cm.
They normally mature in piscivorous birds, and can survive in man. In the Brazilian
Amazon, such larvae are often found in the flesh of the important food fish, Cichla
ocellaris. Inspectors frequently condemn such fish for human consumption, but in any case,
they have little appeal to the consumer.
No treatment for encysted nematode larvae is possible or necessary. Since transmission of

nematodes is usually by way of microcrustacean intermediate hosts, these should be
eliminated from aquaria, when possible. Adult and larval nematodes in the intestinal
lumen can be removed by treatment with D-N-Butyl tin oxide, which is mixed with the
ration at 0.3 % of body weight and given for from one to five days.
Nematodes can be collected by hand sedimentation of the intestinal contents, as

described for trematodes. They can be killed, fixed and preserved by simply dropping then
into 70 % alcohol. If storage for long periods is contemplated, 80 % alcohol to which 10
% glycerin has been added can be used. The glycerin keeps the specimens from drying out
in the event that the alcohol evaporates.
Temporary study slides can be made of nematodes in either glycerin or phenol. For
glycerin mounts, the worms should first be hydrated by passage through water. If phenol
is to be used, the worms should be partially dehydrated in 95 % alcohol first.
If permanent stained preparations are to be made of nematodes, or any other
small invertebrates, they should be killed and fixed first in AFA (85 parts of 85 %
alcohol: 10 parts commercial formalin: 5 parts glacial acetic acid). Next, pass the
specimen directly from AFA to the stain solution (95 % alcohol colored to the
intensity of weak tea with equal parts of eosin and orange-g stains). Stain in this
solution for 10 to 30 minutes and then move the specimens to pure phenol (liquefy
phenol crystals with a small amount of 96 % alcohol to make this solution). The
phenol simultaneously dehydrates, clears and destains the material. When a specimen
appears clear in the phenol, it is already dehydrated (this usually takes a few seconds
or at most a couple of minutes) but if more destaining is desired, it may be left in this
solution longer. After the proper degree of destaining is achieved, the specimen can
be moved to methyl salicylate where the destaining process will stop. After 3 minutes
in the latter, the specimens may be mounted in Canada balsam. Material fixed by
other means and stored in 70 % alcohol can be processed in this same way, but
specimens in aqueous solutions (10 % formalin, for example) must be placed in 70 %
alcohol for a few minutes before staining.
It often happens that a nematode, or other invertebrate, is fixed in an undesirable
position. Regardless of the fixative used, the position can be corrected to some extent
because a specimen in phenol becomes soft and pliable. It can be taken from that liquid,
placed on a dry slide and manipulated into a better position with dissecting needles. When
the desired position is achieved, a coverglass is placed over the specimen to secure it and
methyl salicylate is added. The latter hardens the specimen in a few seconds and it will
retain this position after mounting in balsam.
Using the method described above, it is relatively easy to make permanent “en face”
mounts of nematodes. Place a drop of Canada balsam on a slide. Remove a cleared
nematode from the methyl salicylate and place it on the slide near the drop of balsam.
Then, simply cut off the head of the nematode and shove it into the drop ( micro-scalpel
or razor blade may be used). Finally, adjust in a vertical position and place a coverglass on
top. For permanence, all balsam slides should be dried in an oven at 56 °C for two weeks.
Morphological features used in nematode identification are: 1) size and shape of the body,
2) cuticular ornamentation, 3) form of the mouth and lips, 4) form and size of the buccal
capsule and esophagus, 5) shape of the tail in both sexes, 6) position of the vulva, 7) form
and size of the male spicules, and 8) number and position of the caudal papillae in the
male. Some species can only be separated on the basis of male spicule morphology and
papilla number.
Key to the superfamilies of Nematoda from Amazonian freshwater fishes
238
I. Esophagus a stichosome (surrounded by unicellular glands) ...................................................................................................... Trichinelloidea

II. Esophagus not a stichosome.
A. Three large lips present ..................................................................................................................................................................... Ascaridoidea
B. Lips small or lacking
1. Esophagus provided with a posterior muscular bulb ............................................................................................................ Oxyuroidea
2. Esophagus divided into anterior muscular and posterior glandular regions.
a. Oviparus nematodes ............................................................................................................................................................. Seuratoidea
b. Ovoviviparous nematodes ................................................................................................................................................ Camallanoidea
c. Viviparous nematodes
1) Parasites of connective tissue and body cavity ....................................................................................................... Dranculoidea
2) Parasites of the intestinal tract ............................................................................................................................. Cosmocercoidea
C. Pseudolabia present
1. Cuticle spinous anteriorly ...............................................................................................................................................Gnathostomatoidea
2. Cuticle smooth.
a. Two large pseudolabia present .................................................................................................................................... Physalopteroidea
b. Two small pseudolabia present .................................................................................................................................. Habronematoidea
D. Oral cavity present, simple, without lips ........................................................................................................................................ Thelazioidea
The genera and species included herein are those that have been reported from the Amazon region. Other species that may occur
there are also mentioned. For complete information on the nematodes of freshwater fishes of the Neotropical region, the reader is
referred to the classic work of MORAVEC (1998).
Key to the Ascaridoidea of Amazonian freshwater fishes
I. Esophagus widened posteriorly; ventriculus and 1 or 2 posteriorly directed ventricular appendices usually present ........... Anisakidae
A. Body covered with circular rows of spines ................................................................................................................. Goezia (Fig. 6-21 A-B)
B. Body unspined; mouth large, lips prominent.
1. Interlabia small or absent.
a. Ventricular appendix present; intestinal cecum absent ............................................. Raphidascaris (Sprentascaris) (Fig. 6-20 A-C)
b. Ventricular appendix absent; intestinal cecum present ............................................................................ Terranova (Fig. 6-12 A-B)
2. Interlabia large ................................................................................................................... Raphidascaroides (Figs. 6-20 A-C, 6-60 & 6-61)
II. Esophagus elongate, not widened posteriorly; ventriculus present with 2 anteriorly and 3 posteriorly directed appendices ..................
........................................................................................................................... Acanthocheilidae ...................................................... Brevimulticaecum
Key to the Oxyuroidea (Pharyngodonidae) of Amazonian freshwater fishes
I. Esophagus short, stout; bulb without sclerotized apparatus ................................................................................................ Travnema (Fig. 6-9)
II. Esophagus elongate; bulb with sclerotized apparatus.
A. Body of esophagus expanded anterior to bulb.

1. Buccal capsule present, armed with teeth ............................................................................................................. Cosmoxynema (Fig. 6-6)
2. Buccal capsule absent ........................................................................................................................................... Cosmoxynemoides (Fig. 6-4)
B. Body of esophagus not expanded anterior to bulb.
1. Caudal appendage of male without papillae; caudal alae present ............................................................................................ Ichthyouris
2. Caudal appendage of male with 1 or 2 papillae; caudal alae present or absent..
a. Buccal capsule absent; parasites of Loricariidae ......................................................................................................... Parasynodontisia
b. Buccal capsule present, armed with 3 teeth; parasites of Pimelodidae ........................................................................... Brasilnema
c. Buccal capsule present, without teeth; parasites of Doradidae ....................................................................................... Spinoxyuris
Key to the Camallanoidea (Camallanidae) of Amazonian freshwater fishes
I. Buccal capsule formed from 2 lateral valves ..................................................................................................................................... Camallaninae

A. Valves divided into anterior and posterior parts of similar size .......................................... Paracamallanus (Figs. 6-30 & 6-39 to 6-42)
239
B. Valves undivided, terminating posteriorly in a chitinized ring ................................................... Camallanus (Figs. 6-25 & 6-31 to 6-35)
240
II. Buccal capsule cup-like ..................................................................................................................................................................... Procamallaninae
A. Inner surface of buccal capsule smooth in both sexes ............................................................... Procamallanus (Procamallanus) (Fig. 6-28)

B. Inner surface of buccal capsule with spiraling ridges in both sexes ...............................................................................................................
............................................................................................................................. Procamallanus (Spirocamallanus) Figs. 6-24 A-B, 6-26 & 6-27)
C. Inner surface of buccal capsule of female with spiraling ridges; buccal capsule of male without ridges but with teeth ..................
................................................................................................................................................ Procamallanus (Denticamallanus) (Figs. 6-81& 6-87)
Key to the Cosmocercoidea of Amazonian freshwater fishes
I. Viviparous nematodes with from 1 to 10 formed larvae in the uterus ............................................................................................. Atractidae
A. Head end armed with sclerotized pieces ....................................................................................... Klossinemella (Figs. 6-43 to 6-49 & 6-78)
B. Head end without sclerotized pieces ............................................................................................................ Rondonia (Figs. 6-5 & 6-23 A-B)
II. Oviparous nematodes ............................................................................................................................................................................... Kathlaniidae
A. Mouth with 3 lips; cephalic end separated from rest of body by a cuticular fold at level of nerve ring ...............................................
................................................................................................................................................................................ Myleusnema (Figs. 6-64 to 6-77)
B. Mouth with 6 lips; cephalic end not separated from rest of body ........................................................................ Spectatus (Fig. 6-8 A-B)
C. Pharyngeal part of esophagus armed with cuticular ring ............................................................................. Chabaudinema (Fig. 6-19 A-C)
Key to the Dracunculoidea of Amazonian freshwater fishes
I. Esophagus divided into anterior muscular and posterior glandular parts; glandular part long or stout ............................. Guyanemidae
A. Glandular esophagus long, slender, without posterior appendix ....................................................................... Guyanema (Fig. 6-18 A-B)
B. Glandular esophagus short and stout, with elongate posterior appendix .............................................................................. Travassosnema
II. Esophagus not divided or glandular part very short ...................................................................................................................... Philometridae
A. Cuticle of adult females with small projections or bosses.
1. Bosses rounded ............................................................................................................................................................................. Philometroides
2. Bosses conical ................................................................................................................................................................... Nilonema (Fig. 6-14)
B. Cuticle of adult females smooth.
1. Esophagus expanded anteriorly .......................................................................................................................................................Philometra
2. Esophagus not expanded anteriorly ...................................................................................................................................................... Rumai
Key to the Seuratoidea of Amazonian freshwater fishes
I. Buccal cavity absent ................................................................................................................................................................................ Quimperiidae

A. Anterior end of esophagus expanded.
1. Esophagus divided into two parts; mouth without lips .................................................................................................................. Touzeta
2. Esophagus undivided; mouth with 6 lips .................................................................................................... Paraseuratum (Fig. 6-17 A-B)
B. Anterior end of esophagus not expanded ................................................................................................................................ Neoparaseuratum
II. Buccal cavity present ............................................................................................................................................................................... Cucullanidae
A. Intestinal cecum present ............................................................................................................................................................................Dichelyne

B. Intestinal cecum absent.
1. Caudal extremity conical or pointed ................................................................................................. Cucullanus (Figs. 6-16 A-B & 6-38)
2. Caudal extremity rounded ....................................................................................................................... Neocucullanus (Figs. 6-13 & 6-15)
Key to the Trichinelloidea (Capillariidae) of Amazonian freshwater fishes
I. Spicular sheath of male spinous; stichosome composed of a single row of stichocytes ................................................................ Capillaria
II. Spicular sheath of male not spinous.
A. Spicule absent or very short ........................................................................................................................................................... Freitascapillaria
B. Spicule present, well developed.
Tail of male with membranous bursa supported by two lateral lobes.
1. Caudal lobes of male narrow, finger-shaped ................................................................................................ Paracapillaria (Fig. 6-1 A-B)
2. Caudal lobes of male wide, nearly spherical ................................................................................................................... Capillostrongyloides
241
Key to the Gnathostomatoidea (Gnathostomatidae) of Amazonian freshwater fishes
242
The only species of this family shown to occur in Neotropical fishes is ..................................................Echinocephalus daileyi (Fig. 6-11 A-D)

Key to the Thelazioidea (Rhabdochonidae) of Amazonian freshwater fishes
The only genus of this family known from Neotropical fishes is ................................................................................ Rhabdochona (Fig. 6-2 A-B)
Key to the Physalopteroidea (Physalopteridae) of Amazonian freshwater fishes
The only genus of this family reported from Neotropical freshwater fishes is ..................................................................................... Heliconema
Key to the Habronematoidea of Amazonian freshwater fishes
Only two genera of this Superfamily are known from Neotropical freshwater fishes:
I. Cuticle with circular rows of spines .............................................................................................................................. Spinitectus (Fig. 6-10 A-D)
II. Cuticle smooth ................................................................................................................................................................ Cystidicoloides (Fig. 6-3 A-B)
Clave para superfamílias de Nematoda de peces Amazonicos de agua dulce
I. Esófago un esticosoma (rodeado por glándulas unicelulares) ..................................................................................................... Trichinelloidea

II. Esófago no es un esticosoma.
A. Tres grandes labios presentes .......................................................................................................................................................... Ascaridoidea
B. Labios pequeños o ausentes
1. Esófago provisto con un bulbo muscular posterior ............................................................................................................... Oxyuroidea
2. Esófago dividido en dos regiones siendo la anterior muscular y la posterior glandular.
a. Nemátodos oviparos ............................................................................................................................................................... Seuratoidea
b. Nemátodos ovoviviparos .................................................................................................................................................. Camallanoidea
c. Nemátodos vivíparos
1) Parásitos del tejido conectivo y cavidad del cuerpo ............................................................................................... Dranculoidea
2) Parásitos del tracto intestinal ................................................................................................................................. Cosmocercoidea
C. Pseudolabios presentes
1. Cutíula espinosa anteriormente ....................................................................................................................................Gnathostomatoidea
2. Cutícula lisa.
a. Com dos pseudolabios grandes ................................................................................................................................... Physalopteroidea
b. Con dos pseudolabios pequeños ............................................................................................................................... Habronematoidea
D. Cavidad oral presente, simple, sin labios ........................................................................................................................................ Thelazioidea
Los géneros y especies incluídos aquí son solo aquellos reportados para la región Amazónica. Otras especies que pudiesen ocurrir
allí también están mencionadas. Para una información completa sobre los nemátodos de agua dulce en la región Neotropical, el lector

puede consutar el trabajo clásico de MORAVEC (1998).
Clave para las Ascaridoidea de peces Amazonicos de agua dulce
I. Esófago ensanchado posteriormente; ventrículo y 1 o 2 apéndices ventriculares dirigidos posteriormente generalmente presentes ...
.......................................................................................................................................................................................................................... Anisakidae
A. Cuerpo cubierto con hileras circulares de espinas ..................................................................................................... Goezia (Fig. 6-21 A-B)
B. Cuerpo sin espinas; boca grande, labios prominentes.
1. Interlabios pequeños o ausentes.
a. Apéndice ventricular presente; cecum intestinal ausente ......................................... Raphidascaris (Sprentascaris) (Fig. 6-20 A-C)
b. Apéndice ventricular ausente; cecum intestinal presente ........................................................................ Terranova (Fig. 6-12 A-B)
2. Interlabios grandes ........................................................................................................... Raphidascaroides (Figs. 6-20 A-C, 6-60 & 6-61)
243
II. Esófago alargado, no se engrosa posteriormente; ventrículo presente con 2 apéndices dirigidos anteriormente y 3 apéndices dirigidos
244
posteriormente ............................................................................................... Acanthocheilidae ........................................................ Bevimulticaecum

Clave para las Oxyuroidea (Pharyngodonidae) de peces Amazonicos de agua dulce
I. Esófago corto, robusto; bulbo sin aparato esclerotizado ...................................................................................................... Travnema (Fig. 6-9)
II. Esófago alargado; bulbo con aparato esclerotizado.
A. Cuerpo del esófago expandido anterior al bulbo.
1. Cápsula bucal presente, armada con dientes ........................................................................................................ Cosmoxynema (Fig. 6-6)
2. Cápsula bucal ausente ........................................................................................................................................... Cosmoxynemoides (Fig. 6-4)
B. Cuerpo del esófago sin expansión anterior al bulbo.
1. Apéndice caudal del macho sin papilas; alas caudales presentes .............................................................................................. Ichthyouris
2. Apéndice caudal del macho con 1 o 2 papilas; alas caudales presentes o ausentes.
a. Cápsula bucal ausente; parásitos de Loricariidae ........................................................................................................ Parasynodontisia
b. Cápsula bucal presente, armada con 3 dientes; parásitos de Pimelodidae ..................................................................... Brasilnema
c. Cápsula boca presente, sin dientes; parásitos de Doradidae ............................................................................................. Spinoxyuris
Clave para las Camallanoidea (Camallanidae) de peces Amazonicos de agua dulce
I. Cápsula bucal formada de 2 válvulas laterales .................................................................................................................................. Camallaninae

A. Válvulas divididas en parte anterior y posterior de tamaños similares ............................... Paracamallanus (Figs. 6-30 & 6-39 to 6-42)
B. Válvulas sin division, terminando posteriormente en un anillo quitinizado ........................... Camallanus (Figs. 6-25 & 6-31 to 6-35)
II. Cápsula bucal en forma de copa ..................................................................................................................................................... Procamallaninae
A. Superficie interna de la cápsula bucal, lisa en ambos sexos ....................................................... Procamallanus (Procamallanus) (Fig. 6-28)
B. Superficie interna de la cápsula bucal con salientes en espiral en ambos sexos ...........................................................................................
............................................................................................................................. Procamallanus (Spirocamallanus) Figs. 6-24 A-B, 6-26 & 6-27)
C. Superficie interna de la cápsula bucal de la hembra con protuberancias en espiral; cápsula bucal del macho sin protuberancias pero
con dientes ......................................................................................................................... Procamallanus (Denticamallanus) (Figs. 6-81 & 6-87)
Clave para las Cosmocercoidea de peces Amazonicos de agua dulce
I. Nemátodos vivíparos con 1 a 10 larvas formadas en el útero ............................................................................................................ Atractidae

A. Extremidad anterior armada con piezas esclerotizadas ............................................................... Klossinemella (Figs. 6-43 a 6-49 & 6-78)
B. Extremidad anterior sin piezas esclerotizadas ............................................................................................ Rondonia (Figs. 6-5 & 6-23 A-B)
II. Nemátodos ovíparos ................................................................................................................................................................................. Kathlaniidae
A. Boca con 3 labios; extremidad cefálica separada del resto del cuerpo por un doblez cuticular a nivel del cordon nervisoso ..........
.................................................................................................................................................................................. Myleusnema (Figs. 6-64 a 6-77)
B. Boca con 6 labios; extremidad cefálica sin separación del resto del cuerpo ....................................................... Spectatus (Fig. 6-8 A-B)
C. Parte faríngea del esófago armada con anillo cuticular ................................................................................ Chabaudinema (Fig. 6-19 A-C)

Clave para las Dracunculoidea de peces Amazonicos de agua dulce
I. Esófago dividido en parte anterior musculosa y parte posterior glandular; parte glandular larga o robusta .................... Guyanemidae
A. Esófago glandular largo, delgado, sin apéndice posterior ................................................................................... Guyanema (Fig. 6-18 A-B)
B. Esófago glandular corto y robusto, con apéndice posterior alargado .................................................................................... Travassosnema
II. Esófago sin divisió o parte glandular muy corta ............................................................................................................................. Philometridae
A. Cutícula de hembras adultas con proyecciones pequeñas o abultadas.
1. Abultamientos redondeados ....................................................................................................................................................... Philometroides
2. Abultamientos cónicos ................................................................................................................................................... Nilonema (Fig. 6-14)
B. Cutícula de hembras adultas lisas.
1. Esófago expandido anteriormente ..................................................................................................................................................Philometra
2. Esófato sin expansión anteiror .............................................................................................................................................................. Rumai
245
Clave para las Seuratoidea de peces Amazonicos de agua dulce
246
I. Cavidad bucal ausente ............................................................................................................................................................................ Quimperiidae

A. Terminal anterior del esófago expandido.
1. Esófago dividido en dos partes; boca sin labios ....................................................................................................................... Touzeta
2. Esófago no dividido; boca con 6 labios ................................................................................................ Paraseuratum (Fig. 6-17 A-B)
B. Terminal anterior del esófago no expandido ..................................................................................................................... Neoparaseuratum
II. Cavidad bucal presente ........................................................................................................................................................................... Cucullanidae
A. Ciego intestinal presente ......................................................................................................................................................................Dichelyne
B. Ciego intestinal ausente.
1. Extremidad caudal conica o puntiaguda ................................................................................... Cucullanus (Figs. 6-16 A-B & 6-38)
2. Extremidad caudal rounda ............................................................................................................... Neocucullanus (Figs. 6-13 & 6-15)
Clave para las Trichinelloidea (Capillariidae) de peces Amazonicos de agua dulce
I. Vaina espicular del macho espinosa; esticosoma compuesto de una única hilera de esticocistes ................................................. Capillaria
II. Vaina espicular del macho no espinosa.
A. Espículas ausentes o muy cortas ................................................................................................................................................... Freitascapillaria
B. Espículas presentes, bien desarrolladas.
Cola del macho con bursa membranosa sostenida por dos lóbulos laterales.
1. Lóbulos de la cauda del macho angostos, en forma de dedos ................................................................. Paracapillaria (Fig. 6-1 A-B)
2. Lóbulos del macho anchos, casi esféricos ....................................................................................................................... Capillostrongyloides
Clave para las Gnathostomatoidea (Gnathostomatidae) de peces Amazonicos de agua dulce
La única especie de esta familia encontrada en peces Neotropicales es ....................................................Echinocephalus daileyi (Fig. 6-11 A-D)
Clave para las Thelazioidea (Rhabdochonidae) de peces Amazonicos de agua dulce
El único género conocido de esta familia en peces Neotropicales es ......................................................................... Rhabdochona (Fig. 6-2 A-B)
Clave para las Physalopteroidea (Physalopteridae) de peces Amazonicos de agua dulce
El único género de esta familia reportado para peces Neotropicales de agua dulce es ........................................................................ Heliconema
Clave para las Habronematoidea de peces Amazonicos de agua dulce
Solamente dos géneros de esta Superfamilia son conocidos para preces Neotropicales de agua dulce:
I. Cutícula con hileras circulares de espinas .................................................................................................................... Spinitectus (Fig. 6-10 A-D)
II. Cutícula lisa ...................................................................................................................................................................... Cystidicoloides (Fig. 6-3 A-B)

247
VII. Checklist of Nematoda from Amazonian fishes
Atractidae TRAVASOS, 1919

Oxyuroidea. Small worms with esophagus divided into two parts; posterior
part usually bulbous. Intestine simple. Male: without preanal sucker;
spicules equal, or not; gubernaculum present or absent. Female: tail
pointed; ovary single; vulva posterior; viviparous. Intestine of vertebrates.
Klossinemella COSTA, 1961
Mouth with six small lips. Esophagus divided into two parts; enlarged at
posterior end, but without definite bulb. Tail of both sexes long and
pointed. Male: tail spirally coiled; with four pairs of preanal and five parts
of postanal papillae; spicules unequal, gubernaculum present. Female:
vulva close to anus; viviparous. Intestine of fishes.
K. iheringi (TRAVASSOS, ARTIGAS & PEREIRA, 1928): Piaractus brachypomum, Myleus
(Myloplus) asterias, Mylesinus paraschomburgkii, Leporinus copelandi, L. fasciatus,
Schizodon nasutus, Hoplias malabaricus and Pterodoras granulosus: Brazil (Ama-
zonia). (Figs. 6-7 & 6-43 to 6-49). Body = 2.4-3.5 x 0.09 mm (male); 4.1-4.2
x 0.8 mm (female); Esophagus = 200-210 x 24-32 µm (anterior part); 360-
410 x 48-68 µm (posterior part); Spicules = 88-120 x 17-18 µm.
Rondonia TRAVASSOS, 1919
Mouth with three bilobed lips, each with two papillae. Esophagus with
cylindrical anterior portion which expands posteriorly and terminates in a
bulb. Male: posterior extremity spiraled; spicules subequal; gubernaculum
present; one pair of preanal and four pairs of postanal papillae present.
Female: tail pointed, ovary single, vulva opening into cloaca; viviparous.
Digestive tract of fishes.
R. rondoni TRAVASSOS, 1919: Piaractus brachypomus, Colossoma metrei, Myleus spp.,
Mylesinus paraschomburgkii, Myletes torquatus, M. edulus, Pterodoras granulosus,
Trachydoras paraguayensis, Pimelodus clarias, Pseudopimelodus zungaro and Lu-
ciopimelodus pati: Brazil. (Figs. 6-5 & 6-23 A-B). Body = 6-6.5 x 0.30 mm
(male); 6.5-7.0 x 0.32 mm (female); Esophagus = 520-555 x 80-100 µm
(anterior part); 260-280 x 60-70 µm (posterior part, less bulb); Bulb = 140
µm in diam; Spicules = 100-200 µm; Gubernaculum = 60 µm.
Camallanidae RAILLIET & HENRY, 1915
Camallanoidea. Mouth without lips; buccal capsule prominent, chitinoid,
cup-like, or divided into two shell-like halves. Esophagus with anterior
muscular part and long posterior glandular part. Male: posterior extremity
curved ventrally; caudal alae present; papillae variable in number, frequent-
ly pedunculate; spicule unequal, dissimilar. Female: vulva near middle of
body; vagina directed posteriorly; two uteri present, viviparous. Parasites
of digestive tracts of cold-blooded vertebrates.
Camallanus RAILLIET & HENRY, 1915
Mouth formed of two lateral chitinous halves (valves) with longitudinal
rib-like thickenings internally; trident shaped chitinous processes project
posteriorly from junctions of valves; chitinous ring present between valves

and esophagus. Glandular portion of esophagus enlarged posteriorly.
Male: with about 7 pairs of preanal papillae, 2 pairs of adanal papillae and
several pairs of postanal papillae; spicules unequal and dissimilar, with one
of them lightly chitinized; gubernaculum absent. Female: uteri opposed;
posterior ovary absent; viviparous. In stomach, intestine or pyloric cecae
of fishes, amphibians and reptiles.
C. tridentatus (DRASCHE, 1884): Arapaima gigas: Brazil (Amazonia). (Fig. 6-32).
Body = 3.6 mm (male); 7-12 x 0.17 x 0.23 mm (female); Buccal valves =
62-75 µm (male); 110-120 x 170 µm (female); Spicules = 205 µm; Papillae
= 14 pairs (7 preanal, 2 adanal & 5 postanal).
C. acaudatus FERRAZ & THATCHER, 1990: Osteoglossum bicirrhosum: Brazil (Ama-
zonia). (Figs. 6-31 & 6-33 to 6-35). Body = 1.0-1.8 x 0.08-0.13 mm (male);
12-19 x 0.32-0.44 mm (female); Esophagus = 520-555 x 80-100 µm
(anterior part); 260-280 x 60-70 µm; Buccal valves = 50-65 x 57-107 µm
(male) & 67-95 x 75-107 µm (female); Spicules = 155-197 µm; Papillae =
14 pairs (7 preanal, 2 adanal & 5 postanal).
Paracamallanus YORK & MAPLESTONE, 1928
Mouth formed from 2 lateral chitinous valves with longitudinal rib-like
thickenings and marginal indentations; trident-like chitinous processes at
junction of valves on either side; large chitinous buccal cavity between
valves and esophagus. Intestine of fishes.
P. amazonensis FERRAZ & THATCHER, 1992: Hypophthalmus edentatus: Brazil
(Amazonia). (Figs. 6-30 & 6-39 to 6-42). Body = 2-3 x 0.07-0.10 mm
(male); 4-5 x 0.10-0.15 mm (female); One spicule present = 120-170 µm;
Papillae = 6 pairs (preanal).
Procamallanus (Denticamallanus) BAYLIS, 1923
Medium sized nematodes with almost smooth cuticle. Mouth opening
circular, surrounded by six low elevations (2 lateral and 4 submedian), eight
cephalic papillae in two circlets and two lateral amphids. Basal capsule
orange-brown, approximately as long as wide in male and distinctly shorter
than wide in female; basal ring indistinct. Buccal capsule surrounded by
outer, colorless hyaline layer. Muscular esophagus considerably shorter than
glandular one; both parts of esophagus expanded at their posterior half.
Minute deirids situated just below nerve ring level; excretory pore somewhat
posterior to nerve ring level. Tail conical, its tip bluntly pointed.
Procamallanus (D.) dentatus MORAVEC & THATCHER, 1997: Bryconops alburnoides:
Brazil (Urubu River, Amazonas State). (Figs. 6-81 to 6-87). Body = 5.84-
7.14 x 0.204-0.310 mm (male); 22.78-35.77 x 0.517-0.870 mm (female);
Buccal capsule = 78-105 x 81-108 µm (male) & 108-111 x 153 µm
(female); Spicules = 78 x 117.
Procamallanus (Procamallanus) BAYLIS, 1923
Mouth a circular opening in a large cup or chalice-shaped chitinous buccal
capsule; internal walls of capsule smooth. Male: caudal alae present,
uniting anteriorly; 3 – 10 pairs of preanal papillae present; spicules

unequal. Female posterior ovary lacking; vulva in midbody; viviparous.
Intestine of fishes.
Procamallanus (P.) peraccuratus MAGALHÂES PINTO, FABIO, NORONHA & ROLAS,
1976: Geophagus brasiliensis and Cichlasoma facetum: Brazil (Espírito Santo
State). (Fig. 6-28). Body = 9.4-9.8 x 0.15-0.17 mm (male); 12.8-22.3 x 0.21-
0.40 mm (female); Buccal capsule = 72-87 x 49 µm (male) & 87-110 x 52
µm (female); Spicules = 220 x 240 x 180-200.
Procamallanus (P.) annipetterae KOHN & FERNANDES, 1988: Hypostomus albopuncta-
tus: Brazil (Paraná State). Body = 9.69 mm (male); 21.8 mm (female);
Buccal capsule = 131 x 123 µm (male) & 180 x 187 µm (female); Spicules
= 21 & 16 µm; Papillae = 5 pairs (2 preanal, 2 adanal & 1 postanal).
Procamallanus (Spirocamallanus) BAYLIS, 1923
Buccal capsule with prominent spiraling ridges on inner walls, may also be
provided with teeth in the mouth opening or in the bottom of the cup.
Other characters as in Procamallanus. (note: This taxon is probably poly-
phyletic and may require division into several genera in the future).
Procamallanus (S.) amarali VAZ & PEREIRA, 1934: Leporinus sp.: Brazil (São Paulo
State). Body = 9.25 x 0.28 mm (male); Buccal capsule = 70 x 60 µm; Spicules
= 440 & 240 µm; Papillae = 14 pairs (8 preanal, 3 adanal & 3 postanal).
Procamallanus (S.) barroslimai PEREIRA, 1935; possibly Triportheus sp.: Brazil (São
Paulo State). Body = 3.1 x 0.14 mm (male); Buccal capsule = 30 x 40 µm
(male); Spicules = 50 & 60 µm; Papillae = 6 pairs (3 preanal & 3 postanal).
Procamallanus (S.) cearensis PEREIRA, VIANNA & AZEVEDO, 1936 (considered a
synonym of S. hilarii by KLOSS, 1966): Astyanax bimaculatus: Brazil (Ceará
State). Body = 4.0 x 0.18 mm (male) & 10.3-14.0 x 0.28-0.42 mm (female);
Buccal capsule = 40 µm (male) & 40-50 µm (female); Spicules = 57 µm;
Papillae = 7 pairs (4 preanal & 3 postanal).
Procamallanus (S.) chimusensis TEXEIRA DE FREITAS & IBAÑEZ, 1968: Pygidium
punctatum: Colômbia and Peru. Body = 12-13 x 0.24-0.25 mm (male) & 34-
37 x 0.65-0.67 mm (female); Buccal capsule = 78-83 x 58-67 µm (male) &
93 x 73-80 µm (female); Spicules = 220-230 & 160-170 µm; Papillae = 6
pairs (3 preanal & 3 postanal).
Procamallanus (S.) hilarii VAZ & PEREIRA, 1934: Salminus hilarii: Brazil (São Paulo
State), Peru and Venezuela. Body = 6 x 0.2 mm (male) & 14 x 0.36 mm
(female); Buccal capsule = 57 x 57 µm (male) & 62 x 62 µm (female);
Spicules = 82 & 62 µm; Papillae = 8 pairs (3 preanal, 2 adanal & 3 postanal).
Procamallanus (S.) iheringi TRAVASSOS, 1929: Salminus hilarii: Brazil (São Paulo
State). Body = 12 x 0.3 mm (male) & 16-21 x 0.5-0.8 mm (female); Buccal
capsule = 70-80 x 56-64 µm (male) & 88-96 x 70-80 µm (female); Spicules
= 240-250 & 520-530 µm.
Procamallanus (S.) incarocai TEXEIRA DE FREITAS & IBAÑEZ, 1970 (= syn. of
(P.(S). hilarii): Pygidium punctatum: Peru. Body = 6.6-6.9 x 0.12-0.16 mm
(male); 8.5-27 x 0.16-0.53 mm (female); Buccal capsule = 53-60 x 50-53
µm (male) & 60-67 x 53-60 µm (female); Spicules = 56-71 µm; Papillae =

9 pairs (4 preanal, 1 adanal & 4 postanal).
Procamallanus (S.) inopinatus TRAVASSOS, 1929: Brycon brevicaudatus, B. hilarii and
B. erythropterus: Brazil (São Paulo to Amazonas State) and Venezuela. (Fig.
6-24 A-B). Body = 5.0-8.7 x 0.25-0.35 mm (male); 16-28 x 0.45-0.80 mm
(female); Buccal capsule = 80-100 x 60-90 µm (male) & 100-140 x 100-130
µm (female); Spicules = 90-110 & 100-120 µm (note: PETTER & THATCHER
(1988) have pointed out that this species has an oral aperture provided
with two sharp teeth and 4 cutting plates).
Procamallanus (S.) intermedius MAGALHÃES PINTO, FABIO, NORONHA & ROLAS,
1974 (=syn. of P.(S.) pimelodi): Pimelodus clarias: Brazil (São Paulo State).
Body = 8-10 x 0.15-0.23 mm (male); 11-20 x 0.17-0.31 mm (female);
Buccal capsule = 70 x 10 µm (male) & 70-90 x 10-70 µm (female); Spicules
= 240-280 x 60-63 µm.
Procamallanus (S.) krameri (PETTER, 1974): Hoplerythrinus unitaenlatus: French
Guiana and Venezuela. Body = 7 x 0.25 mm (male); 14 x 0.46 mm
(female); Buccal capsule = 50 x 90 µm (male) & 100 x 100 µm (female);
Spicules = 80 & 75 µm; Papillae = 9 pairs (4 preanal & 5 postanal).
Procamallanus (S.) paraensis MAGALHÃES PINTO & NORONHA, 1976: unidentified
fish; “jeju” (possibly Erythrinidae): Brazil (Pará State). Body = 5.2-7.7 x
0.18-0.31 mm (male); 12.6-15.7 x 0.31-0.44 mm (female); Buccal capsule =
70-80 x 90-100 µm (male) & 80-100 x 100-110 µm (female); Spicules = 70
& 90 µm; Papillae = 19 pairs (7 preanal, 1 adanal & 11 postanal).
Procamallanus (S.) pexatus MAGALHÃES PINTO, FABIO, NORONHA & ROLAS, 1974:
Pygidium brasiliensis: Brazil (Espírito Santo State). Body = 3.7-4.3 x 0.21-
0.24 mm (male); 13-20 x 0.45-0.72 mm (female); Buccal capsule = 53-59 x
46-59 µm (male) & 66 x 53-66 µm (female); Spicules = 90 & 110 µm.
Procamallanus (S.) pimelodus MAGALHÃES PINTO, FABIO, NORONHA & ROLAS,
1975: Pimelodus clarias: Brazil (São Paulo State & Matto Grosso State). (Fig.
6-27). Body = 7.6-11.3 x 0.15-0.22 mm (male); 11.7-23.2 x 0.17-0.23 mm
(female); Buccal capsule = 50-70 x 40-50 µm (male) & 49-55 x 40 µm
(female); Spicules = 160-240 & 39-48 µm.
Procamallanus (S.) pintoi (KOHN & FERNANDES, 1988): Corydoras paleatus: Brazil
(Paraná State) and Venezuela. Body = 4.67 x 0.25 mm (male) & 21.8-26 x
0.95-1.16 mm (female); Buccal capsule = 54 x 45 µm (male) & 60-71 x 49-
56 µm (female); Spicules = 94 & 82 µm; Papillae = 6 pairs (4 preanal & 2
postanal).
Procamallanus (S.) rarus TRAVASSOS, 1929: Pimelodella lateristriga and Rhinodoras
d’orbignyi: Brazil (São Paulo State), Paraguay and Argentina. (Fig. 6-26).
Body = 5.3 x 0.11 mm (male); female unknown; Buccal capsule = 70 x 60
µm; Spicules = 400 & 170 µm.
Procamallanus (S.) solani MAGALHÃES PINTO, FABIO, NORONHA & ROLAS, 1975:
unidentified catfish: Brazil (Pará State). Male - right spicule = 430-450 µm;
Papillae = 9 pairs.
Procamallanus (S.) wrighti (PEREIRA, 1935) (considered a synonym of S.

inopinatus by KLOSS 1966): Astyanax sp., Hoplias malabaricus, Leporinus sp. and
Pygocentrus sp.: Brazil. Body = 3.86-4.6 x 0.18-0.27 mm (male) & 8.2-10.6 x
0.30-0.38 mm (female); Buccal capsule = 40-60 µm (male) & 49-110 µm
(female); Spicules = 45 µm; Papillae = 8 pairs (4 preanal & 4 postanal).
Cucullanidae COBBOLD, 1864
Seuratoidea. Mouth an open slit bordered by two large lobes each bearing
2 papillae and an amphid. Esophagus muscular throughout, expanded
posteriorly. Male: spicules equal or unequal; gubernaculum usually present.
Female: vulva near midbody; vagina pointing anteriorly; 2 ovaries usually
present; oviparous. Intestinal tract of fish and reptiles.
Cucullanus MÜLLER, 1777
Cucullanidae. Anterior extremity bent dorsally; lips not sclerotized; intes-
tine without diverticula. Male: rim of preanal sucker not chitinized;
spicules equal; gubernaculum present. Female: 2 ovaries present; eggs with
thin shells. Intestinal tract of fish and reptiles.
C. brevispiculus MORAVEC, KOHN & FERNANDES, 1993: Auchenipterus nuchalis:
Itaipu reservoir, Paraná State, Brazil. Body = 8.6-11 x 0.38-0.48 (female);
9.6-10 x 0.22-0.23 (male): spicules = 0.21.
C. colossomi DÍAZ-UNGRÍA, 1968: Colossoma macropomum: Venezuela (Amazonas State).
Body = 17 x 0.6 mm (male) & 22 x 0.7 mm (female); Spicules = 1.3 mm long.
C. grandistomis (FERRAZ & THATCHER, 1988): Pseudodoras niger: Brazil (Amazo-
nas). (Fig. 6-38). Body = 3.7-4.7 x 0.09-0.16 mm (male) & 3.0-7.4 x 0.06-
0.15 mm (female); Spicules = 0.78 mm.
C. interrogativus TRAVASSOS, ARTIGAS & PEREIRA, 1928 (= syn. of Neocucullanus
neocucullanus): Salminus maxillosus: Brazil (São Paulo State). Species based on
immature specimens; Body = 10.5 x 0.45 mm (male & 12.9 x 0.50 mm
(female); Mouth lobes with small denticles, each about 5 µm long.
C. mogi TRAVASSOS, 1948: Leporinus sp.: Brazil (São Paulo State). Description unavailable.
C. oswaldocruzi VICENTE, SANTOS & JARDIM, 1979 Pseudopimelodus zungaro: Brazil
(Pará State). Body = 17 x 0.51 mm (male) & 18.2 x 0.54 mm (female);
Spicules = 1.1-1.2 mm long.
C. pauliceae VAZ & PEREIRA, 1934: Paulicea luetkeni: Brazil (São Paulo State).
Body = 18.5 x 0.76 mm (male) & 20.3 x 0.76 mm (female); Buccal capsule
= 0.40 x 0.36 mm; Spicules = 1.2 mm.
C. pimelodellae MORAVEC, KOHN & FERNANDES, 1993: Pimelodella lateristriga:
Paraná State, Brazil. Body = 5-12 x 0.23-0.39 (female); 5-12 x 0.18-0.44
(male); Spicules = 0.35-0.68.
C. pinnai TRAVASSOS, ARTIGAS & PEREIRA, 1928: Pimelodus clarias and Pseudo-
platystoma sp.: Brazil. Body = 8.5 x 0.3 mm (male) & 5.9-9.8 x 0.28-0.30 mm
(female); Buccal capsule = 140 x 140 µm; Spicules = 57 µm.
C. pseudoplatystomae MORAVEC, KOHN & FERNANDES, 1993: Pseudoplatystoma
corruscans: Paraná River, Paraná State, Brazil. Body = 8.6-15 x 0.20-0.27
(female); 9-12 x 0.18-0.27 (male); Spicules = 0.49-0.53.
C. rhamphichthydis MORAVEC, KOHN & FERNANDES, 1997: Rhamphichthys rostratus:

Paraná River, Paraná State, Brazil. Body = 9.3-12.4 x 0.20-0.22 (female);
male unknown
C. schubarti TRAVASSOS, 1948: Paulicea luetkeni: Brazil (São Paulo State). (Fig. 6-
16 A-B). Body = 14.5 x 15.6 x 0.48-0.59 mm (male) & 19.2 x 0.58-0.61 mm
(female); Spicules = 1.0-1.4 mm long.
C. zungaro VAZ & PEREIRA, 1934: Pseudopimelodus zungaro and Paulicea luetkeni:
Brazil (São Paulo State). Body = 5.4-6.1 x 0.36 mm (male) & 8.3 x 0.41 mm
(female); Buccal capsule = 0.14 x 0.14 mm; Spicules = 0.64-0.70 mm.
Neocucullanus TRAVASSOS, ARTIGAS & PEREIRA, 1928
Cucullanidae. Similar to Cucullanus but mouth cavity is less well developed
and tail of male is long and tapered.
N. neocucullanus TRAVASSOS, ARTIGAS & PEREIRA, 1928: Characidae: Brazil
(Minas Gerais State). (Fig. 6-15). Body = 16 x 0.68 mm (male) & 32 x 0.9
mm (female); Spicules = 380 µm; Egg = 60-70 µm long.
Gnathostomatidae LANE, 1923
Gnathostomatoidea. Mouth with prominent trilobed lateral lips having
inner tooth-like ridges; spherical head bulb behind lips provided with
transverse striations or retrospines. Four cervical glands present. Males:
caudal alae supported by pedunculate papillae; spicules equal or unequal.
Females: vulva postequatorial; vagina directed anteriorly; uterus with 2 or 4
branches; oviparous; egg shells thin, ornamented. Digestive tract of fish,
reptiles and mammals.
Echinocephalus MOLIN, 1858
Gnathostomatidae. Head bulb armed with retrospines; body unarmed. Male:
caudal alae small; 8 pairs of papillae present, with anterior pair distant from
others; spicules slightly unequal, long, slender, marked by transverse stria-
tions. Female: tail long, conical; vulva near posterior end; vagina long;
oviparous. From intestinal spiral valve of elasmobranch fishes.
E. daileyi DEARDORFF, BROOKS & THORSON, 1981: Potamotrygon circularis and P.
histrix: Colombia, Venezuela & Brazil. (Fig. 6-11 A-D). (Note: This species is
known from the Amazon and Orinoco River systems). Body = 55-66 x 0.6 x
0.9 mm (male) & 55-85 x 0.9-1.4 mm (female); Head bulb = 0.44-0.53 x
0.61-0.65 mm (male) & 0.48-0.68 x 0.59-0.80 mm (female); Retrospines in
30-34 rows; Gubernaculum present; Spicules equal = 1.4-2.4 mm long.
Guyanemidae PETTER, 1974
Dracunculoidea. Anterior extremity with 6 pairs of cephalic papillae. Buccal
capsule absent. Esophagus divided into 2 parts. Male: spicules equal; caudal
alae present; gubernaculum absent. Female: vulva pre-equatorial; viviparous.
Parasitic in abdominal cavity and digestive tract of fishes.
Guyanema PETTER, 1974
Guyanemidae, with the characters of the family. Male: 4 pairs of preanal
papillae present. Female: monodelphic. From abdominal cavity and intesti-
nal tract of fish.
G. seriei PETTER, 1974: Hoplerythrinus unitaeniatus: French Guiana. (Fig. 6-18 A-B).
Body = 8 x 0.09 mm (male) & 14 x 0.19 mm (female); Spicules = 100 µm.
G. seriei paraguayensis PETTER & DLOUHY, 1985: Hoplerythrinus unitaeniatus:
Paraguay. Body = 4.4 x 0.10 mm (male) & 6.35 x 0.08 mm (female);
Spicules = 80 µm.
G. ancistri PETTER, 1987: Ancistrus sp.: Ecuador. Body = 6.8-9.8 mm (female).
G. baudi PETTER & DLOUHY, 1985: Hoplias malabaricus: Paraguay. Body = 2.55-
2.70 x 0.05-0.06 mm (male) & 5 x 0.05 mm (female); Spicules = 70 µm.
G. longispiculum MORAVEC, PROUZA & ROYERO, 1996: Loricariichthys brunneus:
Orinoco River, Venezuela. Body = 6.8-9.3 x 0.07-0.08 (male); 16.8-30.7
(female); Spicules = 0.48-0.61.
G. raphiodoni MORAVEC, KOHN & FERNANDES, 1993: Raphiodon vulpinus: Itaipu
reservoir, Paraná State, Brazil. Body = 3.7 x 0.07 (male); Spicules 0.15.
Travassosnema COSTA, MOREIRA & OLIVEIRA, 1991.
Guyanemidae, with the characters of the family. Cephalic end rounded;
buccal capsule absent. Esophagus divided into anterior muscular part and
posterior glandular part; latter with elongate dorsal appendix extending
posteriorly. Deirids present. Caudal end of male with alae, preanal and
postanal papillae; two spicules and gubernaculum present. Vulva anterior.
Monodelphic. Ovoviviparous. Anus functional in female. Parasites of
body cavity and tissues of fishes. Type species: Travassosnema travassosi
COSTA, MOREIRA & OLIVEIRA, 1991.
T. travassosi COSTA, MOREIRA & OLIVEIRA, 1991: Acestrorhynchus lacustris: Minas
Gerais State, Brazil. (Fig. 6-94). Body = 2.0-2.4 x 0.03-0.05 (male); 7.5-13.8
x 0.14-0.24 (female); Spicules = 0.05-0.06. MORAVEC et al (1993) divided
this species into two subspecies, namely:
T. travassosi travassosi and T. travassosi paranaensis. The latter was found in the
same host species from Paraná State, Brazil. The differences between the
two are largely based on size.
Anisakidae RAILLIET & HENRY, 1912
Ascaroidea. Esophagus with oblong to cylindrical posterior ventriculus
with one dorsal and one ventral longitudinal suture-like depression, or
with small globular ventriculus giving off posteriorly directed appendix
with such depressions or rarely two appendices. Excretory system asym-
metrical, confined to left lateral chord. Excretory pore situated between
base of subventral lips or near nerve ring. Intestinal caecum present or
absent. Parasites of all classes of vertebrates.
Goezia ZEDER, 1800
Anisakidae. Body with cuticular rings which are provided with retrospines.
Lips flattened and expanded outward, separated from body by constric-
tion; esophagus expanded posteriorly and with 1 or 2 long ventricular
appendages; short anterior cecum present. Male: spicules nearly equal;
gubernaculum absent. Female: vulva pre-equatorial; oviparous. From
digestive tract of fish and reptiles.
G. spinulosa (DIESING, 1839): Astronotus ocellatus and Arapaima gigas: Brazil

(Amazonia). (Fig. 6-21 A-B). Body = 6.3-15.3 x 0.37-0.77 mm (male) & 11.7-
16.2 x 0.80-0.87 mm (female); Spicules = 410-320 µm; Egg = 50 x 38 µm.
Raphidascaris.(Sprentascaris) RAILLIET & HENRY, 1915
Anisakidae. Lips without denticles; interlabia absent; post-labial cuticular
horns present; cervical alae present or absent; esophagus with small
ventriculus and short appendix; intestinal cecum absent. Male: spicules
equal; gubernaculum absent. Female: vulva pre-equatorial; eggs embryo-
nated or not. From digestive tract of siluriform fishes.
R. (Sprentascaris) hypostomi (PETTER & CASSONE, 1984); Hypostomus sp.: Paraguay.
Body = 6.8-8.2 mm (male) & 6.4 x 11.0 mm (female); Esophagus = 0.90-
1.4 mm; Spicules = 150-250 µm.
R. (Sprentascaris) mahnerti (PETTER & CASSONE, 1984); Loricariichthys platymeto-
pon: Paraguay. (Figs. 6-20 A-C & 6-98). Body = 5.5-7.5 mm (male) & 7.5-
11.0 mm (female); Esophagus = 0.70-1.1 mm; Spicules = 120-145 µm.
R. (Sprentascaris) pimelodi (PETTER & CASSONE, 1984); Pimelodus maculatus:
Paraguay. Body = 8.9 mm (male) & 13.6 x 19.0 mm (female); Esophagus =
0.90-1.6 mm; Spicules = 140 µm.
Raphidascaroides YAMAGUTI, 1941
Anisakidae. Rather large nematodes. Lips well developed, with lateral cuticular
flanges, with or without dentigerous ridges. Interlabia well developed. Excretory
pore slightly posterior to nerve ring level. Esophagus muscular, provided with
almost spherical ventriculus; posteriorly directed ventricular appendix present.
Intestinal cecum absent. Spicules equal, alate. Genital papillae numerous. Vulva
pre-equatorial. Oviparous. Parasite of freshwater and marine fishes.
R. brasiliensis MORAVEC & THATCHER, 1997. Pterodoras granulosus. Janauacá Lake,
Amazon River, near Manaus, Brazil. (Figs. 6-60 & 6-61). Body = 31.2-34.4
x 0.6-0.7 mm (male) & 46-76.7 x 0.82-1.03 mm (female); Egg = 45-51 x
36-42 µm long.
Terranova LEIPER & ATKINSON, 1914
Anisakidae. Cephalic end with three short lips bearing fine dentigerous ridges on
inner surface. Interlabia absent. Ventriculus well developed, ventricular appendix
absent. Anterior intestinal cecum present. Excretory pore situated at base of
ventrolateral lips. Spicules equal, gubernaculum absent. Vulva in anterior part of
body. Tail conical. Parasites of digestive tract of fishes and reptiles.
T. diazungriai MORAVEC, 1998: Potamotrygon hystrix, Potamotrygon circularis. Vene-
zuela (Orinoco River Delta). (Fig. 6-12 A-B). Body = 17.0 x 0.5 mm (male)
& 28 x 0.70 mm (female).
T. serrata (DRASCHE, 1884): Arapaima gigas: Brazil (Amazonia). (Fig. 6-12 A-B).
Body = 25 x 0.62 mm (male) & 28 x 0.73 mm (female); Spicules = 1.5 mm;
Egg = 45-50 µm long.
Kathlaniidae YORK & MAPLESTONE, 1926
Cosmocercoidea. Mouth with 3 or 6 well developed lips; teeth present or
absent; interlabia present or absent; esophagus with posterior bulb, usually
preceded by a swelling; intestine without diverticula. Male: pre-cloacal

sucker usually present, may have chitinoid rim; spicules equal or subequal;
gubernaculum usually present. Female: vulva postequatorial. From diges-
tive tracts of cold-blooded vertebrates.
Chabaudinema DÍAZ-UNGRÍA, 1968
Kathlaniidae. Mouth with 6 papillate lips; esophagus with anterior cuticu-
larized ring. From digestive tract of fish.
C. americanum DÍAZ-UNGRÍA, 1968: Colossoma macropomum: Venezuela (Amazo-
nas State). (Fig. 6-19 A-C). Body = 30 x 1.28 mm (male) & 34 x 1.5 mm
(female); Spicules = 0.90-1.1 mm; Eggs = 115-117 µm.
Myleusnema MORAVEC & THATCHER, 1996.
Kathlaniidae. Body large, with cephalic portion separated from remaining
wider portion of body by transverse cuticular fold approximately at level
of nerve ring. Mouth aperture triangular, surrounded by three small lips,
each associated with a cuticularized lamella-like formation demarcating a
poorly developed buccal cavity. Anterior extremity of esophagus not
differentiated into a pharyngeal portion; posterior extremity of esophagus
formed by elongate, expanded isthmus and spherical bulb with valves. Tail
conical, relatively short. Male: ventral precloacal sucker present; posterior
part of cloacal opening with lobe-like formation armed with two conspic-
uous horns associated with a gubernaculum; spicules simple, alate and of
equal length (similar); numerous paired preanal and postanal papillae
present; caudal alae absent. Female: vulva postequatorial; reproductive
apparatus prodelphic; eggs numerous, oval, thin walled, containing moder-
ately developed embryos. Intestinal parasites of Neotropical fishes. Type
species: M. bicornis MORAVEC & THATCHER, 1996
M.bicornis MORAVEC & THATCHER, 1996: Myleus ternetzi: Takari Tanté Falls,
Sinnamary River, French Guiana. (Figs. 6-64 to 6-73). Body = 14.28-17.41
mm x 1.09-1.12 µm & 16.62-20.13 mm x 1.36-1.54 mm (female); Eggs =
135 x 150 x 96-105 µm.
M.brasiliense MORAVEC & THATCHER, 1999. Myleus sp.: Serra da Mesa, Tocantins
River, Pará State, Brazil. (Figs. 6-74 to 6-77). Body = 15.16-22.63 mm x
1.02-1.31 µm & 18.33-24.18 mm x 1.14-1.56 mm (female); Eggs = 122 x
150 x 82-95 µm.
Spectatus TRAVASSOS, 1923
Kathlaniidae. Mouth with 6 lips each of which bears a papilla; esophagus
with posterior bulb containing valves; excretory pore anterior to esoph-
ageal bulb. Male: spicules subequal, similar, falcate; preanal sucker ellipsoi-
dal, with chitinoid rim; Papillae = 11 pairs (5 preanal, 3 adanal & 3
postanal). Female: vulva in midbody; ovijector long; eggs large, thick-
shelled. From intestine of freshwater fish.
S. spectatus TRAVASSOS, 1923: Colossoma brachypomum: Brazil. (Fig. 6-8 A-B).
Body = 10-12 x 0.7 mm (male) & 12-14 x 0.8 mm (female); Spicules = 0.7-
1.0 mm; Egg = 92 x 63 µm.
Pharyngodonidae TRAVASSOS, 1919

Oxyuroidea. Oral aperture surrounded by four large cephalic papillae and
two lateral amphids. Lateral alae present or absent. Genital cone in male
often supported by a V-shaped sclerotized structure. Male caudal alae
present or absent. Eggs usually elongate-oval, asymmetrical, often provid-
ed with operculum or filaments. Parasites of cold-blooded vertebrates,
rarely of archaic mammals.
Brasilnema MORAVEC, KOHN & FERNANDES, 1992.
Pharyngodonidae. Body small, with distinct transverse striations; lateral
alae present in both sexes. Mouth opening hexagonal, without lips,
surrounded by six triangular lamellae. Buccal capsule large. Tail with very
long terminal spike in both sexes. Male: gubernaculum absent; spicule
simple, moderately sclerotized; genital papillae clearly separated into
anterior group on protruding genital cone and one posterior pair; caudal
alae absent. Female: vulva pre-equatorial; uterus opposed; eggs larvated,
provided with long polar filaments. Intestinal parasites of fishes. Type
species: Brasilnema pimelodellae MORAVEC, KOHN & FERNANDES, 1992.
B. pimelodellae MORAVEC, KOHN & FERNANDES, 1992: Pimelodella lateristriga.
Paraná River near Guaira, Paraná State, Brazil. (Fig. 6-95 A-B). Body =
0.99-1.20 mm x 82-136 µm (male including tail spike) & 1.96-2.64 mm x
249-326 µm (female); Eggs with numerous filaments in both poles = 144-
156 x 63-96 µm in size.
Cosmoxynema TRAVASSOS, 1948
Pharyngodonidae. Body fusiform, transversely striated, with lateral flanges
throughout body length. Mouth circular; buccal capsule spherical, with
three conical teeth at base; esophagus claviform, followed by bulb
containing valves. Male: unknown. Female: vulva pre-equatorial; ovaries in
midbody; unpaired portion of uterus divided into 2 branches; eggs
elongate, curved, not embryonated. Intestinal parasites of fish.
C. viannai TRAVASSOS, 1948: Curimata gilberti: Brazil. (Fig. 6-6). Body = 3.1-4.6 x
0.33 mm (female; male unknown); Buccal capsule = 30-40 µm; Egg = 163-
174 x 38-40 µm.
Cosmoxynemoides TRAVASSOS, 1948
Pharyngodonidae. Body fusiform, transversely striated; lateral flanges
present throughout body length. Cephalic end smooth, separated from
body by circular groove; buccal capsule small; esophagus claviform,
followed by bulb with valves. Male unknown. Female: vulva pre-equatorial;
uterus as in Cosmoxynema; egg elongate, bent. Intestinal parasite of fish.
C. aguirrei TRAVASSOS, 1948: Curimata gilberti and Geophagus brasiliensis: Brazil
(São Paulo State). (Fig. 6-4). Body = 3.3-3.8 x 0.29-0.30 mm (female; male
unknown); Egg = 190-197 x 38-45 µm.
Ichthyouris INGLIS, 1962.
Pharyngodonidae. Body small, cuticle transversely striated. Lateral alae
present. Oral aperture bounded by six small labial lobes, each associated
with small, anteriorly protruding lobe-like lamella growing out from its
base. Tail with long terminal spike in both sexes. Male: gubernaculum
absent; spicule simple, well sclerotized; genital papillae around anus, two
pairs of them being long, pedunculate; broad caudal alae present.
Female: vulva pre-equatorial, uterus prodelphic; eggs larvated mostly
provided with filaments. Intestinal parasites of fishes. Type species:
Ichthyouris ro INGLIS, 1962.
I. brasiliensis MORAVEC, KOHN & FERNANDES, 1992: Pterygoplichthys aculeatus and
Megalancistrus aculeatus: Brazil (Paraná River near Guaira, Paraná State) and
Paraguay (Province Alto Paraná). Body = 1.28-1.88 mm x 109-150 µm
(male including tail spike) & 3.16-4.50 mm x 231-422 µm (female); Eggs
with numerous filaments on both poles = 93-99 x 33-36 µm in size.
I. bursata MORAVEC & PROUZA, 1995: Symphysodon discus: Intestine. Introduced
in Czech Republic together with fishes from South America. Body = 1.24-
1.39 mm x 109 µm (male including tail spike) & 2.39-3.17 mm x 231-340
µm (female); Eggs with only one filament on each pole = 105-111 x 39-42
µm in size.
I. laterifilamenta MORAVEC, KOHN & FERNANDES, 1992: Trachydoras paraguayensis:
Intestine. Paraná River basin, Brazil and Paraguay. Body = 1.16-1.44 mm x 82-
109 µm (male including tail spike) & 2.56-3.33 mm x 190-299 µm (female);
Eggs with numerous lateral filaments = 117-138 x 51-69 µm in size.
I. ovifilamentosa MORAVEC & THATCHER, 2001: Cichlasoma: Intestine. Rio Negro,
São Gabriel da Cachoeira, Amazonas, Brazil. (Figs. 6-53 to 6-55). Body =
1.7-1.8 mm x 163 µm (male including tail spike) & 3.9-4.583 mm x 231-286
µm (female); Eggs = 99-102 x 30-33 µm.
I. ro INGLIS, 1962: Cichlasoma festivum and Geophagus steindachneri: Intestine.
Guyana & Peru. Body = 0.83-1.10 mm x 57-87 µm (male including tail
spike) & 2.10-2.40 mm x 150-190 µm (female); Eggs = 36-11 µm.
Parasynodontisia MORAVEC, KOHN & FERNANDES, 1992:
Pharyngodonidae. Small nematodes; females distinctly larger than males.
Cuticle with fine transverse striations. Lateral alae absent. Mouth opening
triangular, surrounded by six low triangular lamellae in hexagonal pattern.
Four large circumoral cephalic papillae and one pair of elongate amphids
present. Buccal capsule absent. Anterior end of esophagus forming well
developed “pharynx” containing three depressions in its anterior region,
which correspond to its three longitudinal sectors. Esophagus of almost
uniform diameter except for posterior bulb containing sclerotized appara-
tus. Nerve ring encircling esophagus approximately at border of its first
and second quarters. Excretory pore situated anterior to level of bulb. Tail
conical (in larger specimens more elongate), ending in sharp cuticular
spine. Type specie: P. petterae MORAVEC, KOHN & FERNANDES, 1992.
P. petterae MORAVEC, KOHN & FERNANDES, 1992: Rhinelepis aspera: Intestine. Paraná
River in Brazil and Paraguay. Body = 1.39-2.46 mm x 82-122 µm (male including
tail spike) & 2.35-4.09 mm x 177-340 µm (female); Eggs = 69-33 µm.
Spinoxyuris PETTER, 1994

Pharyngodonidae. Small nematodes with transversely striated cuticle. Lateral
alae in both sexes, each ending posteriorly as a spike. Buccal cavity of female
surrounded by six small oral membranous lamellae, four oval cephalic
papillae and two lateral amphids. Esophagus consisting of cylindrical corpus,
short isthmus and bulb with sclerotized apparatus. Excretory pore posterior
to esophageal bulb. Type specie: S. oxydoras PETTER, 1994.
S. annulata MORAVEC & THATCHER, 2001: Myleus ternetzi: Sinnamary River,
French Guiana. (Figs. 6-50 to 6-52 & 6-56 to 6-59). Body = 1.8-1.9 mm x
109-122 µm (male including tail spike) & 4.0-4.4 mm x 340 – 408 µm
(female); Eggs = 150-180 x 66-84 µm.
S. oxydoras PETTER, 1994: Oxydoras kneri: Paraná River, Province Alto Paraná,
Paraguay. Body = 1.00-1.05 mm x 55-70 µm (male including tail spike) &
1.93-2.42 mm x 150-200 µm (female); Eggs = 110 x 50 µm. With few
filaments growing out irregularly from egg surface.
Travnema PEREIRA, 1938
Pharyngodonidae. Mouth circular; lips lacking; 4 papillae present; buccal
capsule small; pharynx and esophagus both flask-shaped. Male: lateral alae
extending throughout most of body length; 1 spicule present; gubernaculum
absent. Female: cervical alae well developed; lateral alae reduced; vulva post-
equatorial; 2 ovaries present; originating at posterior extremity; ovijector
short; egg large, asymmetrical, operculate, embryonated. Intestine of fish.
T. travnema PEREIRA, 1938: Curimata elegans: Brazil (São Paulo State). (Fig. 6-9).
Body = 1.1 x 0.10 mm (male) & 2.1-3.1 x 0.24-0.30 mm (female); Spicules
= 45 µm; Egg = 164-169 x 74-82 µm.
T. araujoi FERNANDES, CAMPOS & ARTIGAS, 1983: Curimata gilberti: Brazil (São
Paulo State). Body = 2.1 x 0.16 mm (male) & 3-5 x 0.23-0.39 mm (female);
Spicules = 36 µm; Egg = 177-229 x 52-81 µm.
Philometridae BAYLIS & DAUBNEY, 1926
Dracunculoidea. Body long, slender. Anterior end rounded, mouth simple,
without lips, but 6 or 8 papillae may be present; esophagus elongate,
divided into 2 parts. Male: 2 spicules present; spicules equal, slender, finely
pointed; gubernaculum may or may not be present. Female: vulva small, or
absent; vagina rudimentary, or absent, in gravid females; ovaries short, at
opposite ends of body; uterine branches opposed, forming continuous
tube, viviparous. Parasites of body cavity or tissues of fishes.
Nilonema KHALIL, 1960.
Female body large, with narrowed ends. Cuticle with large, cone-shaped
excrescences. Esophagus cylindrical, without anterior inflation. Cephalic
papillae small. Ovaries opposed. Anus and vulva atrophied. Males un-
known. Parasites of freshwater fishes.
N. senticosa BAYLIS, 1927: Arapaima gigas: Brazil (Amazonia) and Peru. (Fig. 6-
14 A-B). Body = 90-125 x 0.62-0.85 mm (female); Cuticle with spine-like
protuberances, irregularly placed; Embryos = 300 µm.
Philometra COSTA, 1845

Philometridae. Body filiform; female much larger than male; both extrem-
ities rounded; head and tail papillae may be present. Mouth with or
without lips; esophagus short, expanded anteriorly; rudimentary ventricu-
lus present. Male: cloaca terminal, bordered by 2 lips; spicules equal,
needle-like; gubernaculum present. Female: anus and vulva atrophied;
vulva in posterior one-third of body in young females; ovaries small;
uterus voluminous; viviparous. Tissue and body cavity parasites of fish.
P. amazonica TRAVASSOS, 1960: Callophysus macropterus: Brazil. Body = 265-950 x
0.5-1.0 mm (female); Embryos = 416-445 x 14 µm.
P. baylisi VAZ & PEREIRA, 1934: Pimelodus clarias: Brazil. Body = 95 x 0.29 mm
(female); Mouth with 3 lips; Caudal spine = 10 µm.
P. maplestonei TRAVASSOS, ARTIGAS & PEREIRA, 1929: Salminus hilarii: Brazil.
Body = 144 x 0.22 mm (female); Cuticle with irregularly placed, small,
rounded protuberances; Embryos = 320 x 18 µm.
Quimperiidae BAYLIS, 1930
Seuratoidea. Lips small or absent; buccal capsule absent; esophagus divided
or not. Male: preanal sucker usually present, lacks chitinized rim; spicules
equal; gubernaculum present or not. Female: vulva post-equatorial; ovipa-
rous. Parasites of digestive tract of fish and occasionally amphibians.
Neoparaseuratum MORAVEC, KOHN & FERNANDES, 1992
Quimperiidae. Head rounded; mouth hexagonal, surrounded by six small
lip-like formations; four submedian cephalic papillae and two lateral
amphids present. Esophagus long, undivided, its posterior of greater
diameter than anterior. Cephalic region of body surrounded by numerous
longitudinal bands of inflated cuticle. Deirids small, at esophagus level;
excretory pore postesophageal. Male: caudal alae absent, oblique muscle
bands in preanal region present. Spicules equal, short gubernaculum
present. Female: vulva postequatorial, uterus opposed. Eggs not embryo-
nated in uteri. Intestinal parasites of South American fishes. Type species:
Neoparaseuratum travassosi MORAVEC, KOHN & FERNANDES, 1992.
N. travassosi MORAVEC, KOHN & FERNANDES, 1992: Pterodoras granulosus and
Trachydoras paraguayensis: Basin of Paraná River in Brazil. (Fig. 6-97). Body
= 2.26-8.02 mm x 122-299 µm (male) & 8.41-11.40 mm x 299-381 µm;
Egg = 90-105 x 54-66 µm.
Paraseuratum JOHNSTON & MAWSON, 1940.
Quimperiidae. Cephalic end truncated. Mouth surrounded by six low lips;
four double cephalic papillae and lateral amphids present. Buccal capsule
absent. Esophagus long, with inflated anterior end; small esophastome
present, containing 3 small sclerotized pieces and 3 small teeth in its
lumen. Male without caudal alae; tail conical. Spicules equal. Vulva at
second third of body, vagina short. Uterus opposed. Eggs almost spherical
Tail of female conical. Intestinal parasites of freshwater fishes. Type
species: Paraseuratum tandani JOHNSTON & MAWSON, 1940.
P. albidum, KLOSS, 1966: Astyanax bimaculatus, Tetragonopterus argenteus and Erythri-

nus erythrinus: Paraná River basin, São Paulo, Brazil & Ecuador (Amazon
River drainage system). (Fig. 6-17 A-B). Body = 7.7-11 mm x 200-500 µm
(male) & 9.2-21.5 mm x 206-750 µm; Egg = 70-80 x 45-50 µm.
P. soaresi FABIO, 1982: Hoplias malabaricus. Rio de Janeiro State, Brazil.
Touzeta PETTER, 1987
Quimperiidae. Oesophastome present; esophagus short, divided into 2
parts of different diameters. Male: preanal pseudo-sucker present; spicules
equal; gubernaculum present. Female: vulva post-equatorial; amphidel-
phes. From digestive tract of fish.
T. ecuadoris PETTER, 1987: Cichlidae: Ecuador. (Fig. 6-22 A-B). Body = 4.8 x 0.35
mm (male) & 6 x 0.40 mm (female); Spicules = 140 µm; Embryos = 70 x 50 µm.
Rhabdochonidae SKRJABIN, 1946
Thelazioidea. Mouth with or without lips; buccal cavity funnel-shaped or
cylindrical, with or without teeth; esophagus of 2 parts. Male: posterior
extremity usually rolled or coiled; caudal alae narrow, papillae sessile;
spicules unequal. Female: oviparous. Parasites of digestive tract of fish and
amphibians.
Rhabdochona RAILLIET, 1916
Rhabdochonidae. Body smooth; mouth with 2 reduced lips; buccal capsule
funnel-shaped, with longitudinal ridges terminating in teeth; esophagus of 2
parts. Male: numerous simple preanal papillae and 3-6 pairs of postanal
papillae present; spicules unequal. Female: vulva in mid-body; uterine branches
opposed; oviparous; eggs elliptical. From intestine of freshwater fishes.
R. acuminata (MOLIN, 1860); Brycon falcatus, Pimelodella lateristriga: Brazil: Body =
9-12 x 0.16-0.21 mm (male) & 14-21 x 0.26-0.35 mm (female); Spicules =
530 & 170 µm; Egg = 35 x 22 µm.
R. australis KLOSS, 1966 (= syn. of R. acuminata): Astyanax bimaculatus: Brazil (São
Paulo State). Body = 8.0-8.4 x 0.08-0.12 mm (male) & 12.6-14.9 x 0.12-0.16
mm (female); Spicules = 350-490 & 110-120 µm; Egg = 21-24 x 13-18 µm.
R. colossomi DÍAZ-UNGRÍA, 1968: Colossoma macropomum: Venezuela. Descrip-
tion unavailable.
R. elegans ARTIGAS & PEREIRA, 1929 (= syn. of R. acuminata): Tetragonopterus sp.:
Brazil (São Paulo State). (Fig. 6-2 A-B). Body = 12-13 x 0.2 (male) & 22-28
x 0.3 mm (female); Spicules = 420-450 & 140-150 µm; Egg = 42 x 18 µm.
R. fasciata KLOSS, 1966 (= syn. of R. acuminata): Astyanax fasciatus: Brazil (São
Paulo State). Body = 4.5-6.0 x 0.05-0.08 mm (male) & 18.7-22.2 x 0.24-
0.28 mm (female); Spicules = 380 & 680-710 µm; Egg = 24 x 18 µm.
R. uruyeni DÍAZ-UNGRÍA, 1968: Piabucina sp.: Venezuela. Body = 7.5 x 0.12 mm
(male) & 11.8-14.9 x 0.15 mm (female); Spicules = 320-400 & 65-80 µm;
Egg = 35 x 20 µm.
Cystidicolidae SKRJABIN, 1946
Habronematoidea. Mouth simple, with small lips; buccal capsule sclero-
tized, cylindrical; esophagus long, clearly divided into anterior and posteri-
or parts. Male: Posterior extremity spiraled; caudal alae narrow; papillae

numerous; spicules unequal. Female: uterus divergent; egg thin-shelled.
From digestive tract of fish.
Cystidicoloides SKINKER, 1931
Cystidicolidae. Large or medium sized nematodes. Cuticle sometimes with
ornamentations. Oral opening slit-like or dorsoventrally elongated.
Pseudolabia small, often reduced to small appendix. Cephalic papillae
usually reduced to four at base of pseudolabia. Anterior end of stoma
(vestibule) sometimes armed with small teeth. Esophagus divided into
anterior muscular and posterior glandular sections. Caudal alae in male
present. Eggs oval, larvated, sometimes with special formations on
surface. Parasites of digestive tract and swim bladder of fishes.
C. dlouhyi PETTER, 1984: Sternopygus macrurus: Paraguay. Body = 5.5 x 0.20 mm
(male) & 6.2-9.1 x 0.25-0.38 mm (female); Spicules = 590 & 150 µm.
C. fischeri (TRAVASSOS, ARTIGAS & PEREIRA, 1928): Salminus maxillosus: Brazil (São
Paulo State). (Fig. 6-3 A-B). Body = 9.6 x 0.20 mm (male) & 9.7-12.0 x 0.20-
0.30 mm (female); Spicules = 900-1000 & 160-180 µm; Egg = 40-44 x 32 µm.
C. vaucheri PETTER, 1984: Oxydoras kneri: Paraguay. Body = 16.2 x 0.4 mm
(female); Egg = 40 x 25 µm.
Spinitectus FOURMENT, 1883
Cystidicolidae. Cuticle with transverse rings bearing retrospines; mouth with
indistinct lips; buccal cavity funnel-shaped or cylindrical; esophagus of 2
parts. Male: posterior extremity coiled; caudal alae narrow; preanal and
postanal papillae present, numerous; spicules unequal, dissimilar. Female:
vulva equatorial or post-equatorial; oviparous; eggs small, ellipsoidal, polar
filaments sometimes present. From digestive tracts of fish and frogs.
S. asperus TRAVASSOS, ARTIGAS & PEREIRA, 1928: Prochilodus scrofa: Brazil (São
Paulo State). Body = 8.2 x 0.28 mm (female); Egg = 18-21 x 16 µm.
S. jamundensis THATCHER & PADILLA, 1977: Prochilodus reticulatus: Colombia
(Department of Valle). (Fig. 6-10 A-B). Body = 2.5-4.8 x 0.10-0.18 mm
(male) & 6.0-9.8 x 0.17-0.30 mm (female); Spicules = 200-280 & 90-130 µm;
Egg = 32-40 x 22-30 µm. This species was considered a synonym of S.
asperus by MORAVEC (1998).
S. multipapillata PETTER, 1987: Pimelodella sp.: Ecuador. Body = 6.3-7.4 x 0.11-
0.12 mm (male) & 6.5 x 0.15 mm (female); Spicules = 220-240 µm.
S. pachyuri PETTER, 1984: Pachyurus bonariensis: Paraguay. Body = 10.6-0.065
mm (female); Egg = 40 x 28 µm.
S. rudolphiheringi VAZ & PEREIRA, 1934: Pimelodella lateristriga and Salminus
hilarii: Brazil (São Paulo State). Body = 3.7-4.0 x 0.12 mm (male) & 3.0 x
0.12 mm (female); Spicules = 200 & 120 µm.
S. sternopygi PETTER, 1984: Sternopygus macrurus: Paraguay. Body = 4.7-4.9 x 0.13
mm (male) & 5.9-7.0 x 0.15-0.25 mm (female); Spicules = 200-210 & 120-
130 µm; Egg = 40 x 25 µm. This species was considered a synonym of S.
rudolphiheringi by MORAVEC (1998).
S. yorkei TRAVASSOS, ARTIGAS & PEREIRA, 1928: Pimelodella lateristriga: Brazil (São
Paulo State). Body = 4.2 x 0.08 mm (male) & 11 x 0.16 mm (female);
Spicules = 162 & 40 µm; Egg = 30-34 x 20-23 µm.
Capillariidae RAILLIET, 1915.
Trichinelloidea. Thread-like nematodes. Cuticle with longitudinal bacillary
bands, usually two lateral, one dorsal and one ventral. In males, one more
or less sclerotized spicule present, with smooth or spiny spicular sheath,
able to invaginate or evaginate. Posterior end of male with or without
small membranous bursa. Vulva near distal end of esophagus, sometimes
with elevated lips. Eggs usually barrel-shaped, with polar plugs, and with
smooth or variously sculptured surface. Parasitic in varios organs.
Capillaria ZEDER, 1800
Capillariidae. Body capillary; mouth simple; esophagus long, increases in
diameter posteriorly. Male: anus terminal or subterminal; caudal alae
sometimes present; spicule long and slender, with spinous or smooth sheath.
Female: vulva near posterior end of esophagus; oviparous; eggs elliptical,
with polar plugs. Digestive tract or urinary bladder of vertebrates.
C. minima TRAVASSOS, ARTIGAS & PEREIRA, 1928 (= syn. of Capillostrongyloides
sentinosa): Leporinus sp.: Brazil (São Paulo State). Body = 1 x 0.03 mm (male)
& 1.9 x 0.035 mm (female); Spicule = 130 µm; Egg = 50 x 25 µm.
C. zederi (TEXEIRA DE FREITAS & LENT, 1935) (= syn. of Capillostrongyloides
sentinosa): Hoplias malabaricus: Brazil (São Paulo State). Body = 3.7-3.8 x
0.016-0.048 mm (male) & 3.4-3.5 x 0.024-0.040 mm (female); Spicule =
140-160 µm; Egg = 48-68 x 24-35 µm.
Capillostrongyloides FREITAS & LENT, 1935.
Capillariidae. Stichosome consisting of single row of stichocytes; lateral
caudal alae in male absent; membranous bursa well developed, usually
exceeding considerably posterior end of body, being supported by two
short, wide lateral lobes, each of them bearing papilla at the base; spicule
without rough transverse grooves on its surface, being often insufficiently
sclerotized; spicular sheath not spiny; vulvar appendage absent; parasites
of digestive tract of fishes.
C. sentinosa TRAVASSOS, 1927: Astyanax fasciatus: Brazil (São Paulo State). Body =
2 x 0.04 mm (male) & 4.5-4.7 x 0.07 mm (female); Spicule = 89 µm; Egg =
41-49 x 20-23 µm.
Freitascapillaria MORAVEC, 1982
Capillariidae. Stichosome consisting of single row of stichocytes; lateral
caudal alae in male absent; posterior end of male rounded, distinctly
laterally expanded, without distinct papillae, with terminal cloacal opening;
spicule allegedly absent (short, slightly sclerotized spicule probably
present); spicular sheath not spiny; vulvar appendage absent; parasites of
digestive tract of fishes.
F. maxillosa (VAZ & PEREIRA, 1934): Salminus maxillosus: Brazil (São Paulo State).
Body = 6 x 0.07 mm (male) & 16.3 x 0.10 mm (female); Egg = 62 x 28 µm.
Paracapillaria MENDOÇA, 1963

Capillariidae. Stichosome composed of single row of stichocytes. Lateral
caudal alae in male absent. Membranous bursa present, supported by two
lateral digital projections (rays) bent along posterior border of bursa to
dorsal side; dorsal caudal projection absent; pair of large sessile adanal or
postanal papillae present. Spicule without rough transverse grooves on its
surface. Spicular sheath not spiny. Vulvar appendage absent. Parasites of
digestive tract of cold-blooded vertebrates.
P. piscicola (TRAVASSOS, ARTIGAS & PEREIRA, 1928): Acestrorhamphus sp.: Brazil
(São Paulo State). (Fig. 6-1A-B). Body = 4.2 x 0.044 mm (male) & 6 x
0.056 mm (female); Egg = 45 x 18 µm.
Physalopteridae RAILLIET, 1893.
Physalopteroidea. Cephalic end with two large, unlobed lateral pseudola-
bia, usually provided with teeth on their inner surface. Interlabia absent.
Papillae of external circle fully fused together, situated on pseudolabia.
Stoma strongly reduced. Inflation of cuticle (cephalic collarette) posterior
to pseudolabia sometimes present. Male caudal papillae pedunculate,
caudal alae well developed, often forming unification on ventral side of
body (vesicule); or caudal papillae sessile and caudal alae absent. Vulva of
female usually pre-equatorial. Uteri sometimes numerous.
Heliconema TRAVASSOS, 1919.
Physalopteridae. Internal border of each pseudolabium carrying only an
iternolateral tooth and, sometimes, a simple tooth at each dorsoventral
extremity. Spicules of male differing in size and shape. Usually ten pairs
of pedunculate caudal papillae present. Vulva of female slightly pre-
equatorial in position. Ovejector very long. Two uteri present. Ovipa-
rous. Parasites of fishes.
H. izecksohni FÀBIO, 1982. Hoplias malabaricus. Brazil (Rio de Janeiro). (Fig. 6-
96). Body = 32 x 0.15 mm (male) & 23 x 0.17 mm (female); Egg = 65-72
x 44-50 µm.
VIII. Plates of Nematoda (Figs. 6-1 to 6-98)
6-1 A
6-1 B
40 µm
6-1. Paracapillaria piscicola: female; A. entire; B. egg (after TRAVASSOS et al. 1928).
6-2 A
6-2 B
6-3 A
6-3 B
6-2. Rhabdochona elegans: male; A. head; B. tail (after TRAVASSOS et al. 1928); 6-3. Cystidicoloides fischeri:
male; A. tail; B. head (after TRAVASSOS et al. 1928).
6-6
6-4 6-5
6-7
6-8 A
6-9
6-8 B
6-4. Cosmoxynemoides aguirrei: female, head; 6-5. Rondonia rondoni: female, head; 6-6. Cosmoxynema viannae:
female, head; 6-7. Klossinemella iheringi: female, head; 6-8. Spectatus spectatus: male; A. head; B. tail; 6-9.
Travnema travnema: female, head (Figs. 6-4 to 6-9 after TRAVASSOS et al. 1928).
6-10 B
6-10 A
6-10 C 6-10 D
6-10. Spinitectus jamundensis: female; A. head; B. “en face”; C. tail; D. tail; male (after THATCHER &
PADILHA 1977).
6-11 A
6-11 B
6-11 C
6-11 D
6-11. Echinocephalus dailey: female; A. anterior end; B. lips (lateral view); C. lips (apical); male; D.
anterior end (all after DEARDORFF et al. 1981).
6-12 A 6-12 B
6-13
6-14 A
6-15
6-14 B
6-12. Terranova serrata: male; A. head; B. spicules; 6-13. Neocucullanus neocucullanus: female, head; 6-14.
Nilonema senticosa: female; A. head; B. tail. 6-15. Neocucullanus neocucullanus: male, tail (6-12 to 6-15 after
TRAVASSOS et al. 1928).
6-16 A
6-16 B
6-17 A
6-17 B
6-16. Cucullanus schubarti: A. head end (female); B. tail (male) (after PETTER 1987); 6-17. Paraseuratum
albidum: A. head end (female); B. tail (male) (after PETTER 1987).
6-18 A
6-19 A
6-18 B
6-19 C
6-19 B
6-18. Guyanema seriei: A. head end (female); B. tail (male) (after PETTER 1987); 6-19. Chabaudinema
americana: female; A. head end; B. head; male; C. tail (after DÍAZ-UNGRÍA 1968).
6-20 A
6-20 C
6-20 B
6-21 A
6-20. Raphidascaris (Sprentascaris) mahnerti: A. “en face”; B. tail (male); C. head end (female) (after PETTER
& CASSONE 1984); 6-21 A. Goezia spinulosa: tail (male) (after SANTOS, VICENTE & JARDIM 1979).
6-22 A
6-21 B
6-22 B
6-21 B. Goezia spinulosa: head end (female) (original); 6-22. Touzeta ecuadoris: A. head end; B. tail (male)
(after PETTER 1987).
6-23 A 6-23 B
6-24 A
6-23. Rondonia rondoni: female; A. anterior extremity; B. head and mouth; 6-24 A. Procamallanus
(Spirocamallanus) inopinatus: female; apical view.
6-24 B
6-25
6-24 B. Procamallanus (Spirocamallanus) inopinatus: female, buccal capsule; 6-25. Camallanus tridentatus:
female, buccal capsule.
6-26 6-27
6-28 6-29 6-30
6-26 to 6-30. Head ends of Camallanidae: 6-26. Procamallanus (Spirocamallanus) rarus (showing teeth in
base of capsule); 6-27. Procamallanus (S.) pimelodus; 6-28. Procamallanus (Procamallanus) peraccuratus; 6-
29. “Procamallanus sp.” (showing 3 teeth in base of capsule); 6-30. Paracamallanus sp. (showing post-
valvular chamber and tridents). (All original photomicrographs.)
6-31 6-32
6-33 6-34
6-31 to 6-34. Camallanus spp.: 6-31. C. acaudatus (male; head); 6-32. C. tridentatus (female; apical view);
6-33. C. acaudatus (3rd stage larva; showing post-valvular chamber); 6-34. C. acaudatus (3rd stage larva
in transition to 4th stage). (All original photomicrographs).
6-35
A B
6-35. Camallanus acaudatus: female; A. anterior expansion; B. anterior extremity; C. buccal capsule.
(All original photomicrographs).
6-36 6-37
6-38
6-36. Spinitectus sp. 6-37. Spinitectus asperus: female, head end; 6-38. Cucullanus grandistomis: female,
head and mouth. (All original photomicrographs).
6-40
6-39
6-41
6-42
Paracamallanus amazonensis: 6-39. female head, lateral view (scale = 50 µm); 6-40. female head “en-face”
(scale = 25 µm); 6-41. male head, ventral view (scale = 25 µm); 6-42. Male tail, ventral (scale = 50 µm).
6-43
6-45
6-46
6-48
6-47
6-44
6-49
Klossinemella iheringi: 6-43. anterior of female, lateral view; 6-44. female entire (showing eggs); 6-45.
female mouth “en face”; 6-46. head female, dorso-ventral view; 6-47. spicules and gubernaculum;
6-48. tail of female; 6-49. tail of male (from MORAVEC & THATCHER 1997 a). Figs. 6-43 to 6-98 are
reproduced here by permission of the editors of Folia Parasitologica, Czech Republic.
6-50
6-53
6-54
6-55
6-51
6-52
6-50 to 6-52. Spiroxyuris annulata: female; 6-50. mouth “en face”; 6-51. entire; 6-52. egg; 6-53 to 6-55.
Ichthyouris ovifilamentosa: female; 6-53. mouth “en face”; 6-54. entire; 6-55. egg (from MORAVEC &
THATCHER 2001).
6-56
6-58
6-59
6-57
Spinoxyuris annulata: female; 6-56. anterior end, ventral view; 6-57. caudal region, dorsal view; 6-58.
mouth “en face”; 6-59. mouth, sub-lateral view (from MORAVEC & THATCHER 2001).
6-60
6-61
6-62 6-63
Raphidascaroides brasilienses: 6-60. anterior extremity and lips; 6-61. posterior extremity of female showing
papillae; 6-62. “Spirocamallanus” sp. “en face” from Myleus schomburgkii; 6-63. Cucullanus sp. “en face”
from Mylesinus paraschomburgkii (from MORAVEC & THATCHER 1997c).
6-67
6-64 6-68
6-65
d
n 6-66
6-69
Myleusnema bicornis: 6-64. mouth of female “en face”; 6-65. head of female, lateral view; 6-66. anterior end of
female, lateral view; d = deirid, n = nerve ring; 6-67. female entire; 6-68. male entire; 6-69. egg (from
MORAVEC & THATCHER 1996).
6-70
6-71
6-72 6-73
Myleusnema bicornis: 6-70. mouth female “en face”; 6-71. tail of female; 6-72. cloacal aperture of male
showing horns and spicules; 6-73. posterior extremity of male (from MORAVEC & THATCHER 1996).
6-75
6-74
6-76
6-77
Myleusnema brasiliense: 6-74. mouth “en face”; 6-75. tail of male; ventral view; 6-76. head of female,
lateral view; 6-77. cloacal aperture of male showing postcloacal horns and spicules (from MORAVEC
& THATCHER 1999).
6-79
6-78
6-80
6-78. Klossenemella iheringi; 6-79. Procamallanus (D.) dentatus; 6-80. Procamallanus (S.) inopinatus (from
MORAVEC & THATCHER 1997a).
6-81 6-82
6-83
6-84
6-85
6-86 6-87
Procamallanus (Denticamallanus) dentatus: 6-81. head of female; 6-82. head of male; 6-83. apical view of
female; 6-84. tail of male; 6-85. anterior end of male; 6-86. posterior end of female; 6-87. posterior
end of male (from MORAVEC & THATCHER 1997 b).
6-88
6-90
6-89
6-91
6-92
6-93
Cucullanus mogi from Leporinus sp.: 6-88. anterior end of female; 6-89. “en face” view of female; 6-90.
posterior end of male; 6-91. egg; 6-92. posterior extremity of female; 6-93. posterior extremity of
male (from MORAVEC 1998).
6-94 6-95 A
6-95 B
6-96
6-98
6-97
6-94. Travassosnema travassosi: anterior end of female; 6-95. Brasilnema pimelodella: A. anterior extremity
female; B. head of female; 6-96. Heliconema izecksohni: head end of female; 6-97. Neoparaseuratum travassosi:
anterior extremity; 6-98. Terranova serrata: anterior extremity (all redrawn from MORAVEC 1998).
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FERRAZ, E. & V.E. THATCHER (1991): Paracamallanus amazonensis sp. n. (Nematoda: Camallanidae) um
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Camallanidae) from Trachycorystes insignis (STEINDACHNER) (Pisces: Doradidae) in Colombia. -
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HAMANN, M.I. (1982a): Parásitos del pacú (Colossoma mitrei) del rio Paraná Medio, República
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HAMANN, M.I. (1982b): Parásitos em peces de la familia Doradidae del río Paraná Medio, República
Argentina (Pisces Siluriformes). - Hist. Nat. 2: 193-199.
HAMANN, M.I. (1983): Parásitos del sábalo (Prochilodus platensis HOLMBERG, 1889) del río Paraná
Medio, República Argentina (Pisces, Tetragonopteridae). - Hist. Nat. 2: 193-199.
HAMANN, M.I. (1984): Nemátodos parásitos de peces pimelodidos del río Paraná Medio, República
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HAMANN, M.I. (1985): Presencia de Cucullamus pinnai TRAVASSOS, ARTIGAS y PEREIRA (1928) en peces
del río Paraná Medio, Provincia de Corrientes, República Argentina (Nematoda, Cucul-
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KLOSS, G.R. (1966): Helmintos parasitos de espécies simpátricas de Astyanax (Pisces, Characidae). I. -
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KOHN, A. & B.M.M. FERNANDES (1988): Helminth parasites of fishes from the hydroelectric power
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(Nematoda: Camallanidae) from the reservoir of “Salto Osório”. - Mem. Inst. Oswaldo Cruz
83(3): 293-298.
KOHN, A., FERNANDES, B.M.M., MACEDO, B. & B. ABRAMSON (1985): Helminth parasites of
freshwater fishes from Pirassununga, SP, Brazil. - Mem. Inst. Oswaldo Cruz 80(3): 327-336.
KOHN, A., FERNANDES, B.M.M., PIPOLO, H.V. & M.P. DE GODOY (1988): Helmintos parasitos de
peixes das usinas hidroeléctricas de Eletrosul (Brazil). II. Reservatórios de Salto Osório e de
Salto Santiago, Bacia do rio Iguaçu. - Mem. Inst. Oswaldo Cruz 83(3): 229-303.
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MAGALHÃES PINTO, R. & D. NORONHA (1972): Redescrição de Procamallanus inopinatus TRAVASSOS,
ARTIGAS & PEREIRA, 1928 (Nematoda, Camallanoidea). - Atas Soc. Biol. Rio de Janeiro 15(2):
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MAGALHÃES PINTO, R., FÁBIO, S.P. DE, NORONHA, D. & T. ROLAS (1974): Procamallanus brasileiros -
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MAGALHÃES PINTO, R., FÁBIO, S.P. DE, NORONHA, D. & T. ROLAS (1975): Novas contribuições ao
conhecimento do gênero Procamallanus (Nematoda, Camallanoidea). - Mem. Inst. Oswaldo
Cruz 73(3): 183-191.
MAGALHÃES PINTO, R., FÁBIO, S.P. DE, NORONHA, D. & T. ROLAS (1976a): Novas considerações
morfológicas e sistemáticas sobre Procamallanus brasileiros. - Mem. Inst. Oswaldo Cruz 74(1): 77-84.
MAGALHÃES PINTO, R., FÁBIO, S.P. DE, NORONHA, D. & T. ROLAS (1976b): Procamallanus brasileiros
(Nematoda, Camallanoidea). Considerações finais, com chave para determinação das espécies. -
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MORAVEC, F. (2001): Trichinelloid nematodes parasitic in cold-blooded vertebrates. - Academia,
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MORAVEC, F. & V.E. THATCHER (1996): Myleusnema bicornis gen. et sp. n. (Nematoda: Kathlaniidae) an
intestinal parasite of a freshwater serrasalmid fish, Myleus ternetzi, from French Guiana. - Folia
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MORAVEC, F. & V.E. THATCHER (1997a): New data on the morphology and systematic status of
Klossinemella iheringi (Nematoda: Atractidae) from an Amazonian serrasalmid fish. - Folia
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MORAVEC, F. & V.E. THATCHER (1997b): Procamallanus (Denticamallanus subgen. n.) dentatus sp. n.
(Nematoda: Camallanidae) from the characid fish Bryconops alburnoides in the Brazilian
Amazon. - Parasite (France) 4: 239-243.
MORAVEC, F. & V.E. THATCHER (1997c): Raphidascaroides brasiliensis n. sp. (Nematoda: Anisakidae), an
intestinal parasite of the thorny catfish, Pterodoras granulosus from the Brazilian Amazon. -
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MORAVEC, F. & V.E. THATCHER (1999): Myleusnema brasiliense sp. n. (Nematoda: Kathlaniidae) a new
intestinal parasite of the serrasalmid fish, Myleus sp. in Brazil. - Folia Parasit. 46: 216-220.
MORAVEC, F. & V.E. THATCHER (2001): New oxyuroid nematodes of the genera Ichthyouris and
Spinoxyuris from South American freshwater fishes. - Folia Parasit. 48: 311-320.
MORAVEC, F., KOHN, A. & B.M.M. FERNANDES (1990): First record of Raphidascaris (Sprentascaris)
hypostomi (PETTER et CASSONE, 1984) comb. n. and R. (S.) mahnerti (PETTER et CASSONE, 1984)
comb. n. (Nematoda: Anisakidae) from Brazil with remarks on the taxonomic status of the
genus Sprentascaris PETTER et CASSONE, 1984. - Folia Parasitol. 37: 131-140.
MORAVEC, F., KOHN, A. & B.M.M. FERNANDES (1992a): Nematode parasites of fishes of the Paraná River,
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MORAVEC, F., KOHN, A. & B.M.M. FERNANDES (1992b): Three new species of oxyuroid nematodes,
including two new genera, from freshwater catfishes in Brazil. - Syst. Parasitol. 21: 189-201.
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(Nematoda: Quimperiidae), a new parasite from thorny catfish Pterodoras granulosus in Brazil. -
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MORAVEC, F., KOHN, A. & B.M.M. FERNANDES (1993a): Nematode parasites of fishes of the Paraná
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MORAVEC, F., KOHN, A. & B.M.M. FERNANDES (1993b): Nematode parasites of fishes of the Paraná
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MORAVEC, F., KOHN, A. & B.M.M. FERNANDES (1994a): Structure of the cephalic end of two little-
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MORAVEC, F., KOHN, A. & B.M.M. FERNANDES (1994b): Two new species of the genus Goezia, G.
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MORAVEC, F., KOHN, A. & B.M.M. FERNANDES (1997): New observations on seuratoid nematodes
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MORAVEC, F., URAWA, C. & O. CORIA (1997): Philonema percichthydis sp. n. (Nematoda: Philometridae
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PETTER, A.J. (1984): Nématodes de poisons du Paraguay. II. Habronematoidea (Spirurida). Descrip-
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a Colombian freshwater fish, Prochilodus reticulatus STEIND. - Rev. Brasil. Biol. 37(4): 799-801.
TRAVASSOS, L. (1919): Informações sobre o material helmintológico colecionado na Ilha da
Trinidade em 1916. - Arch. Mus. Nac. Rio de Janeiro 22: 159-167.
TRAVASSOS, L. (1923): Informações sobre a fauna helmintológica de Mato Grosso, Oxyuroidea,
Kathlaniidae. - Folha Méd. 4(4): 29-30.
TRAVASSOS, L. (1927): Una nova espécie parasita de peixes de água doce, Capillaria senticosa n. sp. -
Bol. Biol. 10: 215-217, 4 figs.
TRAVASSOS, L. (1948a): Contribuição ao conhecimento dos helmintos dos peixes de água doce do
Brasil. III. Duas novas espécies do gênero Cucullanus MUELLER, 1877. - Mem. Inst. Oswaldo
Cruz 45(3): 551-554.
TRAVASSOS, L. (1948b): Contribuição ao conhecimento dos helmintos dos peixes de água doce do
Brasil. IV. Dois novos gêneros de Cosmocercidae (Nematoda) e uma nota de nomenclatura
helmintológica. - Mem. Inst. Oswaldo Cruz 46(3): 633-637.
TRAVASSOS, L. (1960): Sobre nematódeos cavitários de peixes do Rio Amazonas. - Atas Soc. Biol. Rio
de Janeiro 4(2): 15-20, 14 figs.
TRAVASSOS, L. & A. KOHN (1965): Lista dos helmintos de peixes encontrados na Estação
Experimental de Biologia de Emas, Pirassununga, Estado de São Paulo. - Pap. Avuls. Dept.
Zool. Univ. São Paulo 17: 35-42.
TRAVASSOS, L., ARTIGAS, P. & C. PEREIRA. (1928): Fauna helmintológica dos peixes de água doce do
Brasil. - Arq. Inst. Biol. São Paulo 1: 5-68.
VAZ, Z. & C. PEREIRA (1935): Contribuição ao conhecimento dos nematóides de peixes fluviais do
Brasil. - Arq. Inst. Biol. São Paulo 5: 87-103.
YAMAGUTI, S. (1961): Systema Helminthum. Vol. 3, parts 1 & 2. The Nematodes of Vertebrates. -
Intersci. Publ. New York: 1261 pp.
7.
ACANTHOCEPHALA
Acanthocephalans are stout to elongate worms provided anteriorly with an eversible spiny
proboscis. The body cavity is a pseudocoel and there is no mouth or digestive tract. The
sexes are separate with females normally larger than males of the same species. Larvae are
parasitic in arthropods, adults in vertebrates and there are no free-living stages in the cycle.
Adult acanthocephalans found in Neotropical fishes range in length from less than
two millimeters to more than 80 centimeters, but most are in the one to three centimeters
range. The body is composed of the proboscis, bearing hooks or spines, an unspined neck
region and a trunk, which may or may not be spinous. Living acanthocephalans may be
either flattened or rounded in cross section and they are sometimes confused with either
nematodes or tape worms. Recognition features are: the proboscis, proboscis sheath,
elongate glands associated with the sheath (lemnisci) and a reticulate network of tubules
in the body wall (lacunar system). Most of these characters can usually be seen in living
specimens under a dissecting microscope.
The armature of the proboscis consists of thorn-like hooks with widened roots or
simple rootless spines. The number and arrangement of these hooks are fairly constant
within a species and are of primary importance in the systematics of the phylum. The
hooks are usually placed in longitudinally spiraling rows or horizontally circular ones. The
neck region, between the proboscis and the trunk, is unspined.
The trunk is a tubular structure with longitudinal and circular muscle bands in its
walls. Anteriorly, there is a muscular sac (proboscis sheath) to receive the inverted
proboscis and this sheath may be either single or double walled. Within the hypoderm of
the body wall, there is a conspicuous network of channels, called the lacunar system. The
latter consists of a pair of dorso-ventral or lateral longitudinal canals with anastomosing
horizontal branches. There is a similar but separate lacunar system within the proboscis,
proboscis sheath and lemnisci but these channels are more difficult to see. The lacunar
system probably functions as a primitive circulatory system to distribute nutrients to the
cells. The hypodermal layer itself is a syncytium with numerous small fragmentary nuclei
or a small number of giant nuclei. The number, form and position of the giant nuclei are
useful taxonomic features of acanthocephalans.
The primary reproductive organs are formed within the ligament sacs, two tube-like
structures extending from the posterior end of the proboscis sheath to the posterior end
of the body. The male system consists of two oval or elongate testes followed by cement
glands, a cement reservoir and ducts leading to a penis, which projects into a copulatory
bursa. The cement glands consist of a cluster of six or eight units, each with its nucleus, or
they may be fused into a syncytial mass with a number of giant nuclei. The number and
form of the cement glands and the number of giant nuclei in syncytial cement organs are
relatively constant and are useful in the classification of these worms. Within the genital
sheath, which surrounds the ducts leading to the penis, there is an additional sac-like
organ in some acanthocephalans that is known as Saefftigen’s pouch. This structure is
thought to lubricate the eversion of the copulatory bursa.
The ovaries arise in the ligament sacs of the female as one or two spherical masses
that soon break up to form numerous ovarian balls. The ovarian balls float freely within
the ligament sacs and in some acanthocephalans they are released into the body cavity by
rupture of the walls of the sacs. Eggs released by the ovarian balls also float freely until
they are mature at which time they enter the expanded proximal portion of the uterus
(uterine bell). From there, they pass down the uterus and through the vagina to be
expelled through the vaginal pore. Mature eggs usually consist of fully formed larvae,
armed with hooks, surrounded by three or four membranes.
In copulation, the male covers the posterior extremity of the female with his copulatory
bursa and glues it in place with secretions from the cement glands. The penis may then be
inserted into the vagina and spermatozoa injected into the uterus. The copulatory bursa
of the male sticks to the female with such tenacity that sometimes copulating pairs can be
collected. Also, when the bursa is removed by the male, a cap of cement usually remains
on the female for some time. It is therefore possible to tell if a female has mated.
Acanthocephalan eggs contain six-hooked larvae called acanthors when expelled
into the environment in the feces of the definitive host. The larvae do not develop further
until the eggs are eaten by appropriate intermediate hosts. In the case of aquatic
acanthocephalans, the intermediate host is usually a microcrustacen (ostracod, copepod,
amphipod or isopod), In the gut of the crustacean, the egg hatches and the acanthor larva
penetrates the gut wall by means of its hooks. In the hemocoel of the intermediate host,
the acanthor goes through a transformation losing its larval hooks and growing an adult
type hooked proboscis. This second larval stage is called an acanthella.
If the intermediate host containing an acanthella is eaten by the right definitive host,
development to the adult stage proceeds in the gut of the latter. If the acanthella is
ingested by an inappropriate host, it penetrates the intestinal wall and encysts to await later
passage to the right definitive host. Such hosts are called transport hosts since no further
development of the acanthella occurs in them. Fish can serve as either definitive hosts or
transport hosts, so it is common to find encysted acanthellae in the mesenteries, liver and
other organs of small fish. Fish then acquire adult acanthocephalans either by ingesting
crustacean intermediate hosts or small fish that serve as transport hosts.
III. Pathology
Little pathology has been observed in Neotropical fish in association with acantho-
cephalan infection. Of course, some of them embed their hooked proboscides in the
intestinal wall provoking localized inflammatory reactions. In the case of Megapriapus
ungriai in the stingray, the spherical proboscis is buried in the intestinal wall long
enough for a fibrotic capsule to be formed around it. It has been suggested that
acanthocephalans can perforate the wall of a host and bring on peritonitis. We have
not yet been able to verify this phenomenon in Neotropical fish infections. We have
seen, however, a considerable amount of liver damage brought about by the presence
of acanthellas and young adults of Palliolisentis sp. in Triportheus elongatus. The same
fish hosts had adults of the same acanthocephalan in their digestive tracts. No
explanation is presently available for this phenomenon.
Aquarium fish should not be fed wild-caught microcrustaceans if the build-up of

encysted acanthellas in them is to be avoided. No treatment is known for encysted forms,
but adults in the intestinal tract can frequently be expelled by giving Di-N-Butyl Tin
Oxide mixed in the ration at 0.3 % of body weight for one to five days.
Acanthocephalans are best collected alive from recently killed fish. If the proboscides are
embedded in the intestinal wall, it is necessary to dissect them out carefully with needles.
Once removed from the fish, the worms should be placed in water. It will be found that
most acanthocephalans invert the proboscis and they should not be killed and fixed in this
position. If left in water for some hours, the water penetrates the body making the worms
turgid with the proboscis extended. When this stage is reached, they can be killed and
fixed by dropping them into hot AFA solution. After fixing overnight, permanent whole-
mount slides may be prepared with Mayer’s carmalum stain and alcohol dehydration
(explained in detail in Chapter 4). Acanthocephalans are difficult to dehydrate, however,
so after fixing and before staining, their body walls should be perforated in two or three
places preferably with glass needles.
VI. Identification and key
302
Some of the characters used in the identification of the acanthocephalans are: body shape and size, proboscis shape and size; proboscis

hook shape, size and number; proboscis sheat size, form and number of layers in the wall, lemnisci length; testes size and position;
cement gland form and number of nuclei; and egg size and form.
Key to Acanthocephala of Neotropical freshwater fishes
I. Trunk spines present.

A. Trunk spines in limited number (4 – 10) circular rows at anterior extremity; hypodermic nuclei large, frequently dendritic.
1. Proboscis with 18 hooks located in 4 circular rows .................. Acanthogyridae ............................................. Acanthodelta (Fig. 7-6)
2. Proboscis with more than 18 hooks ...................................................................................................................................... Quadrigyridae
a. Proboscis hooks in 6 – 12 longitudinal rows of 6 – 8 hooks each ................................................................................ Palliolisentis
b. Proboscis hooks in 4 circular rows of 5 hooks each ...................................................................................... Quadrigyrus (Fig. 7-3)
B. Trunk spines extensive sometimes reaching to near posterior extremity or limited to 8 – 10 circular rows anteriorly.
Hypodermic nuclei small, oval ............................................................................................................................................ Rhadinorhynchidae
1. Trunk spines extensive ................................................................................................................................... Rhadinorhynchus (Fig. 7-1 A-I)
2. Trunk spines limited to 8 – 10 circular rows anteriorly
a. Proboscis hooks numerous .................................................................................................................... Polyacanthorhynchus (Fig. 7-26)
b. Proboscis with 18 hooks ................................................................................................................ Pseudogorgorhynchus (Fig. 7-25 A-D)
II. Trunk spines absent.
A. Hypodermic nuclei small.
1. Neck long frequently twisted and may have bulbous swelling ....... Pomphorhynchidae ............................... Pomphorhynchus (Fig. 7-16)
2. Neck short, straight ............................................................................................................................................................ Echinorhynchidae
a. Proboscis cylindrical, hooks in 14 longitudinal rows; from characins ............... Echinorynchus (Figs. 7-18, 7-19, 7-29 & 7-30)
b. Proboscis spherical or fusiform; hooks in 20 – 26 longitudinal rows; from rays ...................Megapriapus (Figs. 7-31 & 7-32)
c. Proboscis club-shaped; hooks in 45 – 48 longitudinal rows; from catfish ............................................. Paracavisoma (Fig. 7-17)
B. Hypodermic nuclei large (giant) ...................................................................................................................................... Neoechinorhynchidae
1. Proboscis with 88 hooks (22 longitudinal rows of 4 each) .................................................................................... Pandosentis (Fig. 7-8)
2. Proboscis with 48 hooks (3 circular rows of 16 each) .......................................................................................... Wolffhugelia (Fig. 7-2)
3. Proboscis with 36 hooks (3 circular rows of 12 each) ............................................................................................................ Gracilisentis
4. Proboscis with 21 – 30 hooks (3 circular rows of 7 – 10 each) ......................................... Octospiniferoides (Figs. 7-7, 7-20 to 7-24)
5. Proboscis with 18 hooks
a. Hooks in 3 circular rows of 6 each; dorsal “fin-fold” present on trunk ..................... Gorytocephalus (Figs. 7-5, 7-20 to 7-24)
b. Hooks in 6 diagonal rows of 3 each; dorsal “fin-fold” absent .................................. Neoechinorhynchus (Figs. 7-4 & 7-10 A-E)
VI. Identificación y clave
Algunos caracteres usados en la identificación de acantocéfalos son: forma y tamaño del cuerpo; forma y tamaño de la proboscis; forma,
tamaño y número de ganchos en la proboscis; tamaño del receptáculo de la proboscis; forma y número de capas en la musculatura del
receptáculo de la proboscis; largo de los lemniscos, tamaño y posición de los testes; forma y número de núcleos en las glándulas de

cemento y forma y tamaño de los huevos.
Clave de identificación para Acanthocephala de peces Neotropicales de agua dulce
I. Con espinas en el tronco.

A. Tronco con espinas en número limitado (4 – 10) filas circulares en la extremidad anterior; núcleos hipodermicos grandes,
frecuentemente dendrítico.
1. Proboscis con 18 ganchos localizados en 4 filas circulares ....... Acanthogyridae ............................................. Acanthodelta (Fig. 7-6)
2. Proboscis con más de 18 ganchos ......................................................................................................................................... Quadrigyridae
a. Ganchos de la proboscis en 6 – 12 filas longitudinales con 6 – 8 ganchos cada una ................................................ Palliolisentis
b. Ganchos de la proboscis en 4 filas circulares con 5 ganchos cada una ...................................................... Quadrigyrus (Fig. 7-3)
B. Espinas del tronco extensas alcanzando algunas veces hasta cerca de la extremidad posterior o limitadas a 8 – 10 filas circulares
anteriores; núcleos hipodérmicos pequeños y ovales .......................................................................................................... Rhadinorhynchidae
303
1. Espinas del tronco extensivas ...................................................................................................................... Rhadinorhynchus (Fig. 7-1 A-I)
304
2. Espinas del tronco limitadas a 8 – 10 filas circulares anteriores
a. Numerosos ganchos en la proboscis .................................................................................................... Polyacanthorhynchus (Fig. 7-26)

b. 18 ganchos en la proboscis ........................................................................................................... Pseudogorgorhynchus (Fig. 7-25 A-D)
II. Espinas ausentes en el tronco.
A. Núcleos hipodérmicos pequeños.
1. Cuello largo frecuentemente torcido, puede tener hinchazón en forma de bulbo ...............................................................................
.............................................................................................................. Pomphorhynchidae ............................... Pomphorhynchus (Fig. 7-16)
2. Cuello corto, recto .............................................................................................................................................................. Echinorhynchidae
a. Proboscis cilíndrica, ganchos en 14 filas longitudinales; en caracídeos ............ Echinorynchus (Figs. 7-18, 7-19, 7-29 & 7-30)
b. Proboscis esférica o fusiforme; ganchos en 20 – 26 filas longitudinales; en rayas ................. Megapriapus (Figs. 7-31 & 7-32)
c. Proboscis en forma de bate; ganchos en 45 – 48 filas longitudinales; en bagres ................................. Paracavisoma (Fig. 7-17)
B. Núcleos hipodérmicos grandes (gigantes) .................................................................................................................... Neoechinorhynchidae
1. Proboscis con 88 ganchos (22 filas longitudinales con 4 por fila) ........................................................................ Pandosentis (Fig. 7-8)
2. Proboscis con 48 ganchos (3 filas circulares con 16 por fila) .............................................................................. Wolffhugelia (Fig. 7-2)
3. Proboscis con 36 ganchos (3 filas circulares con 12 por fila) ................................................................................................ Gracilisentis
4. Proboscis con 21 – 30 ganchos (3 filas circulares con 7 – 10 cada una) ........................... Octospiniferoides (Figs. 7-7, 7-20 a 7-24)
5. Proboscis con 18 ganchos
a. Ganchos en 3 filas circulares con 6 cada una; tronco con aleta dorsal .........................Gorytocephalus (Figs. 7-5, 7-27 & 7-28)
b. Ganchos en 6 filas diagonales de 3 cada una; aleta dorsal ausente ........................... Neoechinorhynchus (Figs. 7-4 & 7-10 A-E)
VII. Checklist of Acanthocephala from Neotropical freshwater fishes
In the Checklist, body sizes are given in millimeters and egg sizes in µm (= micrometers).
Acanthogyridae THAPAR, 1927

Neoechinorhynchidea. Trunk spines in circular or longitudinal rows.
Proboscis small, with small number of hooks in transverse rows; proboscis
sheath single-layered. Lacunar system with regular circular canals. Cement
gland double. Intestine of fishes.
Acanthodelta DÍAZ-UNGRÍA & GRACIA RODRIGO, 1958
Body long, slender; proboscis spherical with 18 hooks in four circular rows
(first 3 rows of 4 hooks each; 4th row with 6 hooks). Trunk with 8 rows of
small spines; giant nuclei present. Proboscis sheath single; lemnisci long,
slender (more than 5 times length of proboscis). Testes tandem, in anterior
half; cement glands elongate. Female unknown. Intestine of fishes.
A. scorzai (DÍAZ-UNGRÍA & GRACIA RODRIGO, 1957): catfish (Pimelodidae):
Venezuela (Orinoco River. (Fig. 7-6). Body of male = 16.6 x 0.95.
Echinorhynchidae COBBOLD, 1879
Body long, rarely spinous; trunk nuclei small, numerous. Proboscis usually
elongate, with numerous hooks; sheath wall double, inserted at base of
proboscis. Lemnisci short. Testes oval; cement glands pyriform, 4-8 in
number. Eggs elliptical to fusiform. Intestine of fishes.
Echinorhynchus ZOEGA in MÜLLER 1776 (Figs. 7-18, 7-19, 7-29 & 7-30).
Echinorhynchidae. Proboscis hooks smaller towards proboscis base. Cen-
tral ganglion near middle of proboscis sheath. Digestive tract of fishes.
E. briconi MACHADO FILHO, 1959: Brycon hilarii: Brazil. (Fig. 7-18). Body = 9-15
(male); 15-22 (female); Hooks = 14 longitudinal rows with 16-17 per row.
Egg = 121 x 16 µm.
E. gomezi MACHADO FILHO, 1949: Pacu nigricans: Brazil. (Fig. 7-19). Body = 10-
12 (male); 20-22 (female); Hooks = 14 longitudinal rows with 13 per row.
Egg = 168 x 21 µm.
E. jucundum TRAVASSOS, 1923: Colossoma bidens: Brazil. Body = 10-15 (male);
20-25 (female); Hooks = 14 longitudinal rows with 16-17 per row. Egg =
125 x 21 µm.
E. paranense MACHADO FILHO, 1959: Triportheus paranensis: Brazil (Matto Gros-
so). Body = 7-10 (male); 9-12 (female); Hooks = 14 longitudinal rows with
11 per row.
E. salobrense MACHADO FILHO, 1948: Mylosoma paraguayensis: Brazil (Matto
Grosso). Body = 8-10 (male); 10-14 (female); Hooks = 14 longitudinal
rows with 16-17 per row.
Meapriapus GOLVAN, GRACIA RODRIGO & DÍAZ-UNGRÍA, 1964
Body elongate, slender. Hypodermic nuclei small, fragmented. Proboscis
large, spherical in female; nearly fusiform in male; hooks of three types,
largest being anterior in female and middle of proboscis in male; hooks in
20-26 longitudinal rows of 14-20 per row. Sheath double-walled, inserted

at base of proboscis; lemnisci shorter than sheath. Testes ovoid, separated.
Egg elongate, elliptical. Intestine of freshwater rays.
M. ungriai (GRACIA RODRIGO, 1960): Potamotrygon hystrix: Brazil & Venezuela. (Figs. 7-
9, 7-31 & 7-32). Body = 17-25 x 1.1-1.2 (both sexes). Egg = 70-75 x 15-20 µm.
Paracavisoma KRITSCHER, 1957
Echinorhynchidae. Proboscis claviform, widest near anterior end; hooks
numerous. Intestine of siliruform fishes.
P. impudica (DIESING, 1851) KRITSCHER, 1957: Pseudodoras niger: Brazil & Peru.
(Fig. 7-17). Body = 12-14 X 1.0-1.2 (female); Hooks = 45-48 longitudinal
rows with 23-27 per row.
Neoechinorhynchidae VAN CLEAVE, 1919
Body elongate or ovoid, without spines. Proboscis variable; sheath inserted
at base; wall single. Trunk nuclei few, large. Two lemnisci present. Testes
oval to elliptical; cement gland syncytial, cement reservoir present. Eggs
elliptical, without polar prolongations of middle shell. Gut of fish,
amphibians and turtles.
Gorytocephalus NICKOL & THATCHER, 1971
Body stout, with prominent dorsal crest. Giant nuclei irregular in shape; 5
nuclei dorsal and 1 ventral. Proboscis small, cylindrical, with 18 hooks in 3
circular rows of 6 hooks per row. Distal hooks in two levels; 2 lateral
hooks larger and more anterior than others of row. Single-walled sheath
surrounded by muscular sling; spherical central ganglion at base of sheath.
One lemniscus with 2 nuclei, other with 1; cement reservoir spherical;
Saefftigen’s pouch present. Eggs elliptical. Intestine of freshwater fishes.
G. elongorchis THATCHER, 1979: Hypostomus carinatus: Brazil (Amazon River).
(Figs. 7-27 & 7-28). Body = 12-16 x 1.2-1.6 (male); 12-22 x 0.7-1.7
(female). Egg = 43-46 x 13-14 µm.
G. plecostomorum NICKOL & THATCHER, 1971: Hypostomus plecostomus: Panama. (Figs.
7-11 & 7-12). Body = 10-12 x 07 (both sexes). Egg = 41-43 x 14-17 µm.
G. spectabilis (MACHADO FILHO, 1959): Curimata elegans: Brazil. (Fig. 7-5). Body
= 4-5 x 0.4-0.45 (male); 5.5-6.5 x 0.6-0.5 (female). Egg = 30 x 10 µm.
Neoechinorhynchus HAMANN, 1892
Neoechinorhynchidae. Lacunar system of dorsal and ventral longitudinal
tubules and anastomosing circular ones. Giant hypodermic nuclei usually
4-5 dorsally and 1-2 ventrally. Proboscis short; hooks in 6 spiral rows of 3
per row; anterior hooks larger than posterior ones. Sheath single-walled;
central ganglion near base. Lemnisci with giant nuclei. Testes usually
ovoid; cement gland syncytial; reservoir rounded. Eggs oval to elliptical.
Intestine of fish, amphibians and turtles.
N. buttnerae GOLVAN 1956: Colossoma macropomum: Brazil (Amazonia). Body =
15-22 (male); 20-32 (female). Egg = 22-25 x 9-10 µm.
N. golvani SALGADO-MALDONADO, 1979: Cichlasoma aureum: Mexico. (Fig. 7-13).
Body = 0.9-1.0 (male); 0.7-3.2 (female).
N. paraguayensis MACHADO FILHO, 1959, NICKOL & PADILHA, 1979: Geophagus

brasiliensis and Hoplias malabaricus: Brazil & Paraguay. (Figs. 7-14 & 7-15). Body
= 1.6-3.5 x 0.5-1.0 (male); 2.9-3.8 x 1.1-1.4 (female). Egg = 26-38 x 22-26 µm.
N. prochilodorum NICKOL & THATCHER, 1971: Prochilodus reticulatus: Colombia
(Cauca River). Body = 4.5-8.0 x 1.0-1.7 (both sexes). Egg = 26-29 x 7-12 µm.
N. pterodoridis THATCHER, 1981: Pterodoras granulosus: Brazil (Amazon River).
(Fig. 7-10 A-E). Body = 1.7-2.2 x 0.6-0.9 (male); 2.1-3.3 x 0.6-1.0 (female).
Egg = 24-27 x 7-9 µm.
N. roseum SALGADO-MALDONADO, 1979: Achiurus mazatlanus: Mexico. Body = 6-
7 (male); 7.8-9.3 (female). Egg = 28-36 x 8 µm.
Gracilisentis VAN CLEAVE, 1919
Neoechinorhynchidae. Body small; hypodermic giant nuclei mainly in mid-
dorsal line. Proboscis short, cylindrical, with constriction above basal hook
row; hooks in 3 circular rows of 12 hooks per row; sheath subcylindrical.
Testes contiguous; cement gland syncytial, with 12 nuclei. Eggs elliptical.
Intestine of fishes.
G. variabilis (DIESING, 1856): Hoplias malabaricus, Hypostomus auroguttatus, H.
lituratus, H. plecostomus, H. melanopterus, Monochirus maculipennis and Pleuronect-
es sp.: Brazil. Body = 9-25 (male); 9-50 (female).
Octospiniferoides BULLOCK, 1957
Neochinorhynchidae. Body short, stout. Giant hypodermic nuclei round-
ed, 4-5 dorsally and 1 ventrally. Proboscis short, rounded; hooks in 3
circular rows of 7-10 per row; all hooks rooted. Sheath thin-walled. Testes
ovoid contiguous. Lemnisci longer than sheath; possess giant nuclei. Egg
long, slender. Intestine of fishes.
O. australis SCHMIDT & HUGGHINS, 1973: Chilodus punctatus: Colombia. Body =
4.2-5.0 x 0.70-0.75 (female); Hooks = 3 circular rows of 8 per row. Egg =
30-34 x 10-12 µm.
O. incognita SCHMIDT & HUGGHINS, 1973: Schizodon fasciatum: Bolivia. (Figs 7-7 &
7-20 to 7-24). Body = 2.0 x 0.4 (both sexes).
Pandosentis VAN CLEAVE, 1920
Neoechinorhynchidae. Body small; hypodermic giant nuclei median or
lateral. Proboscis short, cylindrical, with 22 longitudinal rows of 4 hooks
per row. Lemnisci shorter than sheath. Testes contiguous, in middle body
third. Cement gland syncytial, with 16 nuclei. Intestine of fishes.
P. iracundus VAN CLEAVE, 1920: Aequidens pulcher and Crenicichla geayi: Venezue-
la. (Fig. 7-8). Body = 0.6-1.0 (male); 1.0-1.5 (female). Egg = 22-28 x 11 µm.
Wolffhugelia MAÑÉ-GARZÓN & DEI-CAS, 1974
Neoechinorhynchidae. Body small, cylindrical. Hypodermic giant nuclei
ovoid; 5 nuclei dorsal and 1 ventral. Proboscis short, subcylindrical, with 3
circular rows of 16 hooks per row; hooks of three sizes, with anterior row
hooks largest and posterior row smallest. Lemnisci longer than sheath.
Testes contiguous in anterior half of body. Cement gland syncytial, with 8
nuclei. Intestine of fishes.
W.matercula MAÑÉ-GARZÓN & DEI-CAS, 1974: Fitzroyia lineate: Uruguay. (Fig. 7-

2). Body = 0.85-1.9 x 0.35-0.57 (male); 0.95-2.7 x 0.3-0.9 (female). Egg =
27-33 x 10-12 µm.
Pomphorhynchidae YAMAGUTI, 1939
Trunk without spines. Lacunar system of lateral main vessels and reticular
anastomoses. Proboscis cylindrical; hooks numerous. Neck long, cylindri-
cal or spirally twisted, with or without bulbous swelling. Sheath long,
double-walled, inserted at base of proboscis. Testes oval, in mid-body;
cement glands 4 or 6, compact. Egg with polar prolongations of middle
shell. Intestine of fishes.
Pomphorhynchus MONTICELLI, 1905
Pomphorhynchidae. Proboscis long, with 12-20 longitudinal rows of 9-14
hooks per row. Lemnisci short or long. Genital pore of both sexes terminal.
Egg fusiform, with prolongations of middle shell. Intestine of fishes.
P. yamagutii SCHMIDT & HUGGHINS, 1973: Percichthys melanops: Chile. (Fig. 7-
16). Body = 7.0 x 0.6-0.7 (male); 10.0-12.5 x 0.6-0.9 (female). Egg = 66-
70 x 8-10 µm.
Quadrigyridae VAN CLEAVE, 1920
Body small, elongate, with spines in circular rows. Proboscis globular to
elongate, with few hooks in spiral rows; sheath single-walled, with central
ganglion near its base. Hypodermic giant nuclei few or fragmented.
Lacunar system with or without main vessels. Cement gland syncytial. Egg
rounded or elliptical. Intestine of fishes.
Palliolisentis MACHADO FILHO, 1960
Quadrigyridae. Trunk with 5 circular rows of spines anteriorly. Hypodermic
giant nuclei dendritic. Lacunar system annular, without main longitudinal
vessels. Proboscis cylindrical, with 6-12 longitudinal rows of rooted hooks.
Lemnisci claviform, longer than sheath, with one nucleus in each. Testes
contiguous, post-equatorial. Uterus expanded. Intestine of freshwater fishes.
P. ornatus MACHADO FILHO, 1960: Triportheus paranensis: Brazil. Body = 9.7 x
0.8 (male). Trunk spines in 5 rows of 16 per row. Hooks = 6 longitudinal
rows of 8 hooks per row.
P. polyonca SCHIMDT & HUGGHINS, 1973: Acestrorhynchus falcatus: Colombia.
Body = 3.5 x 0.46 (male); 8-10 x 0.76-0.96 (female). Trunk spines in 3-5
anterior rows. Proboscis hooks = 12 longitudinal rows of 6-7 row.
P. quinqueungulis MACHADO FILHO, 1960: Triportheus paranensis and T. angulatus:
Brazil (Matto Grosso). Body = 8.0-8.5 x 0.8-0.85 (male); 13-15 x 1.0-1.2
(female). Trunk spines in 5 rows of 20 per row. Proboscis hooks = 6
longitudinal rows of 8 per row. Egg = 33-54 x 25-30 µm.
Quadrigyrus VAN CLEAVE, 1920
Trunk with 4-10 circular rows of spines anteriorly; sometimes interrupted
dorsally. Hypodermic giant nuclei of two types: anterior ones middorsal and
midventral, others lateral and dendritic. Proboscis hooks in 3-4 circular rows.
Egg oval. Intestine of fishes.
Q. brasiliensis MACHADO FILHO, 1941: Hoplerythrinus unitaeniatus and Hoplias

malabaricus: Brazil. (Fig. 7-3). Body = 10-15 x 0.6-1.3 (male); 13-25 x 1-2
(female). Trunk spines in 3 rows of 12 per row. Proboscis hooks = 4 rows
of 5 per row. Egg = 13-14 x 6-7 µm.
Q. nickoli SCHIMDT & HUGGHINS, 1973: Hoplerythrinus unitaeniatus: Colombia.
Body = 6 x 0.76-0.85 (male); 9.5-10 x 0.94-1.2 (female). Trunk spines in 4
rows of 23-29 per row. Proboscis hooks = 4 rows of 5 per row. Egg = 55-
65 x 28-32 µm.
Q. torquatus VAN CLEAVE, 1920: Hoplias malabaricus, Symbranchus marmoratus,
Crenicichla geayi, Gephyrocharax valenciae and Astyanax bimaculatus: Surinam,
Venezuela, Colombia & Brazil. Body = 8-10 x 0.6 (male); 10-20 x 0.9-1.2
(female). Trunk spines in 3-4 rows of 10-18 per row. Proboscis hooks = 4
rows of 5 per row.
Rhadinorhynchidae TRAVASSOS, 1923
Trunk long, slender, or expanded in anterior or middle third; trunk spined
anteriorly, sometimes spines extend to posterior extremity. Hypodermic nuclei
large and few, or small and numerous. Proboscis usually long; sheath
cylindrical, with ganglion at base or in middle. Testes contiguous, or not;
cement glands 2-8, of various shapes. Egg elongate or oval. Intestine of fishes.
Rhadinorhynchus LÜHE, 1911
Rhadinorhynchidae. Body cylindrical. Hypodermic nuclei small, numer-
ous. Lacunar system with lateral main vessels and reticular anastomoses.
Trunk spines divided into two groups, or not. Proboscis elongate, with 8-
26 longitudinal rows of 8-37 hooks per row. Sheath elongate, with
equatorial central ganglion and double walls. Lemnisci variable. Testes
elongate; cement glands 2-8. Egg elongate, with polar prolongations of
middle shell. Intestine of fishes.
R. plagioscionis THATCHER, 1980: Plagioscion squamosissimus: Brazil (Amazonia).
(Fig. 7-1 A-I). Body = 8.8-11 x 0.7-1.0 (male); 14.5-16.4 x 0.85-1.0 (female).
Trunk spines to near posterior extremity. Proboscis hooks = 12 longitudinal
rows of 23 and 24 hooks in alternate rows. Egg = 110 x 23 µm.
Polyacanthorhynchus TRAVASSOS, 1926
Rhadinorhynchidae. Trunk long, slender, with 8-10 circles of spines
anteriorly. Hypodemic nuclei small, oval, numerous. Proboscis elongate,
claviform, with 16-18 longitudinal row of hooks posteriorly and 26-28
rows anteriorly; hooks of similar shape but diminishing in size posteriorly.
Sheath single-walled; central ganglion equatorial. Spined anterior part of
trunk retractile. Lemnisci longer than sheath. Testes elongate, contiguous,
cement gland 8, long and tubular. Egg oval. Intestine of freshwater fish
and crocodilians.
P. macrorhynchus (DIESING, 1856): Arapaima gigas: Brazil (Amazonia). Body =
60-70 x 1.5 (male); 140 x 2.0 (female). Hooks in 16 longitudinal rows at
base and 15 rows at apex of proboscis, with about 60 hooks per row. Egg
= 71-86 x 38-60 µm.
P. rhopalorhynchus (DIESING, 1851): Arapaima gigas: Brazil (Amazonia). (Figs. 7-

17 & 7-18). Body = 400-555 x 1.5-3.0 (male); 400-700 x 2.5-3.5 (female).
Hooks in 18 longitudinal rows at base and 28 rows at apex of proboscis,
with 55-65 hooks per row. Egg = 220 x 104 µm.
(note: Both of the above species have also been reported from Amazonian
crocodilians, but since the latter often eat Arapaima, these may have been
temporary or spurious infections).
Pseudogorgorhynchus MORAVEC, WOLTER & KÖRTING, 1999.
Rhadinorhynchidae. Trunk long and slender, armed with circular rows of
spines anteriorly. Proboscis short and claviform; hooks few, with simple
roots, arranged in spiral rows; middle hooks distinctly broader than
anterior and posterior hooks; posterior hooks about the same length as
middle hooks. Proboscis receptacle longer than proboscis, with cephalic
ganglion at its base. Lemnisci longer than proboscis receptacle. Testes near
middle of trunk; cement glands 4; seminal vesicle, cement ducts and
Saefftigen’s pouch prominent. Eggs fusiform with polar prolongation of
fertilization membrane. Gonopore subterminal. Type species: Pseudogorgo-
rhynchus arii MORAVEC, WOLTER & KÖRTING, 1999.
P. arii MORAVEC, WOLTER & KÖRTING, 1999: Intestine of Ariopsis seemanni:
Colombia. Male: Body = 3.0-5.7 x 0.45-0.73 mm. Proboscis hooks = 18
hooks arranged in 6 spiral rows of 3 hooks each. Trunk spines in 10-13
transverse rows. Female: Body = 4.4-7.6 x 0.68-0.83 mm.
VIII. Plates of Acanthocephala (Figs. 7-1 to 7-32)
7-1 A
LE
B PS
TE
CG
5.0 mm
C
0.5 mm
7-1 A-C. Rhadinorhynchus plagioscionis (after THATCHER 1980): A. proboscis; B. female (entire); C male
(entire); CG = cement glands; LE = lemnisci; PS = proboscis sheath; TE = testes.
1.0 mm
LE
1.0 mm
NG
CB
PE
7-1
7-1 D-E. Rhadinorhynchus plagioscionis (after THATCHER 1980): D. anterior trunk; LE = lemnisci; NG =
central nerve ganglion; E. posterior region of male; CB = copulatory bursa; PE = penis.
7-1
UF
H
F
G
0.10
0.02
2.0
I
0.20
OB
VS
VA
UE
7-1 F-I. Rhadinorhynchus plagioscionis (after THATCHER 1980): F. egg; G. trunk spine; H. posterior region
of female; OB = ovarian ball; UE = uterus with eggs; UF = uterine funnel; I. posterior extremity of
female; VA = vagina; VS = vaginal sphincter; all scales in mm.
7-2
0.25 mm
7-3
0.5 mm
7-5
7-4
1.0 mm
1.0 mm
7-2. Wolffhugelia matercula: male; anterior end (after MAÑÉ-GARZÓN & DEI-CAS 1974); 7-3. Quadrigyrus
brasiliensis: male; anterior end (after MACHADO FILHO 1941 b); 7-4. Neoechinorhynchus roseum: female;
head end (after SALGADO-MALDONADO 1978); 7-5. Gorytocephalus spectabilis: male; entire (after MACHADO
FILHO 1959 a).
7-6
7-7
7-8
7-9
7-6. Acanthodelta scorzai: head end (after GRACIA-RODRIGO 1959); 7-7. Octospiniferoides incognita: head
end (after SCHMIDT & HUGGHINS 1973); 7-8. Pandosentis iracundus: head end of male (after VAN CLEAVE
1920); 7-9. Megapriapus ungriai: head end of male (after GRACIA-RODRIGO 1959).
7-10 A
C
0.50 mm
0.25 mm
7-10. Neoechinorhynchus pterodoridis: A. male (entire); B. male (proboscis); C. female (tail); D. egg (all
after THATCHER 1981).
DC
7-11 PS MS
NG
LE
7-12
7-13 7-14
7-15
7-11 to 7-12. Gorytocephalus plecostomorum: female (after NICKOL & THATCHER 1971); 7-11. anterior
end; DC = dorsal crest; LE = lemnisci; MS = muscular sling; NG = central nerve ganglion; PS =
proboscis sheath; 7-12. Proboscis; 7-13. Neochinorhynchus golvani: male; entire (after SALGADO-
MALDONADO 1978); 7-14 to 7-15. Neoechinorhynchus paraguayensis: male (after NICKOL & PADILLA 1979);
7-14. entire; 7-15. proboscis.
7-16
7-17
7-18 7-19
7-16. Pomphorhynchus yamagutii: male (after SCHMIDT & HUGGHINS 1973 a); proboscis lateral; 7-17.
Paracavisoma impudica: female, proboscis (after SCHMIDT & HUGGHINS 1973 b); 7-18. Echinorhynchus
gomezi: male proboscis (after MACHADO FILHO 1941 b); 7-19. Echinorhynchus briconi: female; proboscis
(after MACHADO FILHO 1959 a).
7-20
7-21
1 mm
200 µm
7-20. Octospiniferoides incognita SCHMIDT & HUGGHINS, 1973 (after THATCHER 1998): female, entire;
7-21. Octospiniferoides incognita SCHMIDT & HUGGHINS, 1973 (after THATCHER 1998): female, terminal
genitalia.
200 µm
7-23
7-22 1 mm
7-24
100 µm
7-22 to 7-24. Ostospiniferoides incognita SCHMIDT & HUGGHINS, 1973 (after THATCHER 1998): 7-22.
proboscis; 7-23. male lateral view; 7-24. proboscis hooks.
7-25 A B
7-25. Pseudogorgorhynchus ari (redrawn from MORAVEC, WOLTER & KÖRTING 1999): A. male, lateral
view; B. proboscis of female; C. anterior, middle and posterior hook; D. egg.
7-26 7-27
7-28
7-26. Proboscis of Polyacanthorhynchus rhopalorhynchus: male; 7-27. anterior end of Gorytocephalus elongorchis:
male; 7-28. posterior end of Gorytocephalus elongorchis: male.
7-29 7-30
7-31 7-32
7-29. Proboscis of Echinorhynchus sp.: male, from the intestine of Auchenipterichthys longimanus; 7-30.
Proboscis of Echinorhynchus sp.: male, from the intestine of Leporinus fasciatus; 7-31. anterior proboscis
hooks of Megapriapus ungriai: female; 7-32. proboscis of Megapriapus ungriai: female.
CONWAY MORRIS, S. (1982): The origins and evolution of the Acanthocephala. - Biol. Rev. 57: 85-115.
DÍAZ-UNGRÍA, C. & A. GRACIA-RODRIGO (1957): Revisión de la familia Quadrigyridae (Acantho-
cephala) con descripción de Deltania scorzai gen. n. sp. n. - Novedades Cient. (Caracas), ser.
Zool. 22: 1-29.
DÍAZ-UNGRÍA, C. & A. GRACIA-RODRIGO (1959): Acantocéfalos de Venezuela. - Rev. Vet. Venez.
6(31): 82-105.
DÍAZ-UNGRÍA, C. & A. GRACIA-RODRIGO (1960): Revisión del género Polyacanthorhynchus TRAVASSOS,
1920 (Acanthocephala) con descripción de una especie nueva. - In: Libro Homenaje al Dr. E.
CABALLERO y CABALLERO. - Esc. Nac. Cienc. Biol. Y Inst. Politec. Nac. Secret. Educ. Pub.
México: 429-433.
GOLVAN, Y.J. (1956): Acanthocéphales d’Amazone. Redescription Oligacanthorhynchus iheringi TRAVAS-
SOS, 1916 et Neoechinorhynchus buttnerae n. sp. (Neoacanthocephala, Neoechinorhynchidae). -
Ann. Parasit. Hum. Comp. 31(5/6): 500-524.
GOLVAN, Y.J. (1958): Le phylum des Acanthocephala. Premiére note. Sa place dans l’échelle
zoologique. - Ann. Parasit. Hum. Comp. 33(5/6): 538-602.
GOLVAN, Y.J., GRACIA-RODRIGO, A. & C. DÍAZ-UNGRÍA (1964): Megapriapus ungriai (GRACIA-RODRIGO,
1960) n. gen. (Palaecanthocephala) parasite d’une pastenague d’eau douce du Venézuèla
(Potamotrygon hystrix). - Ann. Parasit. Hum. Comp. 39(1): 53-59.
GRACIA-RODRIGO, A. (1959): Acantocéfalos parásitos de los peces de Venezuela. - Ph.D.-thesis, Fac.
Vet. Univ. Cent., Madrid: 32 pp.
MACHADO FILHO, D.A. (1941a): Pesquisas helmintológicas realizadas no Estado de Mato Grosso -
Acanthocephala. - Mem. Inst. Oswaldo Cruz 35(3): 593-601.
MACHADO FILHO, D.A. (1941b): Sôbre alguns acantocéfalos provenientes do Estado de Mato
Grosso. - Rev. Brasil. Biol. 1(1): 57-61.
MACHADO FILHO, D.A. (1947): Revisão do gênero Polyacanthorhynchus TRAVASSOS, 1920 (Acantoceph-
ala, Rhadinorhynchidae). - Rev. Brasil. Biol. 7(2): 195-201.
MACHADO FILHO, D.A. (1954): Uma nova espécie de gênero “Neoechinorhynchus” (HAMANN)
(Neoechinorhynchidae, Acanthocephala). - Rev. Brasil. Biol. 14(1): 55-57.
MACHADO FILHO, D.A. (1959a): “Neoechinorhynchus spectabilis” sp. n. (Neoechinorhynchidae, Acanto-
cephala). - Rev. Brasil. Biol. 19(2): 191-194.
MACHADO FILHO, D.A. (1959b): Uma nova espécie do gênero “Neoechinorhynchus” HAMANN, 1892
parasita de “peixe-martim” do Paraguai (Neoechinorhynchidae, Archiacanthocephala). - Rev.
Brasil. Biol. 19(4): 379-381.
MACHADO FILHO, D.A. (1960): Um novo gênero da família Quadrigyridae VAN CLEAVE, 1929
(Metacanthocephala, Palaeacanthocephala). - Rev. Brasil. Biol. 20(1): 79-84.
MAÑÉ-GARCÓN, F. & E. DEI-CAS (1974): Un acanthocéphale nouveau Wolffhugelia matercula n. gen., n.
sp. d l’intestin de Fitzroyia lineate (JENYNS) (Poisson) en Uruguay. - Ann. Parasit. Hum. Comp.
49(1): 83-89.
MORAVEC, F., WOLTER, J. & W. KÖRTING (1999): Some nematodes and acanthocephalans from exotic
ornamental freshwater fishes imported into Germany. - Folia Parasit. 46: 296-310.
NICKOL, B.B. & T.N. PADILHA (1979): Neoechinorhynchus paraguayensis (Acanthocephala: Neoechino-
rhynchidae) from Brazil. - J. Parasitol. 65(6): 987-989.
NICKOL, B.B. & V.E. THATCHER (1971): Two new acanthocephalans from Neotropical fishes:
Neoechinorhynchus prochilodorum sp. n. and Gorytocephalus plecostomorum gen. et sp. n. - J. Parasitol.
57(3): 576-581.
PETROCHENKO, V.I. (1956): Acanthocephala of domestic and wild animals. Vol. 1. - Akad. Nauk.
U.S.S.R., Moscow (Translated from Russian by Israel, Prog. Sci. Transl., Jerusalem, 1971): 465 pp.
SALGADO-MALDONADO, G. (1978): Acantocéfalos de peces IV. Descripción de dos especies nuevas de
Neoechinorhynchus HAMANN, 1892 (Acanthocephala: Neoechinorhynchidae) y algunas consid-
eraciones sobre este género. - An. Inst. Biol. Univ. Nac. Autón. México 49, ser. Zool. 1: 35-48.
SCHMIDT, G.D. & E.J. HUGGHINS (1973a): Acanthocephala of South American fishes. Part 1.
Eoacanthocephala. - J. Parasitol. 59(5): 829-835.
SCHMIDT, G.D. & E.J. HUGGHINS (1973b): Acanthocephala of South American fishes. Part 2.
Palaeacanthocephala. - J. Parasitol. 59(5): 836-838.
THATCHER, V.E. (1979): Uma nova espécie de Gorytocephalus NICKOL & THATCHER, 1971 (Acantho-
cephala: Neoechinorhynchidae) do acari bodó (Pisces: Loricariidae) da Amazônia, Brasil. -
Acta Amazonica 9(1): 199-202.
THATCHER, V.E. (1980): Rhadinorhynchus plagioscionis n. sp. (Acanthocephala: Rhadinorhynchidae) de
pescada (Plagioscion squamosissimus) da Amazônia brasileira. - Acta Amazonica 10(4): 835-839.
THATCHER, V.E. (1981): Neoechinorhynchus pterodoridis n. sp. (Acanthocephala: Neoechinorhynchidae)
de bacu liso (Pterodoras granulosus) da Amazônia brasileira. - Acta Amazonica 11(3): 445-448.
THATCHER, V.E. (1998): Description of adults of Octospiniferoides incognita SCHMIDT & HUGGHINS
1973, (Acanthocephala, Neoechinorhynchidae) from a fish of Rondônia State, Brazil. -
Amazoniana 15(1/2): 51-55.
THATCHER, V.E. (2001): Brasacanthus sphaeroides gen. et. sp. n. (Acanthocephala: Echinorhynchidae)
from a coastal marine fish of Paraná State, Brazil. - Rev. Brasil. Zool. 18: 1319-1323.
THATCHER, V.E. & B.B. NICKOL (1972): Some acanthocephalans from Panama and Colombia. - Proc.
Helminthol. Soc. Wash. 39(2): 245-248.
VAN CLEAVE, H.J. (1920): Two new genera and species of acanthocephalous worms from
Venezuelan fishes. - Proc. U. S. Nat. Mus. 58: 455-466.
YAMAGUTI, S. (1963): Systema Helminthum. Vol. V. Acanthocephala . - Intersci. Publ., New York:
423 pp.
8.
COPEPODA
Copepods are microcrustaceans, usually less than 3 mm in length, that form an

important part of the zooplankton. As such, they serve as food for many fish species.
Free-living copepods abound in both fresh and salt waters where they feed predaciously
on smaller animals or eat phytoplankton and detritus. Although fish devour copepods,
the latter as a group have gotten their revenge in that they have evolved some forms
that can eat fish tissue.
Among fish parasitic copepods, the ergasiloids (Ergasilidae and related families) are
the most common and best known. In this group, only the females attach to fish though
both sexes have grasping antennae. Species of the genus Ergasilus are well known pests in
pisciculture ponds the world over.
Ergasilidae
Ergasilids superficially resemble the free-living genus Cyclops, and many believe they
evolved from a similar progenitor. The parasitic forms can be recognized at a glance,
however, because they have distinct pigmentation, whereas the free-living species do
not. The pigment granules are usually in the blue spectrum or else in the range of
magenta to purple, and they frequently form distinctive patterns in their distribution
(Figs. 8-57 to 8-60). The tone, intensity and distribution of the pigment granules can be
useful in separating species. Most free-living copepods are white or nearly transparent
except for a pigmented eye. Some high altitude forms are red or orange, but the
coloration in these cases is the result of oil droplets stored in the body as food and not
from the presence of pigment granules.
The ergasilid head is thought to have been derived by the fusion of 6 primitive
segments. Five of these are marked by paired appendages, namely: first antennae (=
antennules); second antennae (= antennae); mandibles; first maxillae (= maxillules); and
second maxillae (= maxillae). The first thoracic segment, which is also fused to the head
unit (= cephalon), bears a pair of maxillipeds (only in the male). In some species, the
second thoracic segment, which bears the first pair of swimming legs, is also fused to the
cephalon. Following the cephalon (also called cephalothorax), there are 3 or 4 free
thoracic segments, each provided with a pair of biramous swimming legs. There are
normally 4 pairs of functional swimming legs in ergasilids and a fifth pair that is vestigial
(in the female). In the male, there is an additional sixth pair of legs that is also vestigial.
Following the free thoracic segments, there is an expanded genital segment which
bears the reproductive pores. This segment is believed to represent the fusion of an
abdominal segment with a thoracic one. This hypothesis is partially supported by the
presence of the sixth legs of the male on the genital segment since abdominal segments
are without appendages. The abdomen consists of four cylindrical segments in the male
and three in the female. The terminal segment is indented medially and contains the anal
aperture. This segment also bears two caudal rami (= uropods) provided with two or three
elongate setae (= caudal filaments).
Ergasilid mouthparts consist of paired: mandibles, which are usually 2-segment-
ed, with spinules or teeth on the terminal part; mandibular palps, that are also
spinulous; first maxillae, which often have 2 setae; and second maxillae, that are larger
and frequently pilose.
The swimming legs of ergasilids (= legs 1-4) are biramous. The inner branch is
termed the endopod and the outer the exopod. Both branches are normally 3-segmented
in free-living cyclopoids and in all known species of ergasilids, legs 2 and 3 have 3-
segmented rami. In some species, the rami of leg 1 are also 3-segmented, but in many
Amazonian forms, the first endopod has only two segments. Some have regarded the 2-
segmented first endopod as an adaptation for parasitism, but it is more probably a
neotenic character resulting from a shortening of the life-cycle. The 4th leg also shows a
reduced number of segments in most species, possibly for the same reason.
The ergasiloid leg was designed for swimming and in most species, it still serves
that function. The legs are provided with feather-like, pilose setae to push better
against the water. Most legs also have a few stout spines which may aid the animal in
grasping a gill filament. Typically, the 2-segmented first endopod has a single medial
seta on the first segment and 5 setae plus 2 stout spines on the terminal segment. The
3-segmented exopod of the first leg usually has a single lateral spine on segment one,
a single medial seta on segment 2 and 5 setae plus 2 spines on the terminal joint. The
3-segmented endopod of legs 2 and 3 frequently has one seta on the first segment, 2
on the second and 4 setae plus one spine on the third. The exopod of the second and
third legs is usually similar to that of the first leg except that it has one more seta and
one fewer spines on the terminal segment. In the Acusicolinae, the parasitic female
has evolved a secure antennal latching mechanism which has made the need for and
the capacity for swimming obsolete. In species of this group, there has been a
reduction in the number and size of the setae and some of those that remain are
pectinate rather than pilose.
The most characteristic feature of ergasiloid copepods that distinguishes them from
their free-living relatives is the prehensile antenna. Both males and females have grasping
antennae, but those of males are usually less than half as large. In Ergasilus, the antenna
consists of a basal segment, about half as long as it is wide, followed by two elongate
segments and a curved claw (Figs. 8-20 to 8-23). Such an antenna has a total of four
segments, if we count the claw. There is some confusion in the literature as to the number
of segments in the ergasiloid antenna because some have considered the claw to be a mere
appendage of the third segment. We have counted the claw as a segment because the joint
between it and the third segment is similar to that between segments two and three. Also,
there is a pit-like sensory organ (sensillum) on it that is similar to those found on other
segments. The antennae of Ergasilus are used to clasp the gill filament and hold the animal
in place. The claws sometimes pierce the tissues of the gill filament.
Several variations on the ergasiloid antenna have been reported. In Acusicola for
example, the same segments are present but the claw is more or less reduced (Figs. 8-18 &
8-19). Segment three is also modified in that it has a groove and segment two has a
cuticular flap that extends out over the groove. This combination of features is used to
latch one antenna to the other. These animals never pierce fish tissue with their claws.
Instead, each claw fits into the groove in the opposite antenna and the flaps partially cover
the latch (Fig. 8-13). These antennae encircle the gill filament as a ring goes around a
finger. Amplexibranchius has antennae of a similar design, but the claw is very small,
segment three is small, segment two is extremely long and the flap is large enough to cover
the claw points when the antennae are latched (Fig. 8-37 F & I).
An entirely different concept in antennal design and use is found in Brasergasilus. In
this genus, there are only three segments in the antennae (Figs. 8-26 to 8-28). This type of
antenna would appear to have evolved from the typical antenna by a failure of the claw to
separate from the preceding segment. The result is an antenna with an unusually long and
slender claw, ideal for piercing tissue. In truth, species of this genus insert the entire
length of the claws into the gill filament and cling very effectively.
An additional attachment method is that of Prehendorastrus BOEGER & THATCHER,
1990, in which the antennae are used individually, one above the other for clinging to a gill
raker. The second, third and fourth (claw) antennal segments are provided with tooth-like
projections one of which seems to serve as a latch. The others may facilitate attachment at
different levels on a tapered gill raker.
Vaigamidae
Vaigamidae is a family of copepods similar to Ergasilidae but differing in a number of

important respects. The most striking difference is the presence of a pair of moveable
retrostylets projecting dorso-laterally from the first thoracic segment (Figs. 8-34 to 8-
36). Some genera also have a rostral spine on the ventral side, between the antennae.
The antennae of Vaigamus are similar to those of Ergasilus, except that they are
proportionally smaller (Fig. 8-29 D-E). In other genera of the family, however, the
antennae end in two moveable claws (Figs. 8-31 B & 8-33 D), or in one fixed spine and
two moveable claws (Fig. 8-32 B). The functions of these antennal structures are not
immediately apparent. Plainly, these antennae would not be effective instruments for
clinging to fish. Vaigamids have no need for strong attachment devices, however, since
they live within the mucous that is found in the nasal fossae of fish. As in Ergasilidae,
only the females are found associated with fish. Males and copepodids are free-living
and not infrequently turn up in plankton.
Vaigamid mouthparts are similar to those of Ergasilidae. Males have a pair of 4-

segmented maxillipeds (Fig. 8-29 C), which females lack. Other features of vaigamids are
also similar to those of ergasilids.
Lernaeidae
The parasitic females of the family Lernaeidae attain a large size due to the fact that they
undergo a kind of metamorphosis after copulating as small free-living females. The
elongate body of the post-metamorphic female is relatively slender and the head has
lateral extensions which serve to anchor it within fish tissues. These parasites are known to
many pisciculturists and they have been called “anchor worms” because of their shape
and method of attachment.
THATCHER & PAREDES (1985 a) described the first lernaeid found in the Amazon
region. The description was based on specimens from Iquitos, Peru, but the same species
has been found to occur in the vicinity of Manaus, Brazil, as well. These parasites are
causing problems in local pisciculture ponds since they infect an important food fish,
namely; Colossoma macropomum. The parasite was named Perulernaea gamitanae to indicate the
country of origin and the fish of origin, since the host is know as “gamitana” in that part
of the Amazon. These parasites embed their heads in the nasal fossae, under the tongue,
in the walls of the esophagus or on the inner walls of the opercula. They suck blood to the
extent that they may weaken or kill fish in captivity.
In recent years, several additional species of Lernaeidae have been found in South
America. At least seven native species occur on this continent including another Perulernaea.
Female lernaeids, prior to metamorphosis, and adult males are found free-living in
the zooplankton. They are similar in size (about 1 mm long) and appearance to ergasiloid
copepods. They can be distinguished at once, however, by their mouthparts (Fig. 8-50).
The main mouthparts of lernaeids are a pair of bifid second maxillae and a pair of 5-
clawed maxillipeds (Fig. 8-40 C).
Therodamasidae
THATCHER (1986) described a new copepod that he called Amazonicopeus elongatus and
proposed a new family to include it. AMADO & ROCHA (1996) pointed out the
similarities between this genus and Therodamas KR ǾYER, 1863, and transferred elongatus
to that genus. TRIPATHI (1960) proposed the family Therodamasidae to include three
genera, Therodamas, Mugilicola and Paeonodes. Although all three have elongate neck-like
regions, that of Therodamas is formed from the head and is between the antennae and
mouthparts. The “necks” of the other two genera are posterior to the mouth and
therefor formed from thoracic elements. This was pointed out by BOXSHALL (1986)
who considered that all three genera should be placed in Ergasilidae. In spite of this
suggestion, PIASECKI et al. (1991) recognized Therodamasidae as valid and placed a
new species of Mugilicola within it.
Therodamas (= Amazonicopeus) differs from all other parasitic copepods in the

following ways: 1) There is a distinct head capsule, but no cephalothorax; 2) The
elongate “neck” is formed from head segments and does not contain any thoracic
elements; 3) The mouth is located at a considerable distance from the head capsule, is
on the thoraco-abdomen and is external to the host tissue; 4) The mouthparts are
similar to those of ergasiloids but in addition there is a protrusible mouth tube, which
possibly represents an extension of the esophagus and 5) The last two abdomimal
segments are divided longitudinally to accompany the uropod on either side. (Fig. 8-39
A-O). For these reasons, we propose that Therodamasidae should be recognized as a
valid family containing only the species of the genus Therodamas. T. elongatus was
described from a fish of the family Sciaenidae but since then, it has been found that this
genus is widely distributed in Brazil. Specimens have been seen from Rondônia,
Amazonas, Pará and Rio de Janeiro States. Most of the material seen was from
siluriform hosts of the families Doradidae and Loricariidae. Pre-metamorphic females
have also been found in plankton samples from near Belém, Pará State (Fig. 8-51).
The post-metamorphic female of T. elongatus inserts its head end through the soft
tissues of the gill arch and clings to the bony arch itself with strong 3-segmented antennae
(Fig. 8-39 B). The thoraco-abdomen, which bears the mouth, hangs outside in close
association with a gill filament. Apparently, these parasites feed by externally digesting the
gill filaments and ingesting the liquid diet through the mouth tube (Fig. 8-39 C, F & O).
Mature female copepods characteristically have a pair of egg sacs projecting from
their genital segments or posterior body regions. These sacs are usually held in place
until the eggs hatch. A free-swimming larva, called a nauplius (Fig. 8-2), breaks out of
each egg. These larvae, by feeding, growing and molting, go through several naupliar
stages and then transform themselves into copepodids. The copepodid stages
resemble the adults which they eventually become. The only complete cycle of an
Amazonian copepod to have been worked out is the study of VARELLA (1985). She
found that in Ergasilus bryconis THATCHER , 1981, there are three naupliar stages and
five copepodid stages. From the third copepodid stage on, males could be distin-
guished from females by the presence of primordial maxillipeds and smaller
antennae. The morphology of the copepodid stages of this species is shown in Figs.
8-3 to 8-5. The experimentally obtained adult male is shown in Fig. 8-7 A-C.
Nothing is known about the life-cycle of the other Amazonian parasitic copepods,
but they are probably similar and vary only in detail. It would seem that the complete cycle
for ergasiloids in the warm waters of the Amazon requires 10 to 20 days. This is such a
short time that populations can build up rapidly on captive fish. In all cases, transmission
is by the direct attack of the female copepod on the fish host.
III. Pathology
Species of Ergasilus are known pathogens and are one of the plagues of pisciculture
throughout the world. In the Brazilian Amazon, the comparative histopathology of
Ergasilus, Brasergasilus and Acusicola was studied by THATCHER & BOEGER (1983 a). They
found that all of these ergasilids produce epithelial hyperplasia, metaplasia and lamellar
fusion in the gill filaments of their hosts (Fig. 8-61). The degree of host response was
found to vary among species of Ergasilus depending on the depth of filament penetration
by the antennae. In the case of Acusicola, which latches one antenna to the other and
completely encircles the gill filament, a tourniquet effect was noted. Species of the genus
tend to reduce blood circulation within the filament and cause necrosis at the tip (Fig. 8-
61). Ergasiloids can and do cause fish deaths by reducing gill efficiency and by opening
the way for secondary bacterial invaders.
The post-metamorphic females of Perulernaea gamitanae can probably produce a
primary anemia in fish. The constant loss of blood no doubt affects metabolism, growth
and resistance to other pathogens, in captive fish.
The pathogenicity of Therodamas elongatus was reported by THATCHER (1986). As the
copepods tunnel through the soft tissues of the gill arch, the “neck” region lengthens so
that the hind body remains outside and adjacent to the gill filaments. The penetration of
the head provokes an inflammatory reaction on the part of the host which extends an
epithelial mass out over the entire “neck”. There is gradual fibrotic encapsulation of the
head and “neck” followed by calcification. This process sometimes destroys the head of
the parasite, but the hindbody may remain alive and reproductive for a while (Fig. 8-39 M-
N). Feeding activity by these copepods may remove some gill filament epithelium, but this
appears to be well tolerated by fish hosts.
Although insecticides such as Neguvon or Malathion are recommended by some as a

treatment for fish infected with Ergasilus, these chemicals have not been shown to be very
effective. Not only do these products not work well against ergasiloids but they are highly
toxic. In reality, no really good remedy for ergasilid infestations is known. Prevention
includes avoiding the introduction of infested fish into clean pounds. If culture fish
become too heavily infested, it is best to sacrifice them, drain and refill the pond or
aquarium and start over.
Ergasiloids are best collected by cutting off the gill filaments to which they are clinging
and dropping them into AFA fixative. After fixing the gill filament for several hours, it will
be possible to remove the ergasilids with dissecting needles, being careful not to break off
the antennae. Temporary study slides can be made by clearing specimens in phenol, after
partial dehydration in 95 % alcohol. Permanent mounts can be made afterwards by means

of the phenol-balsam method which is explained in Chapter 6.
Special preparations are necessary to study the detailed morphology of copepod
legs and antennae. Crush preparations can be made with the specimen under a
coverglass in either phenol or balsam. The excess phenol or balsam is removed with a
piece of filter paper, and the coverglass is pressed down firmly at a slight angle. If
done properly, this method will offer a medial view of all the legs. If crush
preparations do not work with the material at hand, dissections may be required.
These can also be performed on specimens in either phenol or balsam. It is best to
work under a dissecting microscope with a high enough magnification to show the
basipods clearly. Using glass needles, the legs and antennae are simply pushed off
with one needle while holding the body securely with the other. Dissecting small
specimens requires some skill, practice and patience. For maximum stability, the arms
should be resting solidly on the table top and the hands should be steadied on the
sides of the dissecting microscope.
Key to Amazonian freshwater ergasiloid females
I. Lateral retrostylets present on cephalothorax; antennal claw single or double. Rostral spine sometimes present; found in the nasal
fossae of the host ........................................................................................................................................................................................ Vaigamidae
A. Antennal claw single; rostral spine present, sharply pointed ....................................................................................... Vaigamus (Fig. 8-29)
B. Antennal claw double; rostral spine absent, or when present not sharply pointed.
1. Rostral spine absent.
a. Third antennal segment with prominent fixed spine; second antennal segment with numerous spinules ................................
.................................................................................................................................................................... Gamispinus (Figs. 8-32 & 8-35)
b. Third antennal segment without spine; second segment without spinules ................... Gamidactylus (Figs. 8-31, 8-34 & 8-56)
2. Flattened rostral spine with rounded tip present ................................................................................ Gamispatulus (Figs. 8-33 & 8-36)
II. Lateral retrostylets absent; antennal claw single; rostral spine absent; from gill filaments or gill rakers (rarely from nasal fossae) .......
.......................................................................................................................................................................................................................... Ergasilidae

A. Antennae of 3 segments (including claw); only 3 pairs of swimming legs present .............. Brasergasilus (Figs. 8-24 to 8-28 & 8-55)
B. Antennae of 4 segments; four pairs of swimming legs present.
1. Antennae of latching type: third segment provided with groove for reception of opposite claw ............................. Acusicolinae
a. First endopods of 2 segments similar in length to first exopod.
(1) Claw relatively large; points exposed when antennae are latched; first endopod provided with setae ................................
.................................................................................................................................... Acusicola (Figs. 8-12 to 8-14 & 8-18 to 8-19)
(2) Claw small; points covered by sheath-like extension of second antennal segment when antennae are latched; first
endopod without setae .................................................................................................................. Amplexibranchus (Fig. 8-37 A-I)
b. First endopod of 2 segments but is elongated and more than twice the length of the first exopod ........................................
........................................................................................................................................................................... Miracetyma (Fig. 8-53 A-C)
2. Antennae of clampling type; no groove present but antennal segments 3 & 4 form a clamp with segment 3; found in gill rakers
........................................................................................................................................................................... Prehendorastrus (Fig. 8-38 B-C)
3. Antennae of grasping type; third segment without groove.
Antennule of 6 segments ............................................................................................................................................................... Ergasilinae
333
a. Third antennal segment elongate ................................................................ Ergasilus (Figs. 8-1 to 8-11, 8-15, 8-16, 8-20 to 8-23)
334
b. Third antennal segment short
(1) Leg 4 biramous ............................................................................................................................................... Pindapixara (Fig. 8-54)

(2) Leg 4 reduced to one or two setae ........................................................................................................... Rhinergasilus (Fig. 8-30)
Key to the South American genera and species of postmetamorphic female Lernaeidae
I. Head with four anchors.

A. Anchors large, frequently branched; genital pore near posterior extremity; fourth legs on hindbody ...................................................
.................................................................................................................................................................................... Lernaea cyprinacea. (Fig. 8-47)
B. Anchors small, rounded, not branched; genital pore in equatorial region of hindbody; fourth legs on “neck.”
1. Petal-like anterior extensions present on head ................................................................................... Minilernaea floricapitella (Fig.8-46)
2. Petal-like anterior extensions on head absent ....................................................................................... Amazolernaea sannerae (Fig.8-44)
II. Head with two anchors.
A. Anchors short and blunt; only fourth pair of legs on expanded hindbody; mouth posterior to anchors.
1. Glandular area on “neck” extensive; large forms (18-22 mm) found only on Colossoma macropomum ...............................................
.......................................................................................................................................................................... Perulernaea gamitanae (Fig.8-42)
2. Glandular area on “neck” of limited extent; small forms (8-12 mm); found only on Piaractus brachypomus ....................................
........................................................................................................................................................................ Perulernaea irapitingae (Fig. 8-43)
3. “Neck” without glands; collar-like expansion present on hindbody anteriorly ................................. Bedsylernaea collaris (Fig. 8-45)
B. Anchors long and slender; legs 3 and 4 on hindbody; mouth anterior to anchors.
1. Head bilobed ........................................................................................................................................ Taurocherus salminisii (Fig. 8-40 A-C)
2. Head simple, not bilobed ................................................................................................................... Taurocherus taraganophilus (Fig. 8-41)
C. Head without anchors ............................................................................................................................................................... Lamproglena nyasae
Clave para hembras Ergasiloides dulceacuícolas Amazonicas
I. Retroestiletes laterales presentes sobre el cefalotórax; garra antenal simple o doble; espina rostral a veces presente; encontradas en las
fosas nasales del huésped .......................................................................................................................................................................... Vaigamidae
A. Garra antenal única; espina rostral presente, punta afilada ............................................................................ Vaigamus (Figs. 8-29 a 8-30)
B. Garra antenal doble; espina rostral ausente, o cuando presente sin punta afilada.
1. Espina rostral ausente.
a. Tercer segmento de la antena con espina fija prominente; segundo segmento de la antena con numerosas espínulas .........
.................................................................................................................................................................... Gamispinus (Figs. 8-32 & 8-35)
b. Tercer segmento de la antena sin espina; segundo segmento sin espínulas ................. Gamidactylus (Figs. 8-31, 8-34 & 8-56)
2. Espina rostral con punta arredondada .................................................................................................. Gamispatulus (Figs. 8-33 & 8-36)
II. Retroestiletes laterales ausentes; garra antenal única; espina rostral ausente; encontrados en filamentos o rastrillos de las branquias
(raramente en fosas nasales) ....................................................................................................................................................................... Ergasilidae
A. Antena con 3 segmentos (incluyendo garra); solo 3 pares de piernas nadadoras presentes .........Brasergasilus (Figs. 8-24 a 8-28 & 8-55)
B. Antena con 4 segmentos; 4 pares de piernas nadadoras presentes.

1. Antena tipo aldaba: tercer segmento provisto de ranura para recepción de la garra opuesta ............................................................
.......................................................................................................................................................................................................... Acusicolinae
a. Primer endopodito con 2 segmentos similares en largo al primer exopodito.
(1) Garra relativamente grande; puntas expuestas cuando las antena se unen; primer endopodito provisto de setas ...........
....................................................................................................................................... Acusicola (Figs. 8-12 a 8-14 & 8-18 a 8-19)
(2) Garra pequeña; cuando la antena está cerrada puntas cubiertas por una extensión en forma de escudo del segundo
segmento; primer endopodito con setas .................................................................................... Amplexibranchus (Fig. 8-37 A-I)
b. Primer endopodito con dos segmentos pero es alargado y tiene más de dos veces el largo del primer exopodito ...............
........................................................................................................................................................................... Miracetyma (Fig. 8-53 A-C)
2. Antena tipo abrazadera; sin ranura pero los segmentos de las antenas 3 y 4 forman una abrazadera con el segmento 3;
encontrado en los rastrillos branquiales .................................................................................................... Prehendorastrus (Fig. 8-38 B-C)
a. Tercer segmento de la antena alargado ...................... Ergasilus (Figs. 8-1 a 8-11, 8-15, 8-16, 8-20 a 8-23, 8-46, 8-47 & 8-49)
b. Tercer segmento de la antena corto.
335
(1) Pierna 4 birameada ........................................................................................................................................ Pindapixara (Fig. 8-54)
336
(2) Pierna 4 reducida a una o dos setas ......................................................................................................... Rhinergasilus (Fig. 8-30)

Claves para los géneros y especies de Lernaeidae hembras post-metamórficas de América del Sur
I. Cabeza con cuatro áncoras.

A. Ancoras grandes, frecuentemente ramificadas; poro genital cerca de la extremidad posterior; el cuarto par de piernas localizado en
la parte posterior del cuerpo ............................................................................................................................... Lernaea cyprinacea. (Fig. 8-47)
B. Ancoras pequeñas, redondeadas, sin ramificaciones; poro genital en la región ecuatorial de la parte posterior del cuerpo; 4 pares
de piernas en el “cuello”
1. Extensiones anteriores presentes en formas de pétalos en la cabeza ............................................ Minilernaea floricapitella (Fig.8-46)
2. Extensiones anteriores en forma de pétalos ausentes en la cabeza ................................................. Amazolernaea sannerae (Fig.8-44)
II. Cabeza con 2 áncoras.
A. Ancoras cortas y redondeadas; solo el cuarto par de piernas está en la parte posterior del cuerpo; boca posterior a las áncoras
1. Area glandular del “cuello” extensa; formas grandes (18-22 mm); encontradas solamente en Colossoma macropomum ..................
.......................................................................................................................................................................... Perulernaea gamitanae (Fig.8-42)
2. Area glandular del “cuello” de extensión limitada; formas pequeñas (8-12 mm); encontradas solamente en Piaractus brachypomus
...................................................................................................................................................................... Perulernaea pirapitingae (Fig. 8-43)
3. “Cuello” sin glándulas; expansión tipo collar presente en la parte proximal de la parte posterior del cuerpo ..............................
............................................................................................................................................................................ Bedsylernaea collaris (Fig. 8-45)
B. Ancoras largas y finas; piernas 3 y 4 en la parte posterior del cuerpo; boca anterior a las áncoras.
1. Cabeza bilobada .................................................................................................................................. Taurocherus salminisii (Fig. 8-40 A-C)
2. Cabeza simple, no bilobada ............................................................................................................... Taurocherus taraganophilus (Fig. 8-41)
C. Cabeza sin áncoras ..................................................................................................................................................................... Lamproglena nyasae
VII. Checklist of Copepoda from Amazonian freshwater fishes
Species measurements are all in micrometers (µm) except where otherwise indicated.
Ergasilidae NORDMANN, 1832

Copepoda, Poecilostomatoida. Body cyclopoid, tapered posteriorly. Head
fused with first 1 or 2 thoracic segments. Antennule 5 or 6-segmented,
provided with simple setae; antenna 3 or 4 segmented, prehensile, with
clutching, piercing, latching or clamping claw. Mouthparts: mandible 2-
segmented, with simple palp on basal segment; maxillule vestigial, usually
provided with 1 or 2 setae; maxilla large, 2-segmented; maxillipeds 3 or 4-
segmented in male, absent in female. Thorax: 3 – 5 free thoracic segments
present; genital segment enlarged. Abdomen: 3-segmented in female; 4-
segmented in male; uropods simple, with terminal setae. Legs: 3 or 4 pairs
of biramous swimming legs present; vestigial 5th leg of one or 2 setae often
present; vestigial 6th leg of 1 seta sometimes present in male. Male: free-
living; smaller than female. Female: parasitic on gill filaments, gill rakers or
in nares of fish; egg sac usually elongate or elliptical, multiseriate.
Abergasilinae THATCHER & BOEGER, 1983
Ergasilidae. Female: antennae 3-segmented; 3 pairs of swimming legs
present, vestigial legs lacking. Male: unknown. Parasites of gill filaments
and gill rakers of fish.
Brasergasilus THATCHER & BOEGER, 1983
Abergasilinae. Antennae smooth, without bosses.
B. anodus THATCHER & BOEGER, 1983: Anodus elongatus: Brazil (Amazonia).
(Figs. 8-25 A-C & 8-26). Body = 320-370 x 118-163 (female; male
unknown); Eye smalt blue; Body pigmentation sparse, campanula blue;
Claw = 7-88 x 10-15; Caudal setae = 95-163 (maximum),
B. guaporensis MALTA, 1995: Leporinus fasciatus: Guaporé River, Rondônia State,
Brazil. (Fig. 8-55 A-B). Body = 301-384 x 112-150; Cephalothorax = 150-
210 x 112-150.
B. jaraquensis THATCHER & BOEGER, 1983: Semaprochilodus insignis: Brazil (Ama-
zonia). (Figs. 8-24 A-D & 8-28). Body = 380-410 x 120-170 (female; male
unknown); Eye = rose, surrounded by spectrum blue; body pigmentation
spectrum blue ventrally; Claw = 80-95 x 13-18; Caudal setae = 83-143
(maximum).
B. oranus THATCHER & BOEGER, 1985: Anodus elongatus: Brazil (Amazonia). (Fig.
8-27).
Acusicolinae THATCHER, 1984
Ergasilidae. Female with 4 pairs of swimming legs and vestigial 5th pair.
Antennules 5-segmented; antennae 4-segmented, with groove in third
segment for reception of opposite claw when antennae are latched. Male
unknown. Parasites of gill filaments of fish.
Acusicola CRESSEY, 1970, THATCHER, 1984

Acusicolinae. Claws of antennae only partially covered by cuticular flap
from segment 2 when antennae are latched. Legs 1-4 biramous; first
endopod 2-segmented; fourth exopod 2-segmented; all other rami 3-
segmented; leg 5 reduced to 1 or 2 setae; leg 6 absent in female. Male
unknown. From gill filaments of fish.
A. cunula CRESSEY, 1970: Pseudotylosurus angusticeps: Brazil (Amazonia). (Fig. 8-
19). Body = 652 x 290 (female; male unknown).
A. lycengraulidis THATCHER & BOEGER, 1985: Lycengraulis grossidens: Brazil (Ama-
zonia). (Fig. 8-12 A-C). Body = 800-950 x 275-400 (female; male
unknown); Eye = smalt blue; Body pigment campanula blue, scattered
from eye to genital segment; Claw = 43-53 x 13-15; Caudal setae = 150-
200 (maximum).
A. pellonidis THATCHER & BOEGER, 1983: Pellona castelnaeana: Brazil (Amazonia).
(Fig. 8-13 A-D). Body = 1,025-1,275 x 300-425 (female; male unknown);
Eye = cobalt blue; Body pigment cobalt blue and widely scattered; Claw =
50-75 x 15-20; Caudal setae = 140-190 (maximum).
A. tenax (ROBERTS, 1965): Pomoxis annularis and Strongylura sp. Texas &
Guatemala. (Fig. 8-18). Body = 800 x 225 (female; male unknown); Caudal
setae = 175.
A. tucunarense THATCHER, 1984: Cichla ocellaris: Brazil (Amazonia). (Fig. 8-14 A-
D). Body = 720-970 x 250-350 (female; male unknown); Eye = smalt blue;
cerulean blue pigment widely scattered in body; Claw = 64-72 x 15-20;
Caudal setae = 130-175.
Amplexibranchius THATCHER & PAREDES, 1985
Acusicolinae. Female: cephalothorax inflated, or not. Abdomen 3-segment-
ed. Antennule 5-segmented. Antenna 4-segmented; 4th segment (claw)
reduced; first and third segments shorts; segment 2 long, with anterior
cuticular extension which partially encloses segment 3. Maxillipeds absent.
Legs 1-4 biramous; all setae pectinate; first endopod subcylindrical, without
setae (spinules or setules sometimes present); terminal segments of endo-
pods 2-4 sharply tapering, setae small; 4th exopod 2-segmented. Leg 5
represented by 1 or 2 setae. Leg 6 absent. Male unknown; presumably free-
living. Female on gill filaments of freshwater fish.
A. bryconis THATCHER & PAREDES, 1985: Brycon cephalus: Peru & Brazil (Amazo-
nia). (Fig. 8-37 A-I). Body = 825-1,075 x 250-340 (female; male unknown);
Eye = smalt blue; Body pigmentation widely scattered, spectrum blue;
Antenna: segment 2 = 310-440; Claw = 12-18.
Ergasilinae THATCHER & BOEGER, 1983
Ergasilidae. Female with 4 pairs of swimming legs and vestigial 5th pair.
Antennule 5 o 6 segmented; antenna 4-segmented; claw large (for clutching
or piercing). Male: maxillipeds present; abdomen 4-segmented. From gill
filaments, gill rakers or nares of fish.
Ergasilus NORDMANN, 1832

Ergasilinae. Antennule 6-segmented; antenna 4-segmented, with clutching
or piercing claw. Maxilipeds present in male, absent in female. Abdomen 3-
segmented in female, 4-segmented in male. Gill filaments, gill rakers or
nares of fishes.
E. argulus CRESSEY, 1970: Strongylura fluviatilis and S. scapularis: Colombia
(Department of Valle). (Fig. 8-23). Body = 660 x 413 (Female; male
unknown); Caudal setae = 235 (max.).
E. bryconis THATCHER, 1981: Brycon erythropterus: Brazil (Amazonia). (Figs. 8-2
to 8-7). Body = 700-860 x 290-370 (female); 490-610 x 161-218 (male);
Eye = cobalt blue; Body pigment spectrum violet, widely distributed; Claw
= 83-94 x 18-21 (female); 38-42 x 8-10 (male); Caudal setae = 240-340
(female); 180-280 (male).
E. callophysus THATCHER & BOEGER, 1984: Callophysus macropterus: Brazil (Ama-
zonia). (Fig. 8-8 A-D). Body = 800-900 x 200-280 (female; male unknown);
Eye = cobalt blue; Body pigmentation smalt blue, distributed from
antennal bases to third abdominal segment; Claw = 90-110 x 20; Caudal
setae = 230-340.
E. coatiarus ARAUJO & VARELLA, 1998: Cichla monoculus: Amazonas State, Brazil.
Body = 639-717 x 304-334 (female; male unknown); Similar to E. hydrolycus
but smaller.
E. colomesus THATCHER & BOEGER, 1983: Colomesus asellus: Brazil (Amazonia).
(Figs. 8-9 A-E & 8-49 to 8-53). Body = 540-700 x 190-240 (female; male
unknown); Eye = smalt blue; Body pigmentation in longitudinal bands
from eye to 4th thoracic segment; smalt blue anteriorly, shading to
campanula posteriorly; Claw = 53-63 x 15-20; Caudal setae = 128-160.
E. euripedesi MONTÚ, 1980: from larvae of Brevoortia pectinata: Micropogonias
furnieri and Lycengraulis grossidens: Brazil (Rio Grande do Sul State). (Fig. 8-
22). Body = 785 x 292 (female); Claw = elongate, barbed.
E. holobryconis MALTA & VARELLA, 1986: Holobrycon pesu: Brazil (Rondônia
State). (Fig. 8-15). Body = 525-624 x 240-320 (female; male unknown);
Claw = 55-61 x 16-17.
E. hydrolycus THATCHER, BOEGER & ROBERTSON, 1984: Hydrolycus scomberoides:
Brazil (Amazonia). (Fig. 8-1 A-D). Body = 725-875 x 260-310 (female;
male unknown); Eye = cobalt blue; Body with spectrum blue band at level
of mouth; Claw = 97-105 x 50-60; Caudal setae = 200-270.
E. hypophthalmi BOEGER, MARTINS & THATCHER, 1990: Gill rakers of Hypoph-
thalmus edentatus and H. fimbriatus: Amazonas State, Brazil. Body similar to
E. hydrolycus but second thoracic segment is fused to cephalothorax.
E. iheringi TIDD, 1942: Hoplias malabaricus: Brazil. (Fig. 8-16). Body = 799-918
x 238-340 (female; male unknown); Second antennae small, not projecting
beyond margins of cephalotorax.
E. jaraquensis THATCHER & ROBERTSON, 1982: Semaprochilodus insignis: Brazil
(Amazonia). (Fig. 8-10 A-D). Body = 680-750 x 235-275 (female; male
unknown); Eye = cobalt blue; Body pigmentation extensive, cobalt blue;

Claw = 55-69; Caudal setae = 280-360.
E. leporinidis THATCHER, 1981: Leporinus fasciatus: Brazil (Amazonia). (Fig. 8-20).
Body = 530-710 x 270-330 (female; male unknown); Eye = cobalt blue;
Body pigmentation sparse, spectrum violet; Claw = 92-97 x 16-23; Caudal
setae = 270-310.
E. pitalicus THATCHER, 1984: Cichlasoma sp.: Colombia. (Figs. 8-11 & 8-21).
Body = 814-904 x 345-497 (female; male unknown); Eye = spectrum
blue; Body pigmentation extensive, spectrum violet; Claw = 140-154 x
25-28.
E. thatcheri ENGERS, BOEGER & BRANDON, 2000: Rhamdia quelen: Rio Grande do
Sul State, Brazil. Body with blue pigment from cephalothorax to genital
segment. General morphology similar to that of E. callophysus but has row
of spinules on second antennal segment.
E. urupaensis MALTA, 1993: Prochilodus nigricans: Urupá River, Rondônia State,
Brazil. Body = 833-958 (female, male unknown). This is the largest
Neotropical species of the genus.
Miracetyma MALTA, 1993.
Acusicolinae. Female: cephalothorax inflated, or not. Abdomen 3-seg-
mented. Antennule 5-segmented; antenna 3-segmented (not counting claw,
which is very small and grooved; third segment with one or two grooves;
segment two long; first second and third segments with one or two
cuticular expansions. Legs I-IV biramous; all setae pectinate; first endopod
and fourth exopod 2-segmented, all others 3-segmented. First endopod
extremely modified; first segment large, stout and elongate; second
segment subcylindrical, slender and elongate. Leg V represented by one or
two simple setae. Parasitic on the gill filaments of freshwater fishes. Male:
unknown but presumably free-living. Type species: Miracetyma etimaruya.
M. etimaruya MALTA, 1993: Curimata cyprinoides, Potamorhina latior and Psectro-
gaster essequibensis: Rondônia State, Brazil. (Fig. 8-53 A-C). Body = 802-936
x 286-357; Egg sac = 318-535.
M. kawa MALTA, 1993: Rhaphiodon vulpinus: Rondônia State, Brazil. Body =
845-1432 x 357-546; Egg sac = 330-989.
M. piraya MALTA, 1994: Pygocentrus nattereri: Marmoré River, Rondônia State,
Brazil. Body = 978- 1070 x 352-473; Egg sac = 269-474.
Pindapixara MALTA, 1995.
Ergasilinae. Female: Body small. Antennule with 6 segments; antenna with
4 segments (including claw); claw larger that the other segments; third
segment greatly reduced. Legs I-IV biramous first endopod and both rami
of leg IV 2-segmented; other rami 3-segmented. Egg sac uniseriate. Gill
parasites of freshwater fishes. Type species: Pindapixara tarira.
P. taira MALTA, 1995: Hoplias malabaricus: Guaporé River, Rondônia State,
Brazil. (Fig. 8-54). Body = 382-577 (female, male unknown); Caudal
filaments = 17-187.
Prehendorastrus BOEGER & THATCHER, 1990

Ergasilidae. Antennule 5-segmented. Antenna 4-segmented, modified to
individually grasp the gill rakers and latch; second segment with 1 or 2
prominent teeth. First maxilla with 3 setae. Five pairs of biramous legs;
first endopod and fourth exopod 2-segmented; all other rami 3-segmen-
ted; leg 5 reduced to 2 setules. Parasites of gill rakers of fish. Type species:
Prehendorastrus bidentatus BOEGER & THATCHER, 1990.
P. bidentatus BOEGER & THATCHER, 1990; Hypophthalmus edentatus and H.
fimbriatus: Rio Negro, near Manaus, Amazonas State, Brazil. (Fig. 8-38). B
= 931-1,112 x 339-423; Cephalothorax = 494-780 x 339-423.
P. monodontus BOEGER & THATCHER, 1990; Hypophthalmus edentatus and H.
fimbriatus: Rio Negro, near Manaus, Amazonas State, Brazil. B = 1,110-
1,421 x 464-542; Cephalothorax = 622-826 x 462-542.
Rhinergasilus BOEGER & THATCHER, 1988
Ergasilinae. Female: antennule 6-segmented, antenna 4-segmented. Tho-
racic segments V & VI reduced; legs 4 & 5 reduced to single setae.
Parasites in nasal fossae of fish. Male unknown.
R. piranhus BOEGER & THATCHER, 1988: Serrasalmus nattereri: Brazil (Amazonia).
(Fig. 8-17 & 8-30 A-B). Body = 237-282 x 95-102 (female; male unknown);
Claw = 37-52 x 5-7.
Vaigamidae THATCHER & ROBERTSON, 1984
Poecilostomatoida. Female: body cyclopoid; cephalosome completely or
incompletely fused with second thoracic segment; latter bearing moveable
retrostylet on either side; rostrum with or without ventral spine. Thoracic
segments 3-7 free; segments 2-5 bearing biramous legs 1- 4; segment 6
bearing leg 5, composed of 1 or 2 setae; leg 6 absent. Genital segment well
developed; abdomen 3-segmented; uropods cylindrical, with terminal setae.
Antennule 6-segmented, with simple setae; antenna prehensile, 4-segmented.
Mouthparts: mandible 2 or 3-segmented, palp 1-segmented; maxillule
vestigial; maxilla 2-segmented. Egg sac uniseriate, with few eggs. In nasal
fossae of freshwater fish. Male: similar to female but smaller. Cephalosome
lacking retrostylets and rostral spine. Antennae prehensile, but smaller than
those of female. Maxillipeds present, 3 or 4-segmented. Male free-living.
Vaigamus THATCHER & ROBERTSON, 1984
Vaigamidae. Female: retrostylets prominent; rostrum with single tapered
spine ventrally. Antenna ergasiloid, with simple claw. Legs: first endopod
2-segmented; fourth endopod 2 or 3-segmented; fourth exopod 1 or 2-
segmented; all other rami 3-segmented. Male: similar to female, but
smaller. Maxilliped 4-segmented. Leg 5 of 2 setae; leg 6 of 1 seta.
Abdomen 4-segmented.
V. retrobarbatus THATCHER & ROBERTSON, 1984: from plankton, host un-
known: Brazil (Amazonia). (Fig. 8-29 A-E). Body = 573-658 x 168-216
(female); 498-545 x 161-185 (male); Eye = spectrum blue; Body with
single band of spectrum blue in thoracic segment 2; Retrostylet = 108-
110; Rostral spine = 58-72; Claw = 60-74 x 10-14 (female); 29-36 x 6-8
(male); Caudal setae = 291-376.
V. spinicephalus THATCHER & ROBERTSON, 1984: from plankton, host un-
known. Brazil (Amazonia). Body = 461-498 x 223-228 (female); 390-470
x 135-165 (male); Eye = not observed; Body pigmentation limited to
small areas in postero-lateral portion of thoracic segment 1 and in
thoracic segments 3 & 4, also in basipods of legs 1-2; color = smalt blue;
Retrostylet = 91-108; Rostral spine with subbasal swelling = 35-37; Claw
= 35-39 x 8-10 (female); 25-30 x 7-8 (male); Caudal setae = 263-291
(female); 240-280 (male).
Gamidactylus THATCHER & BOEGER, 1984
Vaigamidae. Retrostylets prominent. Rostrum unarmed. Antenna: segment 3
with elongate claw-like moveable spine; segment 4 claw-like. Male: unknown.
Female in nasal fossae of freshwater fish.
G. jaraquensis THATCHER & BOEGER, 1984: Semaprochilodus insignis: Brazil (Ama-
zonia). (Figs. 8-31 A-E & 8-34). Body = 410-470 x 130-160 (female; male
unknown); Eye = spectrum blue; same color widely scattered in body;
Retrostylet = 125-140; Claw = 42-45 x 5-7; Caudal setae = 155-187.
G. bryconis VARELLA, 1995: Brycon pellegrini and B. melanopterus: Rondônia
State, Brazil. (Fig. 8-56 A-B). Body = 464-576 x 162-220; Caudal setae =
160-200.
Gamispatulus THATCHER & BOEGER, 1984
Vaigamidae. Retrostylets short, with medial spatulate processes. Rostrum
with tapered spatulate extension on ventral surface. Antenna: segment 3
with curved moveable spine distally; segment 4 claw-like. Male unknown.
Female in nasal fossae of freshwater fish.
G. schizodontis T HATCHER & B OEGER , 1984: Schizodon fasciatus: Brazil
(Amazonia). (Figs. 8-33 A-D & 8-36). Body = 380-470 x 140-170
(female; male unknown); Eye = spectrum blue; smalt blue pigment
scattered in body; Retrostylet = 90-103; Claw = 23-30 x 5-8; Caudal
setae = 195-205.
Gamispinus THATCHER & BOEGER, 1984
Vaigamidae. Retrostylets present; rostrum unarmed. Antenna 4-segment-
ed; segment 2 spinous on side bearing sensillum; segment 3 with fixed
spines and moveable claw-like spine distally; segment 4 claw-like.
G. diabolicus THATCHER & BOEGER, 1984: Ageneiosus brevifilis: Brazil (Amazonia).
(Figs. 8-32 A-E & 8-35). Body = 570-710 x 160-220 (female; male
unknown); Eye = spectrum blue; same pigment scattered in small spots in
cephalosome and thoracic segments 2-4; Retrostylet = 97-107; Claw = 20-
25; Caudal setae = 152-187.
Lernaeidae COBBOLD, 1879
Cyclopoida. Adult males and pre-metamorphosed adult females small,
cyclopoid; cephalosome, eye, free thorax, genital segment and uropods
present. Antennules sensory; antennae prehensile. Mouthparts: maxillule
vestigial or absent; maxillae usually bifid; maxillipeds strongly developed,

with terminal claws. Legs: 4 pairs of biramous legs present; rami usually 3-
segmented; provided with pilose setae and spines; vestigial legs 5 and 6
frequently present. Metamorphosed female: large, elongate, of varied
morphology; eyes usually absent; head anchors often present; body
segmentation reduced or absent. Mouthparts, antennae and legs usually
same form and size as in pre-metamorphosed female. Uropods present or
absent. Males and pre-metamorphosed females free-living. Metamor-
phosed females parasitic on fish; head usually inserted within fish tissues.
Type genus and species: Lernaea cyprinacea Linnaeus, 1758 (Fig. 8-47).
Amazolernaea THATCHER & WILLIAMS, 1998.
Lernaeidae. Body divisible into head, neck and hindbody. Head bearing
four small petal-like lobes, mouthparts antennae and first pair of thoraco-
pods. Mouthparts: second maxillae bifurcate: maxillipeds with several
terminal spines. Neck long, slender cylindrical. One pair of thoracopods
present on head and three on neck: first neck pair close to head pair.
Hindbody subcylindrical, formed from fused genital segment and abdo-
men; genital pore near midpoint, pregenital prominence small. Egg sacs
multiseriate. Male and premetamorphic female unknown. Type species:
Amazolernaea sannerae THATCHER & WILLIAMS, 1998.
A. sannerae THATCHER & WILLIAMS, 1998: Cichla monoculus and C. temensis: Paciva
River Venezuela and Jatapu River, Amazonas State, Brazil. (Fig. 8-44).
Body (metamorphosed female) = 8.5-10.8 mm long; Hindbody = 4.0-4.5 x
0.79-1.0 mm.
Bedsylernaea THATCHER & WILLIAMS, 1998.
Lernaeidae. Postmetamorphic female: body divisible into head, neck and
hindbody. Three pairs of thoracopods observed. Head bearing two large,
bulbous anchors, mouthparts and antennae. Mouthparts: second maxillae
bifurcate, maxillipeds with five terminal spines. Neck long, slender and
cylindrical, with two pairs of thoracopods. Hindbody subcylindrical, short,
stout, formed from fused genital segment and abdomen; with collar-like
anterolateral projections, one on either side of neck; one pair of thoraco-
pods (probably the fourth) between projections; genital pore anterior to
midpoint, pregenital prominence large. Egg sacs multiseriate. Male and
premetamorphic female unknown. Type species: Bedsylernaea collaris
THATCHER & WILLIAMS, 1998.
B. collaris THATCHER & WILLIAMS, 1998: Hoplias malabaricus: Trombetas River,
Pará State, Brazil. (Fig. 8-45). Body (metamorphosed female) = 13-16 mm
long; Hindbody = 5.0-6.5 x 1.6-2.0 mm.
Minilernaea THATCHER & HUERGO, 2005.
Lernaeidae Postmetamorphic female: Body divisible into head, neck and
hindbody. Head bearing 6 lobes anteriorly, 4 undivided, blunt anchors
posteriorly, antennae, mouthparts and first pair of thoracopods. Mouth-
parts: second maxillae bifurcate; maxilliped with five terminal, recurved
spines. Antennae: antennule with 4 segments; antenna with 2 segments.

Five pairs of thoracopods present: first pair on head; pairs 2-4 on neck;
pair 5 reduced to a simple papillus located on hindbody anterior to genital
pores. Neck subcylindrical, representing about one-third of total body
length. Genital pores equatorial on hindbody; pregenital prominence
absent. Egg sacs elongate, multiseriate. Parasites of freshwater fishes. Male
and premetamorphic female unknown. Type species: Minilernaea floricapitel-
la THATCHER & HUERGO, 2005.
M.floricapitella THATCHER & HUERGO, 2005: Astyanax spp. and Corydoras
ehrhardti: Iguaçu River, Santa Catarina State and Piraquara River, Paraná
State, Brazil. (Fig. 8-46 A-B). Body = 3.4-5.8 mm long; Hindbody = 0.62-
0.82 mm in diameter; Egg sacs = 1.4-1.9 mm long.
Taurocherus BRIAN, 1924.
Lernaeidae. Postmetamorphic female: Body divisible into head, anchors,
neck and hindbody. Head bearing antennae, mouthparts and first pair of
thoracopods. Anchors elongate, cylindrical, projecting laterally between
head and neck. Four pairs of thoracopods present, first on head, second
on neck at juncture of anchors, 3-4 on hindbody. Neck elongate, slender,
subcylindrical. Hindbody cylindrical, expanded anteriorly. Egg sacs short,
multiseriate. Parasites of freshwater fishes. Male and premetamorphic
female unknown. Type species: Taurocheros salminisii BRIAN, 1924.
T. salminisii BRIAN, 1924: Salminus brevidens: Argentina (Patagonia). (Fig. 8-40
A-C). Body = 12-17 mm; Head 2.0-2.2 mm; Neck = 7-9 mm; Egg sac =
2-4 mm.
T. tarangophilus PAGGI, 1976: Hoplias malabaricus: Paraná River, Argentina.
(Fig. 8-41). Body = 11.2-14.5 mm long; Neck = 6.4-8.6 mm; Hindbody =
3.3-5.4 mm long.
Perulernaea THATCHER & PAREDES, 1985
Lernaeidae. Female (pre-metamorphosed): small, cyclopoid, with prominent
hooked maxillae and maxillipeds. Female (post-metamorphosed): body
divisible into head, neck and hindbody. Head bearing: two large blunt lateral
lobes, two small ventral lobes, mouthparts and first pair of legs. Neck region
(thorax between legs 1 and 4) subcylindrical; comprising about two thirds of
body length. Hindbody formed of fusion of last leg-bearing thoracic
segment with genital segment and abdomen. Uropods present. First antenna
5 or 6 segmented, provided with simple setae; second antenna 2 or 3-
segmented, terminating in stout claw and several simple setae. Mouthparts:
maxilla bifurcate; maxilliped with several terminal claws. Four pairs of
biramous legs present; all rami 3-segmented and provided with setae and
spines. Egg sac multiseriate. From nasal fossae, mouth, tongue and gills of
freshwater fish. Male unknown, presumably free-living.
P. gamitanae THATCHER & PAREDES, 1985: Colossoma macropomum: Peru & Brazil
(Amazonia). (Figs. 8-38 A-J, 8-40 & 8-41). Body (metamorphosed female)
= 18-22 mm long; Head and anchors = 1.1-1.2 x 4.5 5.0 mm; Neck = 12-
15 x 0.32-0.93 mm; Hindbody = 5.0-5.5 x 1.2-2.0 mm; Egg sac = 2-4 x

0.43-0.63 mm.
P. pirapitingae THATCHER, 2000: Piaractus brachypomus: Meta River, Colombia
(Fig. 8-43). Body (metamorphosed female) = 8.9-11.4 mm long; Hindbody
= 4.0-5.0 x 0.91-1.1 mm.
Therodamasidae TRIPATHI, 1960 (emended THATCHER, 1986).
Poecilostomatoida. Female (postmetamorphic): Body composed of
head, neck and thoraco-abdomen. Head well developed, usually provid-
ed with two pigmented eyes and two pairs of antennae; head capsule
with or without lateral retrostylets or expansions; long, neck-like exten-
sion of head between antennae and mouth. Antennule 4 or 5-segmented
with simple setae; antenna 3-segmented; terminal segment a stout claw.
Mouth located in anterior part of thoraco-abdomen; mandible 2-
segmented, with palp; maxillule with medial spine; maxilla 2-segmented;
protrusible mouth tube present or absent. Thoraco-abdomen; thorax of
4 leg-bearing segments; abdomen fused to genital segment, except for 2
small free segments on either side preceding uropods; uropods with few
terminal segments. Legs: 4 pairs present on thoraco-abdomen; rami
usually 3-segmented. Egg sac multiseriate. Gill arch parasites of fishes.
Premetamorphic female ergasiloid in appearance. Male unknown but
presumably free-living.
Type and only genus: Therodamas KRǾYER, 1863.
T. elongatus (THATCHER, 1986): gill arches of Plagioscion squamossisimus: Brazil
(Amazonia). (Figs. 8-39 A-O & 8-48, 8-49, 8-51, 8-52). Body = 2.4-7.3 x
0.30-0.46 mm; Thoraco-abdomen = 0.87-1.5 x 0.30-0.46 mm; Head =
0.18-0.20 x 0.10-0.18; Neck of variable length.
T. tamarae AMADO & ROCHA, 1996: gill arches of Plagioscion squamossisimus:
Brazil (Amazonia). Body = 0.58 mm; Neck and cephalic shield = 0.44 mm
long. (Note: This species was distinguished from T. elongatus mainly on the
basis of a 2 segmented first endopod).
VIII. Plates of Copepoda (Figs. 8-1 to 8-62)
8-1 A
8-1. Ergasilus hydrolycus: female; A. entire (dorsal); B. genital segment, abdomen and uropods; C. egg
sac; D. antennule; (scale = 250 µm).
8-2
8-3
8-4
8-5
Ergasilus bryconis: 8-2. nauplius I; 8-3. copepodid I; 8-4. copepodid III; 8-5. copepodid V (all from
VARELLA 1985).
8-6 B
8-6 A
8-7 A
8-7 B
8-7 C
Ergasilus bryconis: 8-6. A. female (entire); B. antenna; C. antenna; 8-7. A. male (entire); B. maxilliped;
C. antenna.
8-8 A
100
250
B
100
C
8-8. Ergasilus callophysus: female; A. entire; B. antenna; C. egg sac; D. antennule; (scales in µm).
100
250
100
8-9 A 100
C
D
50 E
8-9. Ergasilus colomesus: female; A. entire (dorsal); B. antenna; C. ventral thoracic plates; D. genital
segment, abdomen and uropods; E. antennule; (scales in µm).
8-10 A
200
100
B C D
0
10
8-10. Ergasilus jaraquensis: female; A. entire (dorsal); B. egg sac; C. antenna; D. fifth legs, genital
segment, abdomen and uropods; (scales in µm).
8-11
B 50
250
C
100
50
D
8-11. Ergasilus pitalicus: female; A. entire (dorsal); B. antennule; C. base of antennule; D. antenna;
(scales in µm).
8-12
100
A
B
250
100
8-12. Acusicola lycengraulidis: female; A. entire (dorsal); B. genital segment, abdomen and uropods; C.
antenna; (scales in µm).
8-13
A B
150
100
500
100
8-13. Acusicola pellonidis: female; A. entire (dorsal); B. genital segment, abdomen and uropods; C.
antenna; D. latched antennae; (scales in µm).
8-14
50
A
B
250
50
100
8-14. Acusicola tucunarense: female; A. entire (dorsal); B. antenna; C. genital segment, abdomen and
uropods; D. antennule; (scales in µm).
8-16
8-15
8-17
8-18 8-19
Antennae: 8-15. Ergasilus holobryconis; 8-16. E. iheringi; 8-17. Rhinergasilus piranhus; 8-18. Acusicola tenax;
8-19. A. cunula.
8-20 8-21
8-22 8-23
Antennae: 8-20. Ergasilus leporinidis; 8-21. E. pitalicus. 8-22. E. euripidesi; 8-23. E. argulus.
8-24
250
B
50
25
8-24. Brasergasilus jaraquensis: female; A. entire (dorsal); B. genital segment, abdomen and uropods; C.
egg sac; D. antennule; (scales in µm).
250
C
50
8-25
8-25. Brasergasilius anodus: female; A. entire (dorsal); B. genital segment, abdomen and uropods; C.
egg sac; (scales in µm).
25
8-26
25
8-27
25
8-28
Antennae of Brasergasilus: 8-26. B. anodus; 8-27. B. oranus; 8-28. B. jaraquensis; (scales in µm).
8-29
50
C 200
50
D E
50
8-29.Vaigamus retrobarbatus: A. female (dorsal); B. male (dorsal); C. maxilliped (male); D. antenna

(female); E. antenna (male); (scales in µm).
8-30 A
8-30. Rhinergasilus piranhus BOEGER & THATCHER, 1988. Central Amazon, Brazil; A. dorsal view; B.
egg sac; scales = 100 µm.
8-31
A B
200
50
C
8-31. Gamidactylus jaraquensis: female; A. entire (dorsal); B. antenna; C. antennule; D. egg sac; E.
genital segment, abdomen and uropods; (scales in µm).
8-32
A B
8-32. Gamispinus diabolicus: female; A. entire (dorsal); B. antenna; C. antennule; D. egg sac; E. genital
segment, abdomen and uropods.
8-33
C
50
8-33. Gamispatulus schizodontis: female; A. entire (dorsal); B. rostral projection (ventral); C. egg sac; D.
antenna; (scale in µm).
8-34
8-35
8-36
Vaigamid retrostylets: 8-34. Gamidactylus jaraquensis; 8-35. Gamispinus diabolicus; 8-36. Gamispatulus
schizodontis.
8-37
50
A B
250
250
50
C D E
20
8-37 A-E. Amplexibranchius bryconis: female; A. entire (dorsal); B. antennule; C. leg 5; D. egg sac; E.
genital segment, abdomen and uropods; (scales in µm).
8-37
F 50
G
20
250
50
I
H
8-37 F-I. Amplexibranchius bryconis: female; F. latched antennae; G. mouthparts; H. leg 4; I. antenna;
(scales in µm).
C
8-38
E
D
8-38. Ergasiloid antennae: A. Gamispatulus schizodontis; B. Prehendorastrus bidentatus; C. P. monodontus; D.

Brasergasilus sp. (showing 3-segmented antenna); E. undescribed genus (showing 4-segmented antenna).
8-39
500
50
1000
A B C
50
50
8-39. Therodamas elongatus: post-metamorphic female: A. entire (ventral); B. antenna; C. hindbody; D.

antennule; E. uropod and abdominal segments; (scales in µm).
8-39
100
100
50
50
J K
8-39. Therodamas elongatus: post-metamorphic female: F. mouthparts (lateral); G. head capsule (lateral);
H. mouthparts (ventral, suction tube withdrawn); I. leg I; J. leg II (= leg III); K. leg IV; (scales in µm).
8-39
8-39. Therodamas elongatus: L. anterior extremity (lateral view); M. calcifications around “neck”; N.
anterior extremity absorbed by action of host; O. mouth tube (lateral view). (L & O: scale = 50 µm;
M & N: scale = 100 µm).
8-40 A
C A-2
A-1
Mx
Mxp
8-40.Taurocheros salminisii: female; A. entire (ventral); B. anterior extremity (ventral); C. mouthparts;

A-1 = antennule; A-2 = antenna; Mx = maxilla; Mxp = maxilliped (all after BRIAN 1924).
8-41
8-41. Taurocherus tarangophilus PAGGI, 1976. Rio Paraná, Argentina; scale = 2 mm.
8-42
8-42. Perulernaea gamitanae THATCHER & PAREDES, 1985. Iquitos, Peru; scale = 2mm.
8-43
8-43. Perulernaea pirapitingae THATCHER, 2000. Rio Meta, Colombia; scale = 1 mm.
8-44
8-44. Amazolernaea sannerae THATCHER & WILLIAMS, 1998. Venezuela; scale = 1 mm.
8-45
8-45. Bedsylernaea collaris THATCHER & WILLIAMS, 1998. Rio Amazonas, Brazil; scale = 2 mm.
8-46
8-46. Minilernaea floricapitella THATCHER & HUERGO, 2005. Astyanax spp. Rio Iguaçu, Santa Catarina,
Brazil; scales: A = 1 mm; B = 200 µm.
8-47
8-47. Lernaea cyprinacea: L. Rio Tibagi, Paraná, Brazil; scale = 5 mm.

8-48 8-49 8-50
8-51 8-52
Therodamasidae and Lernaeidae: 8-48 & 8-49. anterior extremities of Therodamas sp.; 8-50. head of
premetamorphic female of Perulernaea gamitanae; 8-51. head of premetamorphic female of Therodamas
elongatus; 8-52. posterior extremity of female of T. elongatus.
8-53 8-54
8-53. Miracetyma etimaruya: female. A. dorsal view; scale = 500 µm; B. antenna; C. leg 1; 8-54. Pindapixara
tarira: female, dorsal view; scale = 200 µm.
8-55 A
8-56 A B
8-55. Brasergasilus guaporensis: female. A. dorsal view; scale = 200 µm; B. antenna. 8-56. Gamidactylus
bryconis: female. A. antenna; B. dorsal view; scale = 200 µm.
8-57 8-58
8-59 8-60
8-57 to 8-60. Undescribed Ergasilus females from plankton showing coloration and pigment
distribution.
8-61
8-62
Female ergasilids on gill filaments showing pathology: 8-61. Acusicola tucunarense demonstrating
constriction and terminal necrosis of the filament; 8-62. Ergasilus colomesus, showing epithelial
hyperplasia and accumulation of pigments; scale = 1 mm.
AMADO, M.A.P. & C.E.F. DA ROCHA (1996): Therodamas tamarae, a new species of copepod
(Poecilostomatoida: Ergasilidae) parasitic on Plagioscion squamosissiumus (HECKEL) from the
Araguaia River, Brazil; with a key to the species of the genus. - Hydrobiologia 325: 77-82.
ARAUJO, C.S. DE & A. VARELLA (1998): Ergasilus coatiarus sp. n. (Copepoda, Poecilostomatoida,
Ergasilidae) parasita dos filamentos branquiais de Cichla monoculus SPIX, 1831 (Perciforme:
Cichlidae) da Amazônia Brasileira. - Acta Amazonica 28(4): 417-424.
BOEGER, W.A. & V.E. THATCHER (1988): Rhinergasilus piranhus gen. et sp. n. (Copepoda, Poecilosto-
matoida, Ergasilidae) from the nasal cavities of piranha cajú, Serrasalmus nattereri, in the
Central Amazon. - Proc. Helminthol. Soc. Wash. 55(1): 87-90.
BOEGER, W.A. & V.E. THATCHER (1990): Prehendrorastrus gen. n. (Poecilostomatoida, Ergasilidae)
with descriptions of two new species from the gill rakers of Hypophthalmus spp. (Teleostei,
Siluriformes) from the Brazilian Amazon. - Syst. Parasit. 17: 133-141.
BOEGER, W.A., MARTINS, M. & V.E. THATCHER (1993): Ergasilus hypophthalmus sp. n. (Copepoda,
Poecilostomatoida, Ergasilidae) from the gill rakers of the Amazonian catfishes, Hypophthal-
mus edentatus SPIX, 1826, and H. fimbriatus KNER, 1857 (Teleostei, Siluriformes). - Rev. Brasil.
Biol. 53(3): 421-424.
BOXSHALL, G.A. (1986): A new species of Mugilicola TRIPATHI (Copepoda: Poecilostomatoida) and a
review of the family Therodamasidae. - Proc. Linn. Soc. N.S.W. 108: 183-186.
BRIAN, A. (1924): Descrizione di un nuovo e curioso copepode lerneideo parasita de Salminus
brevidens raccolto dal Prof. Filippo Silvestri nell’America del Sul. - Boll. Lab. Zool. Gen. e
Agrar. R. Scuola Super. Agric. Portici 18: 32-36.
CRESSEY, R.F. & B.B. COLLETTE (1970): Copepods and needlefishes: a study in host-parasite
relationships. - Fish. Bull. 68: 347-432.
ENGERS, K.B., BOEGER, W.A. & D.A. BRANDÃO (2000): Ergasilus thatcheri n. sp. (Copepoda,
Poecilostomatoida, Ergasilidae) from the gills of Rhamdia quelen (Teleostei, Siluriformes,
Pimelodidae) from Southern Brazil. - J. Parasitol. 86(5): 945-947.
KABATA, Z. (1979): Parasitic Copepoda of British fishes. - The Ray Society, London: 468 pp.,
2031 figs.
MALTA, J.C.O. (1993a): Miracetyma etimaruya gen. et. sp. n. (Copepoda, Poecilostomatoida: Ergasil-
idae) from freshwater fishes of the Brazilian Amazon. - Acta Amazonica 23(1): 49-57.
MALTA, J.C.O. (1993b): Miracetyma kawa sp. n. (Copepoda, Poecilostomatoida: Ergasilidae) dos
peixes de água doce da Amazônia Brasileira. - Acta Amazonica 23(2-3): 251-259.
MALTA, J.C.O. (1993c): Myracetyma piraya sp.n. (Copepoda: Ergasilidae) das brânquias de Pygocentrus
nattereri (KNER, 1860) (Characiformes: Serrasalmidae) da Amazônia Brasileira. - Acta Amazon-
ica 23(2-3): 261-269.
MALTA, J.C.O. (1993d): Brasergasilus guaporensis sp.n. (Copepoda: Ergasilidae) das brânquias de
Leporinus fasciatus (BLOCH, 1890) (Characiformes: Anastomidae) da Amazônia Brasileira. - Acta
Amazonica 23(4): 441-447.
MALTA, J.C.O. (1993e): Ergasilus urupaensis sp.n. (Copepoda: Ergasilidae) das brânquias de Prochilodus
nigricans AGASSIZ, 1829 (Characiformes: Prochilodontidae) da Amazônia Brasileira. - Acta
Amazonica 23(4): 449-456.
MALTA, J.C.O. (1994a): Pindapixara tarira gen. et sp.n. (Copepoda: Ergasilidae) das brânquias de
Hoplias malabaricus (BLOCH, 1794) (Characiformes: Erythrinidae) da Amazônia Brasileira. -
Acta Amazonica 24(1/2): 135-144.
MALTA, J.C.O. (1994b): Ergasilus triangularis n. sp. (Copepoda: Ergasilidae) das brânquias de
Laemolyta taeniata (KNER, 1859), (Characiformes: Anostomidae) da Amazônia Brasileira. - Acta
Amazonica 24(3/4): 309-316.
MALTA, J.C.O. & A. VARELLA (1986): Ergasilus holobryconis sp. n. crustáceo parasita de Holobrycon pesu
(MÜLLER & TROSCHELL), um peixe da Amazônia brasileira (Copepoda: Poecilostomatoida:
Ergasilidae). - Amazoniana 10(1): 41-48.
MALTA, J.C.O. & A. VARELLA (1996): Ergasilus turucuyus sp. n. (Copepoda: Ergasilidae) das brânquias
de Acestrorhynchus falcatus (BLOCH, 1794) e A. falcirostris (CUVIER, 1819) (Characidae) da
Amazônia Brasileira. - Acta Amazonica 26(1/2): 69-76.
MALTA, J.C.O. & A. VARELLA (1998): Maxillopoda - Copepoda Poecilostomatoida. Non-marine parasites. -
In: YOUNG, P.S. (ed.): Catalogue of Crustacea of Brazil: 241-249. Rio de Janeiro (Série Livros n. 6).
MONTU, M. (1980): Parasitic copepods of Southern Brazil fishes, I. Ergasilus euripedesi n. sp.
(Copepoda, Cyclopidea). - Iheringia, Ser. Zool. (Porto Alegre, Brazil) 56: 53-62.
PAGGI, J.C. (1976): Una nueva especie de Therodamas (Therodamasidae; Cyclopoida) copépodo
parásito de peces de agua dulce de la República Argentina. - Physis 35(91): 93-102.
PAIVA CARVALHO, J. (1955): Ergasilus xenomelanirisi n. sp. parasito de peixe-rei - Xenomelaniris
brasiliensis (QUOY & GAIMARD). - Bol. Inst. Oceanogr. (Brazil) 6(1-2): 215-224.
PAIVA CARVALHO, J. (1962): Ergasilus cyanopictus sp. nov., parasito de tainha - Mugil cephalus (L.)
(Crustacea, Copepoda - Pisces, Mugiloidei). - Arq. Mus. Nac. Rio de Janeiro 52: 31-36.
PIASECKI, W., KHAMEES, N.R. & F.T. MHAISEN (1991): A new species of Mugilicola TRIPATHI, 1960
(Crustacea, Copepoda, Therodamasidae) parasitic on Iraqui fish. - Acta Ichthyologica et
Piscatoria 21(2): 143-151.
ROBERTS, L.S. (1965): Ergasilus tenax sp. n. (Copepoda, Cyclopoida) from the white crappie, Pomoxis
annularis RAFINESQUE. - J. Parasitol. 51(6): 987-989.
ROBERTS, L.S. (1970): Ergasilus (Copepoda: Cyclopoida): Revision and key to species in North
America. - Trans. Am. Mic. Soc. 89(1): 134-161.
THATCHER, V.E. (1981a): Os crustáceos parasitos de peixes da Amazônia brasileira. 1. Ergasilus
bryconis n. sp. (Copepoda: Cyclopidea) da matrinchã (Brycon melanopterus (COPE)). - Acta
Amazonica 11: 437-442.
THATCHER, V.E. (1981b): Os crustáceos parasitos de peixes da Amazônia brasileira. 2. Ergasilus
leporinidis n. sp. (Copepoda: Cyclopoidea) das branquias de Leporinus fasciatus (BLOCH). - Acta
Amazonica 11: 723-727.
THATCHER, V.E. (1981c): Ergasilus pitalicus n. sp. (Copepoda: Poecilostomatoida: Ergasilidae) a gill
parasite of a cichlid fish from the Pacific Coast of Colombia, South America. - J. Crust. Biol.
4(3): 495-501.
THATCHER, V.E. (1984): The parasitic crustaceans of fishes from the Brazilian Amazon. 7. Acusicola
tucunarense n. sp. (Copepoda: Cyclopidea) from Cichla ocellaris, with an emendation of the
genus and the proposal of Acusicolinae subfam. nov. - Rev. Brasil. Biol. 44(2): 181-189.
THATCHER, V.E. (1986): The parasitic crustaceans of fishes from the Brazilian Amazon, 16.
Amazonicopeus elongatus gen. et sp. n. (Copepoda: Poecilostomatoida) with the proposal of
Amazonicopeidae fam. nov. and remarks on its pathogenicity. - Amazoniana 10(1): 49-56.
THATCHER, V.E. (1998): Copepods and fishes in the Brazilian Amazon. - Journal of Marine Systems
(Holland) 15: 97-112.
THATCHER, V.E. (1999): Anklobrachius marajoensis gen. et sp. nov. (Copepoda, Poecilostomatoida,
Ergasilidae) described from male specimens taken in plankton samples from the Amazon
River. - In: SHRAM, E.J. & J.C. VON VAUPEL KLEIN (eds.): Crustaceans and the biodiversity
crisis: 231-237. Proc. Fourth Int. Crustacean Congr., Amsterdam. Brill, Leiden: 1021 pp.
THATCHER, V.E. (2000): Perulernaea pirapitingae n. sp. (Copepoda: Lernaeidae) a parasite of the serrasalmid
fish, Piaractus brachypomus from the Meta River, Colombia. - Amazoniana 16(1/2): 249-257.
THATCHER, V.E. & W.A. BOEGER (1983a): Patologia de peixes da Amazônia brasileira, 3. Alterações
histológicas em brânquias provocadas por Ergasilus, Brasergasilus e Acusicola (Crustacea:
Cyclopoida: Ergasilidae). - Acta Amazonica 13(2): 441-451.
THATCHER, V.E. & W.A. BOEGER (1983b): The parasitic crustaceans of fishes from the Brazilian
Amazon. 4. Ergasilus colomesus sp.n. (Copepoda: Cyclopoida) from an ornamental fish (Colomesus
asellus: Tetraodontidae) and aspects of its pathogenicity. - Trans. Am. Mic. Soc. 102: 371-379.
THATCHER, V.E. & W.A. BOEGER (1983c): The parasitic crustaceans of fishes from the Brazilian
Amazon, 5. Brasergasilus gen. nov. (Copepoda: Cyclopidea), a “three-legged” ergasilid with two
new species and the proposal of Abergasilinae subfam. nov. - Acta Amazonica 13(1): 195-214.
THATCHER, V.E. & W.A. BOEGER (1983d): The parasitic crustaceans of fishes from the Brazilian
Amazon, 10. Acusicola pellonidis n. sp. (Copepoda: Cyclopidea) from Pellona castelnaeana
(VALLENCIENNES). - Amazoniana 8(2): 273-279.
THATCHER, V.E. & W.A. BOEGER (1984a): The parasitic crustaceans of fishes from the Brazilian
Amazon. 9. Ergasilus callophysus sp.n. (Copepoda: Cyclopoida) from Callophysus macropterus
(LICHTENSTEIN). - Proc. Helminthol. Soc. Wash. 51: 326-330.
THATCHER, V.E. & W.A. BOEGER (1984b): The parasitic crustaceans of fishes from the Brazilian
Amazon. 13. Gamidactylus jaraquensis gen. et sp. n. (Copepoda: Poecilostomatoida: Vaigamidae)
from the nasal fossae of Semaprochilodus insignis (SCHOMBURGK). - Amazoniana 8(3): 421-426.
THATCHER, V.E. & W.A. BOEGER (1984c): The parasitic crustaceans of fishes from the Brazilian
Amazon. 6. Brasergasilus oranus sp.n. (Copepoda: Cyclopoida) from Anodus elongatus SPIX. - Rev.
Brasil. Biol. 44(4): 181-189.
THATCHER, V.E. & W.A. BOEGER (1984d): The parasitic crustaceans of fishes from the Brazilian
Amazon. 15. Gamispatulus schizodontis gen. et sp. n. (Copepoda: Poecilostomatoida: Vaigami-
dae) from Schizodon fasciatus AGASSIZ. - Amazoniana 9(1): 119-126.
THATCHER, V.E. & W.A. BOEGER (1984e): The parasitic crustaceans of fishes from the Brazilian
Amazon. 14. Gamispinus diabolicus gen. et sp. n. (Copepoda: Poecilostomatoida: Vaigamidae)
from the nasal fossae of Ageneiosus brevifilis VALENCIENNES. - Amazoniana 8(4): 505-510.
THATCHER, V.E. & W.A. BOEGER (1985): The parasitic crustaceans of fishes from the Brazilian
Amazon. 8. Acusicola lyncengraulidis n. sp. (Copepoda: Cyclopidea) from Lycengraulis grossidens
CUVIER. - Acta Amazonica 13(5-6): 943-951.
THATCHER, V.E. & V. PAREDES (1985a): A parasitic copepod, Perulernaea gamitanae gen. et sp. nov.
(Cyclopoida: Lernaeidae), from the nasal fossae of a Peruvian Amazon food fish. -
Amazoniana 9(2): 169-175.
THATCHER, V.E. & V. PAREDES (1985b): A parasitic copepod, Amplexibranchius bryconis gen. et sp. nov.
(Ergasilidae: Acusicolinae), from an Amazonian fish and remarks on the importance of leg
morphology in this subfamily. - Amazoniana 9(2): 205-214.
THATCHER, V.E. & J. PEREIRA, JR. (2004): Brasilochondria riograndensi gen. nov., sp. nov. (Copepoda,
Chondracanthidae) a parasite of flounders of Rio Grande do Sul State, Brazil. - Rev. Bras.
Zool. 21: 515-518.
THATCHER, V.E. & B. ROBERTSON (1982): The parasitic crustaceans of fishes from the Brazilian
Amazon, 3, Ergasilus jaraquensis sp. nov. (Copepoda: Cyclopoidea), from the gills of
Semaprochilodus insignis (SCHOMBURGK). - Rev. Bras. Biol. 42: 503-513.
THATCHER, V.E. & B. ROBERTSON (1984): The parasitic crustacean of fishes from the Brazilian
Amazon, 11. Vaigamidae fam. nov. (Copepoda: Poecilostomatoida) with males and females of
Vaigamus retrobarbatus gen. et sp. nov. and and V. spinicephalus sp. nov. from plankton. - Can. J.
Zool. 62: 716-729.
THATCHER, V.E. & E.H. WILLIAMS, JR. (1998): Comparative morphology of four lernaeids
(Copepoda: Cyclopoida) from South American fish with descriptions of two new genera. - J.
Aquat. Anim. Health. 10: 300-308.
THATCHER, V.E. BOEGER, W.A. & B.A. ROBERTSON (1984): The parasitic crustacean of fishes from
the Brazilian Amazon. 12. Ergasilus hydrolycus sp. n. (Copepoda: Poecilostomatoida) from
Hydrolycus scomberoides (CUVIER). - Amazoniana 8(3): 321-326.
TRIPATHI, Y.R. (1960): Parasitic copepods from Indian fishes. 2. Two new families Therodamasidae and
Cataphilidae. – In: Libro de homenaje al Dr. Eduardo Caballero y Caballero, Mexico: 543-548.
VARELLA, A.M. (1985): O ciclo biológico de Ergasilus bryconis THATCHER, 1981 (Crustacea:
Poecilostomatoida, Ergasilidae) parasita das brânquias do matrinxã Brycon erythropterum (COPE,
1872) e aspectos de sua ecologia. - M.Sc.-thesis, INPA/FUA, Manaus, Brazil: 100 pp.
VARELLA, A.M. (1995): Gamidactylus bryconis sp. n. (Copepoda, Poecilostomatoida, Vaigamidae) das
fossas nasais de peixes, Brycon pellegrini HOLLY, 1929 e B. melanopterus (COPE, 1872) da
Amazônia Brasileira. - Acta Amazonica 24(1/2): 145-152.
VARELLA, A.M.B. & J.C.O. MALTA (2001): Brasergasilus mamorensis sp. n. (Copepoda: Ergasilidae)
from the nasal cavities of Hydrolycus pectoralis (GUENTHER, 1866) (Characiformes: Cynodon-
tidae) from the Brazilian Amazon, and considerations about Abergasilinae. - Acta Amazonica
30(1): 323-330.
WILSON, C.B. (1911): North American parasitic copepods belonging to the family Ergasilidae. -
Proc. U. S. Nat. Mus. 39: 263-400.
YAMAGUTI, S. (1963): Parasitic Copepoda and Branchiura of fishes. - Intersci. Publ., New York:
1104 pp., 333 pls.
9.
BRANCHIURA
Branchiurans are small, shield-shaped crustaceans that are ectoparasites on the skin and
within the gill chamber of marine and freshwater fish. They range from 3 to 30 mm in
length and since they are easy to see, they are well known to fishermen and pisciculturists
who have termed them “fish lice”. The term is not inappropriate since branchiurans do
suck blood and can weaken and kill their hosts. Both sexes parasitize fish and they can
leave the host at will. All known branchiurans belong to the family Arguilidae and perhaps
“argulids” is a more appropriate name for the group.
Argulids were once thought to be copepods, but they are morphologically quite
distinct. There is a large concave carapace fused to the head which has wing-like
postero-lateral extensions containing respiratory areas. There are four biramous swim-
ming legs on the thorax, and the abdomen is fused into a single bilobed unit. The first
antennae and both pairs of maxillae are adapted for clinging to fish skin. In Argulus and
Dipteropeltis, the first maxillae are modified to form sucker-like structures and in Dolops
they are prehensile and terminate in strong claws. The ventral surface of the carapace
and thorax is also frequently provided with retro-spines as an additional attachment aid.
The ventral structures of a typical arguilid are shown in Fig. 9-1. Dorsally, argulids have
a pair of moveable compound eyes and often there is green or yellow pigmentation in
definite patterns.
Transmission of argulids is direct since they swim actively up to a fish and attach
themselves. They can leave the host at any time and move to a different fish. They are also
able to skate around over the surface of a fish to find an appropriate feeding site. Argulids
frequently select a site that is less heavily scaled, such as the base of the fins. Some species
show a preference for the inside of the gill chamber.
When mature enough to mate, argulids leave the host and copulate while swimming.
Afterwards, the female lays her eggs by gluing them in linear strings or small clusters (the
pattern is species dependent) on hard surfaces such as submerged logs or rocks. In an
infected aquarium it is sometimes easier to see the egg strings on the glass than the adult
animals. After an incubation period of 10-20 days, the young argulids hatch and
immediately swim out looking for a host fish on which to mature.
In studies made in the Brazilian Amazon in 1979 (MALTA 1981, 1982 a, b, 1983;
MALTA & VARELLA 1983), it was found that argulids on wild fish populations show host
and site preference and seasonal variation in numbers. For example, Argulus multicolor, A.
juparanaensis and Dolops geayi were found only in the gill chambers of their hosts while
Argulus pestifer, Dolops bidentata, D. carvalhoi, D. discoidalis and D. striata infest only the skin.
Of these species, A. pestifer proved to be the most host specific being found only on two
species of Pseudoplatystoma. A. pestifer was also the only species found to be more numerous
on the fish during the low water stage of the river (November and December). All other
species studied were more common during June and July when the river level was high.
III. Pathology
In the act of feeding, argulids perforate the skin of the fish host with their mouth parts
and inject anticoagulants and digestive secretions. The argulid mouth is a tubular structure
with cutting mandibles near its apex. In Argulus there is also a preoral stylet to help the
animal penetrate the fish skin. Argulids ingest blood and epithelial cells and can produce
anemia. There is a localized inflammatory reaction in the vicinity of the bite and
secondary bacterial and fungal infections may also occur. Heavy infestations can bring on
the death of the fish especially in young or small animals.
Argulid infestations are difficult to either prevent or treat. Frequently the first indication
of argulids in an aquarium are the eggs glued on the glass. When argulids are discovered,
it is necessary to treat the fish, move them to a clean aquarium and disinfect the old
aquarium. Fish can be treated in a dip prepared with Neguvon at a concentration of 20
cm3 per liter of water for 2 minutes. The most convenient way to sterilize aquaria is to fill
them with a 5 % solution of Sodium Hypochlorite (household bleach) and leave them for
10 hours. Infested ponds should be drained and quick lime should be scattered over the
muddy bottom. After two days, the ponds can be refilled and used.
Argulids are best removed from the fish with soft forceps and dropped alive into 70 %
alcohol or AFA solution. Temporary study preparations can be made in pure phenol or in
glycerin. Permanent slides can be made by the phenol-balsam method (explained in detail
in Chapter 6).
392
Some characters used in the identification of argulids are: size, shape and color; number of legs covered by carapace; form of maxillae;

and form and number of maxillary teeth.
Key to the Neotropical genera of Branchiura
I. First maxillae sucker-like.

A. Basal plates of second maxillae without teeth ............................................................................................... Dipteropeltis hirundu (Fig. 9-31)
B. Basal plates of second maxillae armed with teeth ................................................................................................................................. Argulus
II. First maxillae simple, armed with claws ........................................................................................................................................................... Dolops
Key to the Neotropical species of Argulus
I. Carapace short, covering only first pair of legs ................................................................................................................... A. elongatus (Fig. 9-9)
II. Carapace longer, covering first 2 pairs of legs.
A. Respiratory areas contiguous.
1. Larger respiratory area with medial indentation .......................................................................................... A. spinulosus (Fig. 9-7 A-D)
2. Larger respiratory area without medial indentation .................................................................................................. A. ichesi (Fig. 9-11)
B. Respiratory areas close together, the smaller ones are subtriangular .................................................................... A. multicolor (Fig. 9-14)
C. Respiratory areas well separated.
1. Small respiratory area hemispherical ................................................................................................................... A. patagonicus (Fig. 9-13)
2. Small respiratory area subtriangular ............................................................................................................ A. violaceus (Figs. 9-1 & 9-19)
III. Carapace long, covering first 3 pairs of legs.
A. Abdominal lobes long and slender, 4 to 5 times longer than wide .................................................................... A. paranaensis (Fig. 9-22)
B. Abdominal lobes short.
1. Lobes of carapace reaching to near base of abdomen.
a. Anterior (antennal) area of carapace supported by rods; respiratory areas contiguous; smaller area medial to larger area .
...................................................................................................................................................................................... A. pestifer (Fig. 9-30)
b. Antennal area without rods; respiratory areas well separated; smaller anterior to larger ........ A. salmini (Figs. 9-10 & 9-18)
2. Lobes of carapace well separated from base of abdomen.
a. Maxillary teeth sharply pointed.
1) Marginal support rods of suckers consisting of stacks of subspherical masses ................................... A. ernsti (Fig. 9-20)
2) Marginal support rods of suckers consisting of L-shaped pieces topped by 2 subspherical masses ..................................
......................................................................................................................................................................... A. carteri (Fig. 9-4 A-D)
b. Maxillary teeth flattened, spatulate ............................................................................................................... A. cubensis (Fig. 9-8 A-C)
IV. Carapace very long, covering all 4 pairs of legs.
A. Carapace nearly spherical.
1. Posterior sinus of carapace wide ................................................................................................................................. A. silvestrii (Fig. 9-6)
2. Posterior sinus of carapace narrow and deep .......................................................................................................... A. nattereri (Fig. 9-5)
B. Carpace not spherical; longer than wide.
1. Maxillary teeth rounded ..................................................................................................................................... A. rhamdiae (Fig. 9-3 A-D)

2. Maxillary teeth squared ................................................................................................................................ A. juparanaensis (Fig. 9-2 A-D)
Key to the Neotropical species of Dolops
I. Basal plate of second maxilla armed with 2 teeth.

A. Ventral surface of carapace spinous .............................................................................................................................. D. bidentata (Fig. 9-33)
B. Ventral surface of carapace without spines ............................................................................................................... D. nana (Fig. 9-23 A-D)
II. Basal plate of second maxilla armed with 3 teeth.
A. Abdominal lobes long and slender.
1. Abdominal lobes longer than carapace; respiratory areas of 3 parts on each side ............................ D. longicauda (Fig. 9-25 A-B)
2. Abdominal lobes shorter than carapace; respiratory areas lacking .............................................................................................. D. geayi
B. Abdominal lobes short, about as wide as long ....................................................................................... D. reperta (Fig. 9-24, 9-28 & 9-29)
393
C. Abdominal lobes of medium length, usually somewhat longer than wide.
394
1. Abdominal sinus shallow ................................................................................................................................................ D. kollari (Fig. 9-26)
2. Abdominal sinus deep.

a. Ventral surface of carapace without spines ......................................................................................................................... D. carvalhoi
b. Ventral surface of carapace spinous.
1) Marginal spination continuous to near posterior border ..................................................................... D. discoidalis (Fig. 9-15)
2) Marginal spination terminates at level of first leg ........................................................................................................... D. striata
Clave para los géneros Neotropicales de Branchiura
I. Primeras maxilas semejantes a una ventosa.

A. Placas basales de las segundas maxilas sin dientes ........................................................................................ Dipteropeltis hirundo (Fig, 9-31)
B. Placas basales de las segundas maxilas, armadas con dientes .............................................................................................................. Argulus
II. Primeras maxilas simples, armadas con garras ...............................................................................................................................................Dolops
Clave para las especies Neotropicales de Argulus
I. Caparazón corto, cubriendo solamente el primer par de piernas .................................................................................... A. elongatus (Fig. 9-9)
II. Caparazón más largo, cubriendo los dos primeros pares de piernas.
A. Areas respiratorias contiguas.
1. Area respiratoria mayor con indentación en el medio ................................................................................ A. spinulosus (Fig. 9-7 A-D)
2. Area respiratoria mayor sin indentación ...................................................................................................................... A. ichesi (Fig. 9-11)
B. Areas respiratorias juntas, las más pequeñas son casi triangulares ........................................................................ A. multicolor (Fig. 9-14)
C. Areas respiratorias bien separadas.
1. Area respiratoria pequeña es hemisférica ........................................................................................................... A. patagonicus (Fig. 9-13)
2. Area respiratoria pequeña es casi triangular .............................................................................................. A. violaceus (Figs. 9-1 & 9-19)
III. Caparazón largo, cubriendo los tres primeros pares de piernas.
A. Lóbulos abdominales largos y delgados, 4 a 5 veces más largos que anchos .................................................. A. paranaensis (Fig. 9-22)
B. Lóbulos abdominales cortos.
1. Lóbulos del caparazón alcanzan hasta cerca de la base del abdomen.
a. Area anterior (antenal) del caparazón es sostenida por varillas; áreas respiratorias contiguas; las pequeñas más cerca a la
línea media ................................................................................................................................................................. A. pestifer (Fig. 9-30)
b. Area antenal sin varillas; áreas respiratorias bien separadas; la menor anterior a la mayor ....... A. salmini (Figs. 9-10 & 9-18)
2. Lóbulos del caparazón bien separados de la base del abdomen.
a. Dientes maxilares afilados.
1) Varillas de soporte marginal de las ventosas consisten en grupos de masas esféricas ......................... A. ernsti (Fig. 9-20)
2) Varillas de soporte marginal de las ventosas consisten en piezas en forma de L sosteniendo 2 masas semiesféricas .....
......................................................................................................................................................................... A. carteri (Fig. 9-4 A-D)
b. Dientes maxilares aplanados, espatulados ................................................................................................... A. cubensis (Fig. 9-8 A-C)
IV. Caparazón bien largo, cubriendo las 4 pares de piernas.
A. Caparazón casi esférico.

1. Espacio entre lóbulos abdominals ancha ................................................................................................................... A. silvestrii (Fig. 9-6)
2. Espacio entre lóbulos abdominals angosto y profundo ......................................................................................... A. nattereri (Fig. 9-5)
B. Caparazón no esférico, más largo que ancho.
1. Dientes maxilares arredondeados ..................................................................................................................... A. rhamdiae (Fig. 9-3 A-D)
2. Dientes maxilares cuadrados ...................................................................................................................... A. juparanaensis (Fig. 9-2 A-D)
Clave para las especies Neotropicales de Dolops
I. Placa basal de la segunda maxila armada con 2 dientes.

A. Superficie ventral del caparazón espinosa ..................................................................................................................... D. bidentata (Fig. 9-33)
B. Superficie ventral del caparazón sin espinas .............................................................................................................. D. nana (Fig. 9-23 A-D)
395
II. Placa basal de la segunda maxila armada con 3 dientes.
396
A. Lóbulos abdominals largos y finos.
1. Lóbulos abdominales más largos que el caparazón; 3 áreas respiratorias de cada lado .................... D. longicauda (Fig. 9-25 A-B)

2. Lóbulos abdominales más cortos que el caparazón; áreas respiratorias ausentes .................................................................... D. geayi
B. Lóbulos abdominales cortos, ancho y largo parecidos .......................................................................... D. reperta (Fig. 9-24, 9-28 & 9-29)
C. Lóbulos abdominales de largura intermedia, con frecuencia más largos que anchos.
1. Espacio abdominal poco profundo ............................................................................................................................. D. kollari (Fig. 9-26)
2. Espacio abdominal profundo.
a. Superficie ventral del caparazón sin espinas ....................................................................................................................... D. carvalhoi
b. Superficie ventral del caparazón espinoso.
1) Espinas marginales continuas hasta cerca del borde posterior ........................................................... D. discoidalis (Fig. 9-15)
2) Espinas marginales terminan al nivel de la primera pierna ........................................................................................... D. striata
VII. Checklist of Branchiura from Neotropical freshwater fishes
All measurements are given in millimeters.
Argulidae LEACH, 1819

Carapace flattened, shield-like. Abdomen bilobed posteriorly, provided
with small papillae medially. Basal portions of first antennae flattened and
armed with hooks and spines; terminal portion cylindrical, 2 or 3-
segmented. Preoral sheathed stylet present or absent. First maxillae tipped
with claws or modified to form sucker-like structures. Second maxillae
with basal plates having spines, teeth or lobes posteriorly. Basal segments
of female fourth legs usually expanded to flattened lobes; basal segments
of posterior legs of male usually modified for reproductive function.
Anterior legs with or without flagella on distal end of basis. Ovary in
posterior portion of thorax; testes in abdomen. Ectoparasites on freshwa-
ter and marine fishes and occasionally amphibians.
Argulus MÜLLER, 1785
Argulidae. Two pairs of antennae present. Preoral stylet usually present.
First maxillae develop into sucker-like cups. Basal plate of second maxillae
armed with teeth. On freshwater and marine fish.
A. carteri CUNNINGTON, 1931: Hoplias malabaricus: Paraguay & Brazil. (Fig. 9-4
A-C). Carapace covers first two legs and base of third; Basal lobe of
female 4th leg pointed; Abdomen slightly longer than wide; Body = 4.0
long; Carapace = 2.8 x 2.1.
A. chromidis KRǾYER, 1863: Chromis sp. and Rhamdia sp.: Mexico (Yucatan).
(Fig. 9-12). Carapace covers first three legs; Lobes separated from thorax;
Body = 6.0 long; Carapace = 4.1 x 4.0; Abdomen = 1.2 x 1.1.
A. cubensis WILSON, 1935: Cichlasoma tetracanthus: Cuba. (Fig. 9-8 A-C). Cara-
pace covers first three legs; Base of second maxilla with rounded teeth.
A. elongatus HELLER, 1857: host unknown: Brazil (Fig. 9-9). Carapace short,
covering only first pairs of legs; Posterior sinus lacking; Thorax elongate;
Caudal sinus deep; Body = 10 long; Carapace = 4.3; Abdomen = 2.0 x 1.9.
A. ernsti WEIBEZAHN & COBO, 1964: Carassius auratus: Venezuela. (Fig. 9-20).
Carapace covers first three legs; Larger respiratory areas deeply indented
medially; Preoral stylet well developed; Body = 4.3-4.6 (male); 3.2-6.9
(female).
A. ichesi BOUVIER, 1910: host unknown: Argentina (Fig. 9-11). Carapace covers
first two pairs of legs; Respiratory areas confluent; Thorax spotted
dorsally; Abdomen longer than wide; sides nearly parallel; Body = 4.5
(male).
A. juparanaensis LEMOS DE CASTRO, 1950: Pachyurus bonariensis and Astyanax
bimaculatus: Brazil (Espírito Santo State). (Figs. 9-2 A-D & 9-17). Carapace
covers all legs in most adult specimens; Respiratory areas confluent;
Maxillary teeth spatulate; Carapace with brown dendriform markings;
Thorax brown-spotted dorsally; Body = 2.0-2.7 (male); 3.2-4.7 (female);

Carapace = 1.4-2.0 x 1.1-1.7 (male); 2.8-3.9 x 2.2-3.5 (female); Abdomen =
0.27-0.72 x 0.30-0.72 (male); 0.50-0.75 x 0.50-0.84 (female).
A. multicolor STEKHOVEN, 1937: Rhaphiodon vulpinus, Colossoma macropomum,
Serrasalmus nattereri, Cichla temensis, Geophagus jurupari and Hydrolycus scombe-
roides: Brazil & Venezuela. (Fig. 9-14). Carapace covers first two pairs of
legs; Length of abdomen nearly double the width; Body = 4.0 (male); 10.0
(female).
A. nattereri HELLER, 1857: Salminus brevidens and Pseudoplatystoma sp.: Brazil &
Argentina. (Figs. 9-5 & 9-16). Carapace nearly circular, covers all legs;
Abdomen short; Body = 5.5 long; Carapace = 4.6 x 4.56 (male).
A. paranaensis RINGUELET, 1943: Salminus maxillosus: Brazil. (Fig. 9-22). Cara-
pace elongate, lobes slender, covers first three legs; Abdominal rami long
and slender; Two pairs of post-maxillary teeth present; Body = 26.5;
Carapace = 14.5 x 9 (female); male unknown.
A. patagonicus RINGUELET, 1943: Percichthys trucha: Argentina. (Fig. 9-13). Cara-
pace short, covering one or two pairs of legs; Abdomen elongate in
female, shorter in male; Abdominal sinus short; Body = 2.5-3.7 (male); 2.5-
3.0 (female); Carapace = 1.6-2.3 x 1.5-2.0 (male); 1.6-1.9 x 1.4-1.7 (female).
A. paulensis WILSON, 1924: Salminus maxillosus and S. hilarii: Brazil. Description
unavailable.
A. pestifer RINGUELET, 1948: Salminus maxillosus, Pseudoplatystoma fasciatum and P.
tigrinum: Argentina & Brazil. (Fig. 9-30). Carapace with stout supporting
rods anteriorly, covers first three pairs of legs; Respiratory areas contigu-
ous, smaller area medial to larger one; Body = 3.8-10.3; Carapace = 3.1-8.7
x 2.6-7.9.
A. rhamdiae WILSON, 1936: Rhamdia guatemalensis: Mexico (Yucatan). (Fig. 9-3
A-B). Carapace tapering anteriorly, covers all legs; Abdomen subcircular;
Sinus shallow; Body = 3.0; Carapace = 2.5 x 1.7.
A. salmini KRǾYER, 1863: Salminus brevidens and S. maxillosus: Argentina &
Brazil. (Figs. 9-10 & 9-18). Carapace wider than long; Posterior sinus wide;
Carapace covers first three legs and base of fourth; Abdomen small; Sinus
narrow; Body = 13; Carapace = 10.4 x 10.6; Abdomen = 2.5 x 2.6.
A. silvestrii LAHILLE, 1926: Pseudoplatystoma coruscans: Argentina. (Fig. 9-6).
Carapace nearly circular; Posterior sinus wide; Abdomen small; Sinus
narrow; Body = 6.2; Carapace = 6.0 x 5.2.
A. spinulosus SILVA, 1980: Hoplias malabaricus: Brazil (Rio Grande do Sul State).
(Fig. 9-7 A-D). Carapace short, covering only first pair of legs and base of
second; Posterior sinus wide; Abdomen long; Sinus narrow; Body = 3.3
(male); 4.3 (female); Carapace = 2.1 x 1.4 (male); 2.9 x 2.4 (female);
Abdomen = 0.79 x 0.26 (male); 0.79 x 0.66 (female).
A. vierai PEREIRA FONSECA, 1939: Cnesterodon decemmaculatus: Uruguay. This
form is herein considered as a “species inquerenda”. The drawings suggest
that the material was immature and less than one millimeter in length.
A. violaceus THOMSEN, 1925: Rhamdia quelen, R. sapo, Loricaria anus, Hypostomus

commersoni, Hoplias malabaricus, Odonthestes bonariensis and Pimelodus albicans:
Argentina. (Figs. 9-1 & 9-19). Carapace covers first two pairs of legs; Sinus
wide; Abdomen elongate, with parallel sides; Sinus narrow; Body = 6.2-7.2
(male); Carapace = 4.5-5.3 x 3.8-4.2 (male).
Dolops AUDOUIN, 1857
Two pairs of antennae present; first antennae without hooks on anterior
margin, but with lateral hooks strongly developed. Second antennae long,
slender, 3 or 4-segmented. First maxillae swollen at base, tipped with stout
claws. Basal plates of second maxillae armed with 2 or 3 teeth each.
Preoral stylet absent. Four pairs of legs present, covered by carapace, with
flagella on first two or three pairs. Fourth pair of legs with wide basal
lobes. On freshwater fishes.
D. bidenta (BOUVIER, 1899): Schizodon fasciatum, Rhytiodus microlepis, Colossoma
bidens, Serrasalmus nattereri and Astronotus ocellatus: Surinam & Brazil (Ama-
zonia). Carapace and abdomen extensively spotted with dark dendriform
markings; Ventral surface spinous; Second maxillae with two teeth on each
basal plate; Body = 2.3-2.8 (male); 2.4-3.2 (female); Carapace = 1.8-2.3 x
2.0-2.3 (male); 1.9-2.6 x 2.0-2.6 (female); Abdomen = 0.56-0.71 x 0.56-0.71
(male); 0.56 x 0.56-0.71 (female).
D. carvalhoi LEMOS DE CASTRO, 1949: Rhaphiodon vulpinus, Pseudoplatystoma
fasciatum, P. tigrinum, Phractocephalus hemiliopterus, Colossoma macropomum, C.
bidens and Pellona castelnaeana: Brazil. Carapace and abdomen covered with
small dark spots; Marginal dark lines present on carapace; Ventral surface
un-spined; Body = 6.9-7.3 (male); 5.0-6.0 (female); Carapace = 5.0-6.9 x
5.3-6.7 (male); 3.5-4.9 x 3.4-4.9 (female); Abdomen = 2.0-2.1 x 1.7-2.4
(male); 1.4-1.6 x 1.4-1.6 (female).
D. discoidalis BOUVIER, 1899: Salminus maxillosus, Hoplias malabaricus, Pseudo-
platystoma fasciatum, P. tigrinum, Hemisorubim sp., Phractocephalus hemilopterus,
Leiarius marmoratus, Arapaima gigas, Hoplerythrinus unitaeniatus and Astronotus
ocellatus: Argentina, Paraguay, French Guiana, Venezuela & Brazil. Cara-
pace and abdomen covered with round whitish blotches; Thorax with
rectangular dark markings; Ventral spination on margins of cephalic lobes
continuous to near posterior border; Body = 9.9-11.9 (male); 8.0-3.0
(female); Carapace = 7.7-9.3 x 9.0-10.0 (male); 7.0-12.0 (female); Abdo-
men = 3.0-3.9 x 3.9-4.9 (male); 2.0-3.3 x 2.4-4.3 (female).
D. geayi BOUVIER, 1897: Hoplias malabaricus, Aequidens pulcher, Salminus
maxillosus, Crenicichia geayi, Astronotus ocellatus and Megalodoras sp.: Argentina,
Paraguay, Brazil & Venezuela. Carapace and abdomen covered with small
dark blotches; Ventral spination lacking; Body = 2.5-4.0 (male); 4.3-5.7
(female); Carapace = 2.3-2.6 x 1.6-2.4 (male); 2.6-3.9 x 1.7-3.6 (female);
Abdomen = 1.0-1.6 x 0.71-0.86 (male); 1.3 x 2.1-0.71-1.4 (female).
D. kollari HELLER, 1857: host unknown: Brazil. (Fig. 9-26). Carapace
completely covers legs; Sinus wide; Thorax with rectangular dark mark-
ings; Abdomen large; Sinus shallow; Body = 12; Carapace = 10 x 9;

Abdomen = 3 x 3.
D. longicauda HELLER, 1857: Salminus maxillosus, S. brevidens, Serrasalmus nattereri,
Pterodoras granulosus and Oxydoras niger: Brazil & Argentina. (Fig. 9-25 A-B).
Carapace covered with small dark spots; Sinus wide; Abdomen usually
longer than carapace; Sinus wide, V-shaped; Three pairs of respiratory
areas present; Body = 16.7-27.5 (male); 11-27.5 (female); Carapace = 8.1-
14.0 x 8.2-15.0 (male); 7.8-14.7 x 8.2-16.5 (female); Abdomen = 3.2-6.0
(male); 3.3-5.4 (female).
D. nana LEMOS DE CASTRO, 1950: Salminus sp.: Brazil. (Fig. 9-23 A-D). Carapace
incompletely covers 4th legs; Ventral surface without spines; Sinus wide;
Abdominal lobes widely separated; Body = 4.7; Carapace = 3.4 x 3.7;
Abdomen = 1.5 x 1.5 (single male specimen described; female unknown).
D. reperta BOUVIER, 1899: Hoplias malabaricus: French Guiana. (Fig. 9-24, 9-28
& 9-29). Carapace nearly circular; Sinus wide; Abdomen short, wide; Sinus
shallow; Body = 7 long.
D. striata BOUVIER, 1899: Hoplias malabaricus, Synbranchus marmoratus, Schizodon
fasciatum and Leporinus fasciatus: Brazil, Argentina, Paraguay & Surinam.
Species similar to D. discoidalis, but marginal ventral spination terminates at
level of first leg; Body = 7.3-11.1 (male); 7.4-11.6 (female); Carapace =
5.8-8.6 x 6.2-9.4 (male); 5.8-9.1 x 6.4-10.4 (female); Abdomen = 2.3-3.3 x
2.1-4.7 (male); 1.7-3.4 x 2.8-4.4 (female).
Dipteropeltis CALMAN, 1912
Two pairs of antennae present; mesial spine lacking. Preoral stylet small.
First maxillae develop into sucker-like cups, but lack marginal supporting
rods. Carapace without spines ventrally; maxillary and post-maxillary teeth
lacking. Legs without flagella. Anal papillae lacking. Lobes of carpace and
abdomen long, slender and delicate. On South American freshwater fish.
D. hirundo CALMAN, 1912: Serrasalmus piraya, Salminus maxillosus, S. brevidens and
Luciopimelodus pati: Argentina, Brazil & Venezuela. (Fig. 9-27, 9-31 A-B).
Carapace elongate; Lobes long and slender, sometimes extending beyond
tips of abdominal lobes; Sinus deep and wide; Abdominal lobes long and
slender; Sinus deep, V-shaped; Body = 11-26; Carapace = 6-26 x 9-14;
Abdominal lobes = 4.0-6.8 x 1.3-1.5.
VIII. Plates of Branchiura (Figs. 9-1 to 9-31)
9-1
A-1
A-2
ME
PS
Mx-1
Mx-2
MO
RA
MT
EN
EX
FL
SR
AP
9-1. Argulus violaceus THOMSEN, 1925 (female, ventral aspect; modified, after RINGUELET 1943): A-1 =
first antenna; A-2 = second antenna; AP = abdominal papilla; EN = endopod; EX = exopod; FL =
flagellum; ME = mesial spine; MO = mouth; MT = maxillary teeth; MX -1 = first maxilla; MX-2 =
second maxilla; PS = preoral stylet; RA = respiratory areas; SR = seminal receptacle.
9-2 A
9-3 A
9-2. Argulus paranaensis (male): A. dorsal aspect; B. antennae; C. sucker support rod; D. second maxilla
(after LEMOS DE CASTRO 1950); 9-3. Argulus rhamdiae (female): A. dorsal aspect; B. second maxilla
(after WILSON 1936).
9-4 A
D
C
9-5
9-6
9-4. Argulus carteri (female): A. abdomen; B. antennae & mesial spine; C. sucker support rods; D.
second maxilla & postmaxillary spines (after CUNNINGTON 1931); 9-5. Argulus nattereri (female; dorsal
aspect) (after WILSON 1944); 9-6. Argulus silvestrii (female; dorsal aspect) (after RINGUELET 1943).
9-7 A
9-8 A C
9-7. Argulus spinulosus (male): A. dorsal aspect; B. antennae; C. second maxilla; D. respiratory areas.
(after SILVA 1978); 9-8. Argulus cubensis (male): A. dorsal aspect; B. antennae; C. second maxilla (after
WILSON 1935).
9-9
9-10
9-11 9-12
9-9. Argulus elongatus (female; dorsal aspect) (after WILSON 1903); 9-10. Argulus salmini (female; dorsal
aspect) (after WILSON 1903); 9-11. Argulus ichesi (female; dorsal aspect) (after RINGUELET 1943); 9-12.
Argulus chromidis (female; dorsal aspect) (after WILSON 1903).
9-13
9-15
9-14
9-16 9-17 9-18
9-19 9-20 9-21 9-22
Respiratory areas: 9-13. Argulus patagonicus; 9-14. Argulus multicolor; 9-15. Dolops discoidalis; 9-16. Argulus
nattereri; 9-17. Argulus juparanaensis; 9-18. Argulus salmini; 9-19. Argulus violaceus; 9-20. Argulus ernsti; 9-
21. Dipteropeltis hirundo; 9-22. Argulus paranensis.
9-23
C
D
9-24
9-23. Dolops nana (male): A. dorsal aspect; B. antennae; C. second maxilla; D. respiratory area (after
WILSON 1902). 9-24. Dolops reperta (female; dorsal aspect) (after WILSON 1903).
9-26
9-25 A
9-25 B 9-27
9-25. Dolops longicauda (male): A. dorsal aspect; B. respiratory areas (after WILSON 1903); 9-26. Dolops
kollari (male; dorsal aspect) (after WILSON 1903); 9-27. Dipteropeltis hirundo (female; ventral aspect)
(after PAIVA CARVALHO 1941).
9-28
9-28. Dolops reperta (female; dorsal aspect).

9-29
9-29. Dolops reperta (female; ventral aspect).

9-30
9-30. Argulus pestifer (female; ventral aspect).

9-31 A 9-31 B
9-31. Dipteropeltis hirundo: A. female; dorsal aspect; B. ventral aspect.

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ecologia de Argulus multicolor STEKHOVEN, 1937, e Argulus pestifer RINGUELET, 1948. - Acta
Amazonica 13(3-4): 489-495.
MALTA, J.C.O. (1988): Maxillopoda. Branchiura. - In: YOUNG, P.S. (ed.): Catalogue of Crustacea of
Brazil: 67-74. Série Livros n. 6. Museu Nacional, Rio de Janeiro.
MALTA, J.C.O. & A. VARELLA (1983): Os argulídeos (Crustacea, Branchiura) da Amazonia brasileira.
3. Aspectos da ecologia de Dolops striata BOUVIER, 1899, e Dolops carvalhoi CASTRO, 1949. - Acta
Amazonica 13(2): 299-306.
MALTA, J.C.O. & A. VARELLA (2000): Argulus chicomendesi sp. n. (Crustacea: Argulidae) parasita de
peixes da Amazônia Brasileira. - Acta Amazônica 30(1): 481-498.
MARTINEZ, R. (1952): Argulus chilensis nov. sp. - Invest. Zool. Chilenas 1(7): 4-9, figs. 1-5.
MELLO LEITÃO, C.F. (1913): Moreirella nome nove para Talaus MOREIRA- Dipteropeltis CALMAN. - A
Lavoura (Rio de Janeiro) 17: 11-12.
MOREIRA, C. (1912): Crustacés du Brésil. - Mem. Soc. Zool. France 25: 145-154, pls. 3-6.
MOREIRA, C. (1915): Les antennas de Dipteropeltis hirundo CALMAN (Talaus riberoi MOREIRA) (Custacea,
Argulidae). - Bull. Soc. Entom. France 25(3-4): 120-122, 2 figs.
PAIVA CARVALHO, J. (1939): Sôbre dois parasitos do gênero Dolops encontrados em peixes d’água
doce. - Rev. Indust. Animal (ser. 1) 2(4): 109-116, figs. 1-6.
PAIVA CARVALHO, J. (1941): Sôbre Dipteropeltis hirundo CALMAN, crustáceo (Branchiura) parasito de
peixes d’água doce. - Bol. Fac. Filos. Cienc. Univ. São Paulo 22(5): 265-275.
PAIVA CARVALHO, J. (1947): Sôbre alguns ectoparasitos de peixes (Argulídeos). - Criação e Veterinaria
2(6): 155-156.
RINGUELET, R. (1941): Sôbre algunos crustáceos argúlidos argentinos. - Physis 19(51): 112.
RINGUELET, R. (1942): Sobre dos espécies de argulidos (Crustacea, Branchiura) en las colecciones
del Museu de Entre Rios. - Mem. Mus. Entre Ríos 16: 3-14.
RINGUELET, R. (1943): Revisión de los argúlidos argentinos (Crustacea, Branchiura) con el catálogo
de las especies neotropicales. - Rev. Mus. La Plata (n.s.) Zool. 3(19): 43-99, pls. 1-13.
RINGUELET, R. (1948): Argúlidos del Museo de La Plata. - Rev. Mus. La Plata (n.s.) Zool. 5: 231-296,
pls. 1-4.
SCHUURMANS STEKHOVEN, J.H., JR. (1951): Investigaciones sobre argúlidos argentinos. - Acta Zool.
Lilloana 12: 479-494.
SILVA, N.M.M. (1978): Uma nova espécie de crustáceo argulídeo do Rio Grande do Sul, Brasil
(Branchiura, Argulidae). - Iheringia, sér. Zool. 52: 3-29.
THOMSEN, R. (1925): Argulus violaceus nov. sp. cangrejo parásito del bagre. - Physis 8: 185-198.
THOMSEN, R. (1942): Notas críticas de dos argúlidos (Branchiura) del Brasil. - An. Acad. Brasil.
Cienc. 14: 37-45.
WEIBEZAHN, F.A. & T. COBO (1964): Seis argúlidos (Crustacea, Branchiura) parásitos de peces
dulceacuícolas en Venezuela, con descripción de una nueva especie del género Argulus. - Acta
Biológica Venez. 4(2): 119-143, pls. 1-15.
WILSON, C.B. (1903): North American parasitic copepods of the family Argulidae, with a
bibliography of the group and a systematic review of all known species. Proc. U. S. Nat. Mus.
25: 635-742.
WILSON, C.B. (1935): Parasitic copepods from the Dry Tortugas. - Carnegie Inst. Washington. Publ.
452: 329-347.
WILSON, C.B. (1936): Copepods from the cenotes and caves of the Yucatan Peninsula, with notes
on the cladocerans. - Carnegie Inst. Washington. Publ. 457: 77-80.
WILSON, C.B. (1944): Parasitic copepods in the United States National Museum. - Proc. U. S.
Nat.Mus. 94(3): 529-582, pls. 20-34.
YAMAGUTI, S. (1963): Parasitic Copepoda and Branchiura of fishes. - Intersci. Publ., New York:
1104 pp., 333 pls.
10.
ISOPODA
Isopods are dorso-ventrally flattened crustaceans found in marine, terrestrial and

freshwater habitats. The name isopod means “equal legs” and most species have a free
thorax (pereon) of seven similar segments (pereomeres) each of which bears a pair of
similar legs (pereopods). The most familiar isopods are commonly known as “sow bugs”
or “pill bugs” and these are found in damp terrestrial habitats. Most of the fish parasitic
isopods belong to the family Cymothoidae, but the family Excorallanidae also occurs on
Amazonian fish. The parasitic isopods have from four to seven pairs of walking legs
modified for grasping and each of these is provided with a stout claw.
Most free living isopods are bottom feeding aquatic scavengers. Some eat
detritus and carrion, while others are more predatory. Many have one or two pairs of
prehensile legs anteriorly to aid in grasping food. Those that evolved more pairs of
prehensile legs and stronger claws were able to cling to fully active fish and make the
transition to parasitism.
In isopods, the sexes are separate and fertilization is internal. Both male and female
cymothoids are found on fish, frequently together. It is believed that all cymothoids go
through a male stage first and then some go on to become females (which would make
them protandrous hermaphrodites). According to this idea, the first young cymothoid to
arrive on a fish, and finding itself alone, would pass rapidly through the male stage and
become a female. The second to arrive would find a female in residence and hence remain
a male. This could be caused by hormonal inhibitors. In any case, adult females are easy to
recognize because they have the entire ventral surface covered by a marsupium formed
from medial flap-like extensions of the coxae. The eggs are laid in and retained within the
marsupium. There they develop into fully functional young cymothoids and eventually go
out into the world through a slit-like posterior pore.
Young cymothoids swim actively and are ready to attack small fish as soon as they
leave the marsupium. In Amazonian waters, it is not uncommon to find small
cymothoids clinging to and feeding on small fish. In the laboratory, we have been able
to observe the hunting behavior of young cymothoids. At first, they swim rapidly and in
very straight horizontal lines. Their swimming posture is with the head forward to the
direction of movement and the dorsal side up. The swimming activity lasts for only 30
to 60 seconds after which they suddenly become immobile and sink to the bottom as
though dead, with the dorsal side down and the legs pointed upward and spread wide
apart. They are capable of waiting motionless in this position for hours. If a fish goes to
investigate, or happens to swim by closely, the apparently dead cymothoid springs its
trap. The legs are all snapped inward towards the midline, and the claws (dactyls) are
engaged in the fish which then swims off with a passenger.
After a young cymothoid has attached itself to a small fish, it begins to feed at
once. They frequently position themselves on the ventral surface of the fish at the level
of the pectoral fins. Wherever they happen to be, they rasp out tissue and frequently,
the pectoral fins are entirely devoured.
The under sides of the opercula are also eaten, and if the fish is small, it soon
succumbs to such an attack. When a fish dies, the young cymothoid loses interest in it
and swims off in search of a live fish. We have seen a single young cymothoid kill and
devour portions of as many as four small fish in a 24 hour period. After young
cymothoids grow larger, they must find a permanent host for they lose their swimming
ability. Adult males detached from the fish can crawl slowly on the bottom, but females
with a marsupium full of young can not move about.
III. Pathology
The pathogenicity of isopods varies according to position on or in the host, feeding

behavior, attachment strategy and size. Four sites are utilized on host fish, namely: the skin
or fins, the gill chambers, the mouth (on top of the tongue) and hollows or pouches that
they make within the body cavity. Each of these sites suffers some modifications from the
presence of isopods so they will be considered separately.
Of the South American species reported from freshwater hosts, only Nerocila armata
is normally found on the skin. These ectoparasites cause considerable damage to their
hosts and are believed to feed on blood, mucous and epithelial cells. They sometimes
perforate the skin of the fish or devour portions of the fins. They may cause primary
anemia from blood loss, and the lesions they provoke may open the way for secondary
invaders such as bacteria and fungi.
Isopods that inhabit the gill chambers of fish feed largely on gill filaments. BRUSCA
(1978) reported lysed red blood cells in the intestines of males of a marine species taken
from such a site. In addition to blood loss, a reduction in the number of gill filaments
decreases respiratory efficiency and thereby affects metabolism and growth. ROMESTAND
& TRILLES (1979) in experimental studies with marine fish infested with isopods found
that the parasites caused delayed growth in the hosts.
The role of isopods living on the tongues of fish is more difficult to assess.
Although the tongues of hosts may be reduced in size or indented, there is no direct
evidence to indicate that the missing tissue has been eaten. ROMESTAND & TRILLES
(1977) found as much as half the tongues of infected hosts to be lacking. The missing
tissue included epidermis, cartilage and some bones. They considered these tongues to
be atrophied or degenerated.
The nutritional habits of tongue inhabiting isopods have been questioned. BRUSCA
(1978) found the intestinal tracts of cymothoid females from tongue sites to be entirely
empty. The same author (1981) suggested, “that females are largely (or perhaps entirely)
non-feeding and hence are best considered obligate commensals”. He further proposed
that these females use nutritional reserves from previous male instars for reproductive
energy. This would appear to be an incredible suggestion, but an even more unusual idea
was put forward by BRUSCA & GILLIGAN (1983). They believed that isopods in the mouths
of fish serve as “tongue replacements”. According to this theory, a fish with an “isopod
tongue” feeds more efficiently than a fish with no tongue at all.
THATCHER (1988) described Asotana magnifica from the mouths of piranhas of the
Brazilian Amazon and made some observations on the habits and pathogenicity of
these cymothoids. He reported one 34 mm isopod on the tongue of a host that was
only 195 mm long (Fig. 10-48). The fish appeared to have been in good health and
had a full stomach. The surface of the tongue of the host was depressed in the exact
shape of the ventral side of the isopod. This indentation in the floor of the mouth
was attributed to pressure atrophy. As shown in Figure 10-48, the head of these
isopods is close to the entrance of the esophagus of the host. The anterior ends of
isopods taken from such sites are frequently found to be covered with an amorphous
and acellular substance. It seems logical to assume that this material represents
regurgitated semi-digested food from the host stomach that is only a few millimeters
from the isopod’s head. We can conclude from this that the real food of the isopods
is regurgitated host food (i.e. vomitus). The term obligate commensal would still apply
in this case. Apparently, Vanamea symmetrica and some species of Braga also utilize the
same feeding strategy.
In freshwater hosts of South America, there are five species of Riggia and two of
Artystone all of which penetrate host tissue and are therefore much more pathogenic.
The mouthparts of these forms are adapted for burrowing and mandible and maxillae
all have hook-like spines pointing outward. Entrance holes are made either at the site of
a missing pectoral fin (which the isopod probably removes) or near the anus. These
entrance holes remain open and the pleotelson can be seen to protrude slightly.
Presumably this position permits the isopod to breathe, an act that is affected by
beating the pleopods in the water. The body of the isopod remains within the body
cavity of the host, frequently in the vicinity of the liver. There may be pressure atrophy
of this or other organs but there is no evident inflammatory reaction or encapsulation
of the parasite on the part of the host. There is also no evidence of secondary invasions
of the open entrance hole. Perhaps the isopods secrete some antibiotic substance that
inhibits secondary invaders. Isopods in this site may be feeding on blood serum
although direct evidence of this is lacking.
Isopods do not ordinarily become a problem to the alert aquarist. Since even young
specimens are large enough to be seen, they can simply be removed. Some chemical dips,
such as a 1:4000 formalin solution, will encourage the isopods to leave their hosts. If
isopods become too numerous in outdoor ponds, the only recourse is to drain and
disinfect them.
When isopods are seen in or on fish, they and their host should be fixed in 10 % formalin
solution. After 24 hours, both may be transferred to 70 % alcohol. The mouthparts, legs,
uropods, pleopods and pleotelson may be removed from isopods with dissecting needles
and permanent slides can be made of them by using the phenol – balsam method
described in Chapter 6.
Neotropical fish parasitic isopods have been little studied. The genera and species are not
well defined and several of these taxa were based on single female specimens. Many new
forms no doubt await future descriptions. Some characters that are useful in distinguish-
ing these animals are: size, shape, color, number of antennal segments and the form of
the cephalon, pleopods, uropods and pleotelson.
South America contains a greater variety of freshwater Cymothoidae than any other
region of the world. At least 27 species, representing nine genera, occur here. For
comparison, there are no freshwater cymothoids in either North America or Europe and
only four species known from Africa. According to BRUSCA (1981), there are some 250
species of cymothoids in 42 genera world-wide and most of these are marine.
Key to the Genera of Cymothoidae from Neotropical freshwater fishes
420
I. Coxal plates on pereonite I free .......................................................................................Anphira (Figs. 10-16, 10-36, 10-47, 10-49 & 10-57)

II. First coxal plates fused to pereonite I.
A. Legs I-VI prehensile and ending in claw-like dactyls; legs of pair VII ambulatory and ending in simple dactyls.
1. Abdomen (pleon) visibly segmented and separate from pleotelson ..................................... Artystone (Figs. 10-1 to 10-6 & 10-46)
2. Pleon and pleotelson fused into a single unit .................................. Riggia (Figs. 10-44 to 10-46, 10-51, 10-52 & 10-59 to 10-60)
B. All 7 pairs of legs prehensile and ending in claw-like dactyls.
1. Anterior margin of pereonite one trisinuate; posterior margins of pereonites II-VII pointed and projecting ..............................
............................................................................................................................................................................................. Nerocila (Fig. 10-58)
2. Anterior margin of pereonite one not trisinuate; osterior margins of pereonites II-VII not pointed and projecting.
a. Anterior margin of cephalon provided with 3 rounded bosses ................................................... Asotana (Figs. 10-37 & 10-38)
b. Anterior margin of cephalon otherwise.
1) Pleopods II-V composed of from 3-4 lamina .................................................................................................................................
................................................................. Braga (Figs. 10-13, 10-14, 10-22, 10-25 to 10-29, 10-31, 10-39 to 10-43 & 10-50)
2) All pleopods bilaminate.
a) Pleotelson narrower than pleonite one.
(1) Body wide anteriorly, tapering posteriorly.
(a) Pleon deeply immersed in pereonite VII ........................................................................................................ Isonebula
(b) Pleon not deeply immersed in pereonite VII .......................................................... Livoneca (Figs. 10-7 to 10-9)
(2) Body with nearly parallel sides ......................................................................................................... Vanamea (Fig. 10-10)
(3) Body tapering anteriorly and posteriorly .................................................................................Phillostomella (Fig. 10-24)
b) Pleotelson wider than pleonite I ....................................................................... Paracymothoa (Figs. 10-11, 10-12 & 10-21)
c) Pleotelson about as wide as pleonite I .......................................................... Telotha (Figs. 10-15, 10-23, 10-61 & 10-62)
Clave para los géneros de Cymothoidae de peces Neotropicales de agua dulce
I. Placas coxales sobre el pereonito I libre .........................................................................Anphira (Figs. 10-16, 10-36, 10-47, 10-49 & 10-57)
II. Primeras placas coxales fundidas al pereonito I.
A. Piernas I-VI prehensiles y terminando en dáctilos parecidos a garras; piernas del par VII ambulatorios y terminando en dáctilos
simples.
1. Abdomen (pleon) visiblemente segmentado y separado del pleotelsón ................................Artystone (Figs. 10-1 a 10-6 & 10-46)
2. Pleón y pleotelsón fusionados en una unidad .............................................................. Riggia (Figs. 10-44 a 10-46 & 10-59 a 10-60)
B. Los 7 pares de piernas prensiles y terminando en dáctilos parecidos a garras.
1. Margen anterior del pereonito uno con 3 sinuosidades; márgenes posteriores de los pereonitos II-VII puntiagudos y
proyectando ....................................................................................................................................................................... Nerocila (Fig. 10-58)
2. Margen anterior del pereonito uno no sinuoso; márgenes posteriores de los pereonitos II-VII sin proyecciones.
a. Margen anterior del cefalón provistos de 3 jorobas redondeadas ................................................ Asotana (Figs. 10-37 & 10-38)
b. Margen anterior del cefalón sin jorobas.
1) Pleopodos II-V compuestos de 3-4 láminas .....................................................................................................................................

..................................................................... Braga (Figs. 10-13, 10-14, 10-22, 10-25 a 10-29, 10-31, 10-39 a 10-43 & 10-50)
2) Todos los pleopodos bilaminados.
a) Pleotelsón más angosto que el pleonito uno.
(1) Cuerpo ancho anteriormente, angostándose posteriormente.
(a) Pleón profundamente inmerso en el pereonito VII ..................................................................................... Isonebula
(b) Pleón no profundamente inmerso en el pereonito VII ........................................... Livoneca (Figs. 10-7 a 10-9)
(2) Cuerpo con lados casi paralelos ...................................................................................................... Vanamea (Fig. 10-10)
(3) Cuerpo se angosta anterior y posteriormente ........................................................................Phillostomella (Fig. 10-24)
b) Pleotelsón más ancho que el pleonito I .......................................................... Paracymothoa (Figs. 10-11, .10-12 & 10-21)
c) Pleotelsón casi tan ancho como el pleonito I ............................................. Telotha (Figs. 10-15, 10-23, 10-61 & 10-62)
421
VII. Checklist of Isopods from Neotropical freshwater fishes
All measurements given in mm.
Cymothoidae SCHIÖDTE, 1866

Flabelllifera. Body symmetrical or asymmetrical; eyes prominent, com-
pound, sessile. Legs (pereopods); all 7 pairs prehensile, or anterior 6
prehensile and terminal pair ambulatory; prehensile legs terminating in
stout claw-like dactyls. Pleopods not ciliated; pleotelson and uropods
usually not ciliated. Antennae reduced to 4-10 segments, without clear
distinction between peduncle and flagellum. Mouthparts: mandibles with
palps; first maxillae spined at tip; second maxillae bilobed at tip; maxilli-
peds provided with 2-segmented palps that have hooks on terminal
segments. Parasitic on skin, in mouth, branchial chamber or within self-
made cavities in fish.
Anphira THATCHER, 1993
Diagnosis: Body ovoid, dorsum of pereon highly convex. Coxae plate-like,
free on all seven pereonites. Cephalon immersed in pereonite I and rotated
downward. Pleonite I slightly immersed in pereonite VII; all pleonites
produced laterally. Mandible rounded, foot-shaped, lacking cutting process
and incisor; palp stout. Pereopods short, II-VI subequal in length, I
shorter and VII longer than others. Uropods slender, rami pointed,
subequal in length. Pleopods with rounded extremities. Pleotelson not
keeled, rounded posteriorly. Parasites of the gill chambers of freshwater
fishes. Type species: Anphira branchialis THATCHER, 1993.
A. branchialis THATCHER, 1993: Serrasalmus spilopleura and S. nattereri: Amazonas
and Roraima States, Brazil. (Figs. 10-16 & 10-47). Female body = 12-21 x
6.7-11.6; Male body = 9.4-13.99 x 5.0-7.
A. guianensis THATCHER, 2002: Acnodon oligacanthus: French Guiana. (Fig. 10-36).
Female body = 12-20 x 6-9; Male body = 7-8 x 3-4.
A. junki ARAUJO & THATCHER, 2003: Triportheus albus and T. flavus: Amazonas
State, Brazil. (Fig. 10-57). Body asymmetrical, posterior part twisted up to
45 degrees between pereonites 6 and 7; Female = 6.9-9.1 x 3.4-5.1; Male =
5.7-6.9 x 2.8-3.4.
A. xinguensis THATCHER, 1995: Ossubtus xinguensis: Xingu River, Pará State,
Brazil. Female body = 5-16 x 2.5-7.0; Male body = 3.5-12 x 2-5.
Artystone SCHIÖDTE & MEINERT, 1881
Cymothoidae. Body symmetrical; head (cephalon) considerably immersed in
first thoracic segment (pereonite). First 6 pairs of legs provided with claw-
like dactyls; terminal pair ambulatory, without claws. Abdomen (pleon)
segmented. Parasitic in self-made pouches within the body cavities of fish.
A. trysibia SCHIÖDTE, 1866: Geophagus brasiliensis and Crenicichla lacustris: Brazil.
(Figs. 10-20 & 10-56). Body = maximum sizes: 20 x 11 (male); 26 x 18
(female).
A. bolivianensis THATCHER & SCHINDLER, 1999: Otocinclus vestitus: Santa Cruz de

la Sierra, Bolivia. (Figs. 10-1 to 10-4). Female body = 5.2-6.2 x 3.5-4.0;
Male body = 2.5-4.5 x 1.2-1.9.
A. minima THATCHER & CARVALHO, 1988: Nannostomus beckfordi: Upper Rio
Negro, Amazonas State, Brazil. (Figs. 10-5 & 10-6). Body = 3.7-4.0 x 1.9-
2.0 (males); 5.2-6.9 x 2.7-3.7 (females); Cephalon = 0.53-0.68 x 0.98-1.0
(males) & 0.62-0.90 x 1.0-1.3 (females); Pleotelson = 0.65-0.75 x 1.1-1.2
(males) & 1.1-1.3 x 1.4-1.8 (females).
Asostana SCHIÖDTE & MEINERT, 1881
Cymothoidae. Female: body large, convex dorsally, cream to straw-colored
with sparse black pigment spots. Cephalon large, subrectangular, not
deeply immersed in pereonite 1, keeled dorsally and with two pairs of
dorsolateral tubercles; frontal margin curved downward and terminating in
three rounded bosses. Mandibles with dorsal and ventral flaps; maxillae
without spines. Pereonite 1 with brown or black roughened patches of
denticles dorsally. Pereopods short, slender, subequal in length; first 6 pairs
of dactyls larger than 7th . Abdomen inserted at its base. Uropods short
and wide. Pleotelson tongue or shield-shaped. Parasites or commensals in
the buccal cavities of freshwater fish.
A. formosa SCHIÖDTE & MEINERT, 1881: host unknown: Peru (probably
Amazonia). (Fig. 10-17). Body = 30 x 12 (female; male unknown); Body
tapers gradually anteriorly and posteriorly from pereonite 5.
A. magnifica THATCHER, 1988: Serrasalmus sp.: Roraima State, Brazil. (Figs. 10-
33, 10-37 & 10-38). Body with abdomen abruptly narrower than thorax,
measures = 30-40 x 14-15.
A. splendida (LEIGH-SHARPE, 1937): host unknown: Ecuador. Female: body
tapers abruptly anteriorly and posteriorly from pereonites 3 & 4; Measures
= 21 x 11.
Braga SCHIÖDTE & MEINERT, 1881
Cymothoidae. Body symmetrical; cephalon little immersed in pereonite 1.
All 7 pairs of pereopods prehensile and provided with stout claw-like
dactyls. Pleotelson usually wider than long. Parasites, or commensals,
found on the tongue or in the gill chambers of freshwater fishes.
B. amapaensis THATCHER, 1996: Acestrorhynchus guyanensis: Amapa State, Brazil. (Figs.
10-25 to 10-29). Female body = 12-18 x 4-5.5; Male body = 8-13 x 3-4.
B. bachmanni STADLER, 1972: Ancistrus cirrhosus. Argentina. (Fig. 10-14). Female
body = 3 x 1.8.
B. cichlae SCHIÖDTE & MEINERT, 1881: Cynopotamus humeralis, Cichla ocellaris and
C. temensis: Brazil (São Paulo, Minas Gerais, Pará & Amazonas States).
(Figs. 10-39 to 10-41). Body about twice as long as wide: Female = 25
maximum length; Male = 15.
B. fluviatilis RICHARDSON, 1911: unidentified catfish: Argentina, Surinam &
Brazil. Female body = 22 x 14; Dactyl 1 of pereopod 1 longer than the
others. (Fig. 10-13).
B. nasuta SCHIÖDTE & MEINERT, 1881: Hypostomus sp.: Brazil (São Paulo, Bahia
& Amazonas States). (Figs. 10-22 & 10-31). Female body = 20-25 long;
Male = about 11 long.
B. patagonica SCHIÖDTE & MEINERT, 1884: Salminus hilarii and Hoplias malabaricus:
Argentina, Paraguay, Surinam & Brazil (São Paulo, Pernambuco, Bahía, Pará
& Amazonas States). (Figs. 10-42 & 43). Female = to 26; Male = to 15.
Livoneca LEACH, 1818
This genus can not be defined at present. The marine and freshwater
forms now assigned to this taxon do not appear to have many characters in
common and they may represent more than one genus.
L. guianensis VAN NAME, 1925: from gills of Leporinus fasciatus, Pimelodus clarias
and Brachyplatystoma sp.: Guyana. (Fig. 10-7). This species is elongate with a
thorax that tapers gradually from pereonite 4 to the abdomen; The
cephalon has a sharp point anteriorly (as seen from above) and the
pleotelson is subquadrangular; Female = 17.7-26 long; Male = unknown.
L. orinoco BOWMAN & DÍAZ-UNGRÍA, 1957: gills of Cichlidae: Orinoco River,
Venezuela (Fig. 10-8 & 10-9). In this species, the head is not much
immersed in the pereon but the abdomen is; The anterior margin of the
cephalon is rounded; The pleotelson is also rounded posteriorly and is
slightly wider than long; Female = 13.9 x 7.0; Male = 8.5 x 4.5.
Nerocila LEACH, 1818
Cymothoidae. Body elongate, considerably flattened. Anterior margin of
pereonite 1 trisinuate. Postero-lateral margins of pereonites 2-7 pointed
and projecting. Coxal plates of pereonites 2-7 also pointed and projecting
posteriorly. Parasites of skin and fins of marine and freshwater fish.
N. armata DANA, 1853: skin of Leporinus fasciatus, Crenicichla saxatilis, Cichla ocellaris
and Pseudauchenipterus nodosus: Uruguay, Guyana & Brazil (São Paulo & Rio de
Janeiro States). (Fig. 10-58). Female body = 18-24 long; Male = 7.5-18 long.
Paracymothoa LEMOS DE CASTRO, 1955 (emend. BOWMAN, 1986)
Cymothoidae. Body oval. Head not deeply immersed in pereonite 1; anterior
margin broad and almost straight, or convex, not curved ventrally. Antennae
short, antennae 1 widely separated at base. Pereonite 1 with short antero-
lateral angles. Pereonite 7 shorter than others. Pleon deeply immersed in
pereon and narrower than latter. Legs short; 1-3 smaller than 4-6. Dactyl of
leg 7 smaller than others. From the mouths of freshwater fish.
P. astyanactis LEMOS DE CASTRO, 1955: Astyanax bimaculatus: Brazil (Rio de
Janeiro & Minas Gerais States). (Fig. 10-21). Female = 10.5 x 5; Rami of
uropods subequal, reach posterior margin of pleotelson.
P. parva TABERNER, 1976: mouth of Hyphessobrycon callistus: Argentina. (Fig. 10-
11). Female = 4.2 x 2; Rami of uropods subequal, reach posterior margin
of pleotelson.
P. tholoceps BOWMAN, 1986: mouth of Hoplias macrophthalmus: Venezuela
(Amazonas State). (Fig. 10-12). Female = 24.3 x 12.2; Outer ramus of
uropod longer, reaches to near equatorial region of pleotelson.
Philostomella SZIDAT & SCHUBART, 1960

Cymothoidae. Body symmetrical; head not immersed. All legs prehensile
and clawed. First antennae: 8-segmented in female; 7-segmented in male.
Second antennae: 8-segmented in female; 9-segmented in male. Pleotelson
about as wide as long, keeled. Mouth parasites of fish.
P. cigarra SZIDAT & SCHUBART, 1960: Cynopotamus humeralis: Brazil (São Paulo
State). (Fig. 10-24). Body = 29 long (female); 15 long (male).
Riggia SZIDAT, 1948
Cymothoidae. Body symmetrical or asymmetrical; head immersed in first
pereonite. Pereopods 7 without claws. Pleon and pleotelson fused together
to form a single unit. Parasites found within self-made pouches in the body
cavities of fish.
R. acuticauda THATCHER, LOPES & FROEHLICH, 2002: Ancistrus sp.: Mato Grosso
do Sul State, Brazil. (Fig. 10-44). Female = 11-16 x 6-10.
R. brasiliensis SZIDAT & SCHUBART, 1959: Leporinus copelandi, L. octofasciatus,
Schizodon nasutus and Leporellus vitattus: Brazil (São Paulo State). (Fig. 10-18).
R. cryptocularis THATCHER, LOPES & FROEHLICH, 2003: Ancistrus sp. and Odontos-
tilbe sp.: Mato Grosso do Sul State, Brazil. (Figs. 10-45 & 10-46). Female =
20 x 13; male = 4.9 x 2.3.
R. nana SZIDAT & SCHUBART, 1959: Leporinus striatus: Brazil (São Paulo State).
(Fig. 10-19). Body = 14 x 10 (female).
R. paranensis SZIDAT, 1948: Curimata platana: Brazil (Paraná River, Paraná
State). (Figs. 10-59 & 10-60). Body = 22 x 16 (female).
Telotha SCHIÖDTE & MEINERT, 1884
Cymothoidae. Body symmetrical; cephalon triangular, not immersed in
pereonite 1, anterior margin curved sharply downward. All legs prehensile
and clawed. Antennules 8-segmented; antennae 9-segmented; antennae
well separated at bases. Pleotelson wider than long. Parasites of gill cavities
of freshwater fish.
T. henselii (VON MARTENS, 1869): Hoplias malabaricus, Pimelodus clarias, Geophagus
sp. and Brachyplatystoma sp.: Guyana, Surinam, Uruguay, Argentina & Brazil
(Rio Grande do Sul). (Figs. 10-61 & 10-62). Body = 16-26 long (female);
Male unknown.
T. lunaris SCHIÖDTE & MEINERT, 1884: Stenarchus brasiliensis: Brazil. (Fig. 10-
15). Body = 28 long (female); Male unknown.
T. silurii SZIDAT & SCHUBART, 1959: Iheringichthys labrosus: Brazil (São Paulo
State). (Fig. 10-23). Body = 13 long (female); Male unknown.
Vanamea THATCHER, 1993.
Female: body elongated, symmetrical; pereon weakly vaulted, sides nearly
parallel. Cephalon not immersed in pereonite 1. Coxae small, not surpass-
ing posterior margins of respective pereonites. Pleon not immersed in
pereonite 7, only slightly narrower then pereon; pleonites subequal in
length; pleopods laminate, with lateral laminar lobes on peduncles. Anten-
nule shorter than antenna; bases set wide apart. Mandible “foot-shaped”
lacking incisor; maxillule with 3 terminal and 2 subterminal recurved

spines: maxilla bilobed with 2-3 recurved spines on each lobe; maxillipedal
palp with 1 terminal and 1 subterminal recurved spine. Pereopods
increasing in length from 1-7; largest dactyls on pereopods 5-6, smallest on
7. Pleotelson inflated anteriorly and medially. Male similar to female but
smaller. Parasites of the buccal cavity of freshwater fishes.
V. symmetrica (VAN NAME, 1925); THATCHER, 1993: Serrasalmus spilopleura, S.
elongatus, S. rhombeus, Myleus rubripennis, Cichla monoculus, Vandelia cirrhosa,
Hemidoras carinatus, Carnegiella strigata and Brachyplatystoma sp.: Guyana,
Venezuela, Brazil (Pará and Amazonas States). (Fig. 10-10). Body 2.3-3.3
times longer than wide; female 18-26 (21) long and 6-11 wide. Male 14-18
(16) long and 5-5.5 (5.6) wide.
Excorallanidae STEBBING, 1904
Flabellifera. Body symmetrical, with nearly parallel sides. Coxal plates
present on all pereonites but the first. Anterior 3 pairs of pereopods
prehensile, posterior 4 pairs ambulatory. Cephalon little immersed, with
conspicuous horn-like processes in male (which are lacking or smaller in
female); eyes prominent. Antennae long, with 2-4 segments in peduncle
and 10-30 in flagellum. Mouthparts: mandible with large curved apical
tooth; mandibular palp 3-segmented, with terminal segment expanded;
first maxilla robust; second maxilla short; maxilliped narrow, free. Uropods
long: pleotelson reduced, subtriangular. Free-living or ectoparasitic on fish.
Excorallana STEBBING, 1904
Excorallanidae, with the characters of the family.
E. berbicensis BOONE, 1918: skin of Lycengraulis grossidens: Guyana. Body = 7.8
(female); 6 (male).
Excorallana sp.: from skin of Ageneiosus brevifilis: Brazil (Pará & Amazonas
States). (Figs. 10-53 to 10-55).
VIII. Plates of Isopoda (Figs. 10-1 to 10-62)
10-1 10-2
10-3
10-4
10-5 10-6
Artystone bolivianensis THATCHER & SCHINDLER, 1999: 10-1. male; 10-2. female; 10-3. male uropod;
10-4. female uropod; Artystone minimia THATCHER & CARVALHO, 1988: 10-5. male; 10-6. female; all
scales = 1 mm.
10-7 10-8 10-9
10-10 10-11 10-12

10-7. Livoneca guianensis; scale = 5mm; 10-8. Livoneca orinoco: female (lateral); scale = 5 mm; 10-9.
Livoneca orinoco: female (dorsal); scale = 5mm; 10-10. Vanamea symmetrica: female; scale = 5 mm; 10-11.
Pracymothoa parva: female; scale = 1mm; 10-12. Paracymothoa tholoceps: female; scale = 5 mm.
10-13 10-14
10-15 10-16
10-13. Braga fluviatilis: female; 10-14. Braga bachmanni: female; 10-15. Telotha lunaris: female; 10-16.
Anphira branchialis: female; all scales = 5 mm.
10-17 10-18
10-19 10-20
10-17. Asotana formosa: female; scale = 5 mm; 10-18. Riggia brasiliensis: female; scale = 10 mm; 10-19.
Riggia nana: female; scale = 2 mm; 10-20. Artystone trysibia: female; scale = 5 mm.
10-21 10-22
10-23 10-24
10-21. Paracymothoa astyanactis: female; scale = 2 mm; 10-22. Braga nasuta: male; scale = 2 mm; 10-23.
Telotha silurii: male; scale = 2 mm; 10-24. Philostomella cigarra: female; scale = 5mm.
10-25 10-26
10-29
10-28
10-27
Braga amapaensis: female; 10-25. dorsal view; scale = 5 mm; 10-26. antenna and antennule; scale = 1
mm; 10-27. pleopod 3; 10-28. pereopod 6; 10-29. pereopod 1.
10-30
10-31
10-34
10-33
10-32
10-35
Mandibles & palps, females: 10-30. Vanamea symmetrica; 10-31. Braga nasuta; 10-32. Riggia brasiliensis;
all scales = 0.5 mm; 10-33. Asotana magnífica; scale = 1 mm; 10-34. Nerocila armata; scale = 0.5 mm;
10-35. Ceratothoa gaudichaudii; scale = 0.5 mm.
10-36
10-36. Anphira guianensis: female, lateral; gill chamber of Acnodon oligocanthus; French Guiana; scale = 2mm.
10-37
10-37. Asotana magnífica: female, dorsal; scale = 5 mm.

10-38
10-38. Asotana magnífica: female, lateral; scale = 5mm.

10-39 10-40
10-41
Braga cichlae: male; Rio Negro, Manaus, Amazonas, Brazil; scales = 5 mm; 10-39. dorsal; 10-40. ventral;
10-41. lateral.
10-42
10-43
Braga patagonica: male, 13 mm long; Rondônia, Brasil; scale = 5 mm; 10-42. dorsal; 10-43. lateral.
10-44
10-44. dorsal. Riggia acuticauda: female, dorsal; scale = 2mm; col. Otavio Froehlich.
10-45
10-46
Riggia cryptocularis : female, length = 20 mm; scale = 5 mm; 10-45. dorsal; 10-46. ventral.
10-47
10-48
10-47. Anphira branchialis: male, in Serrasalmus sp.; scale = 10 mm; 10-48. Asotana magnífica: female, on
tongue of Serrasalmus sp.; scale = 10 mm.
10-49
10-50
10-49. Anphira sp.: gills of Curimatopsis sp.; scale = 5 mm; 10-50. Braga sp.: gills of piranha; scale = 20 mm.
10-51
10-52
Stenarchorhamphus muelleri with Riggia sp.: 10-51. isopod “in situ”; 10-52. isopod removed to show size;
scale = 10 mm.
10-53 10-54
10-37
10-55
Excorallana sp.: male; Amazonas, Brazil; scale = 2 mm; 10-53. dorsal; 10-54. ventral; 10-55. lateral.
10-37
10-56
10-56. Artystone trysibia: female in Geophagus brasiliensis.

10-57
10-57. Anphira junki: female, latero-dorsal view.

10-58
10-58. Nerocila armata: male; length = 20 mm.

10-59
10-59. Riggia paranensis: female, dorsal.

10-60
10-60. Riggia paranensis: female, ventral.

10-61
10-62
Telotha henselii: female; 10-61. dorsal; 10-62. lateral.

ARAUJO, C.S. DE & V.E. THATCHER (2003): Anphira junki n. sp. (Isopoda, Cymothoidae) a gill
chamber parasite of Triportheus albus and T. flavus (Pisces) in the Brazilian Amazon. -
Amazoniana 17(3/4): 283-290.
BASTOS, P.B. & V.E. THATCHER (1997): A redescription of Riggia paranensis SZIDAT, 1948, (Isopoda,
Cymothoidae) based on 32 specimens from curimatid fish of Rio de Janeiro, Brazil, with a
redefinition of the genus. - Mem. Inst. Oswaldo Cruz. 92(6): 755-760.
BOUVIER, G. (1953): De quelques crustacés parasites des poissons d’eau douce de Suisse et du
Brasil. - Bull. Soc. Vaudoise Sci. Nat. 65(283): 423-427, figs. A-F.
BOWMAN, T.E. (1986): Paracymothoa tholoceps, a freshwater parasitic isopod from Southern Venezuela
(Flabellifera: Cymothoidae). - Proc. Biol. Soc. Wash. 99(4): 753-756.
BOWMAN, T.E. & C. DÍAZ-UNGRÍA (1957): Isópodos quimotoideos de peces de las aguas venezolanas.
- Mem. Soc. Cien. Nat. La Salle (Caracas) 17(47): 112-124, figs. 1-4.
BRUSCA, R.C. (1978): Studies on the cymothoid fish symbionts of the Eastern Pacific (Crustacea:
Isopoda: Cymothoidae) II. Systematics and biology of Lironeca vulgaris STIMSON 1857. - Occas.
Pap. Allan Hancock Found. New Ser. 2: 1-19.
BRUSCA, R.C. (1981): A monograph on the Isopoda Cymothoidae (Crustacea) of the Eastern
Pacific. - Zool. J. Linn. Soc. London 73: 117-199.
BRUSCA, R.C. & M.R. GILLIGAN (1983): Tongue replacement in a marine fish (Lutjanus guttatus) by a
parasitic isopod (Crustacea: Isopoda). - Copeia 3: 813-816.
CORDERO, E.H. (1937): Nerocila fluviatilis y otros parásitos de las familias Cymothoidae y Bopyridae
del Uruguay y del Brasil. - An. Mus. Hist. Nat. Montevideo 24(12): 1-11, pl. 1.
GIAMBIAGI DE CALABRESE, D. (1922): Cuatro nuevos isópodos de la Argentina. - Physis 5 (20): 230-
244, pls. 1-4.
GIAMBIAGI DE CALABRESE, D. (1933): Descripción complementaria de un isópodo de agua dulce
Braga fluviatilis RICHARDSON. - An. Mus. Nac. Hist. Nat. (Buenos Aires) 37: 511-515.
HUIZINGA, H.W. (1972): Pathobiology of Artystone trysibia SCHIÖDTE (Isopoda: Cymothoidae), an
endoparasitic isopod of South American freshwater fishes. - J. Wildl. Dis. 8: 225-232.
LEIGH-SHARPE, W.H. (1937): Badroulboudour splendida n. g. et sp., a new parasitic isopod from
Ecuador. - Parasit. 29: 391-394.
LEMOS DE CASTRO, A. (1955): “Pracymothoa astyanactis” g. n. esp. n. Isopode parasita de peixe de água
doce (Isopoda, Cymothoidae). - Rev. Brasil. Biol. 15(4): 411-414.
LEMOS DE CASTRO, A. (1959): Sôbre as espécies sul-americanas do gênero Braga SCHIÖDTE &
MEINERT, 1881 (Isopoda, Cymothoidae). - Arch. Mus. Nac. (Rio de Janeiro) 49: 69-77.
LEMOS DE CASTRO, A. & J. LOYOLA E SILVA (1985): 33. Isopoda. - In SCHADEN, T. (org.): Manual de
identificação de invertebrados límnicos do Brasil.- CNPq, Brasilia: 10 pp.
LEMOS DE CASTRO, A. & J.P. MACHADO FILHO (1946): “Artystone trysibia” SCHIÖDTE, um crustáceo
parasita de peixe d’água doce de Brasil, com descrição do alótipo macho (Isopoda,
Cymothoidae). - Rev. Brasil. Biol. 6(3): 407-413.
MARTENS, E. VON (1869): Südbrasilianische Süss-und Brackwasser-Crustaceen nach den Sammlun-
gen des Dr. REINH. HENSEL. - Arch. Naturgesch 35(1): 1-37, pls. 1-2.
MONOD, T. (1931): Sur un Braga do Paraguay. - Ann. Parasit. Hum. Comp. 9: 363-365.
PAIVA CARVALHO, J. (1939): Sôbre un caso curioso de ectoparasitismo. - Rev. Industr. Animais (Sâo
Paulo) (n.s.) 2(3): 41-44.
RICHARDSON, H. (1911): Description d’un nouvel isopode du genre Braga provenant d’une rivière de
l’Amerique du Sud. - Bull. Mus. Natl. Hist. Nat. (Paris) 17: 94-96.
RINGUELET, R. (1947): Anotaciones sobre copépodos e isópodos parásitos de peces. - Notas Mus.
La Plata Zool. (Argentina) 12(98): 93-107, pls. 1-2.
ROMESTAND, B. & J.P. TRILLES (1977): Dégénérescence de la langue des bogues (Boobs boops L., 1758)
(Téléostéens, Sparidae)) parasitées par Meinertia oestroides (RISSO, 1826) (Isopoda, Flabellifera,
Cymothoidae). - Z. Parasitenk. 54: 47-53.
ROMESTAND, B. & J.P. TRILLES (1979): Influence des cymothoadiens Meinertia oestroides, Meinertia
parallela et Anilocra physodes (Crustacés, Isopodes: parasites de poisons) sur la croissance des
poissons hôtes Boops boops et Pagellus erythrinus (sparidés). - Z. Parasitenk. 59: 195-202.
SCHIÖDTE, J.C. (1866): Krebsdyrenes Sugemund. - Naturhist. Tidsskr. Ser. 3, 4: 169-206, pls. 10-11.
SCHIÖDTE, J.C. & E. MEINERT (1881): Symbolae ad Monographiam Cymothoarum Crustaceorum
Isopodus Familiae. IV Cymothoidae. - Naturhist. Tidsskr. Ser. 3, 14: 221-454, pls. 6-7 & 24-36.
SCHOUTEN, G.B. (1932): Isópodo del género Braga. - Rev. Soc. Cient. Paraguay 3: 105-106.
STADLER, T. (1972): Braga bachmanni, nuevo ectoparásito de Ancistrus cirrosus (Crustacea, Cymothoid-
ae en piece, Loricariidae). - Neotropica 18(57): 141-145.
SZIDAT, L. (1948): Riggia paranensis n. g., n. sp. un isópodo parásito de la cavidad del cuerpo de
“Curimata platana” GÜNTHER, del Rio Paraná (y descripción del huesped por el Prof. ALBERTO
NANI). - Rev. Inst. Nac. Mus. Argent. Cienc. Nat. Bernadino Rivadavia, Cienc. Zool. 1(2): 47-65.
SZIDAT, L. (1965): Sobre la evolución del dimorfismo sexual secundario en isópodos parásitos de la
familia Cymothoidae (Crustacea, Isopoda). - An. Seg. Congr. Lat. Amer. Zool. (1962) 2: 83-87.
SZIDAT, L. & O. SCHUBART (1960): Neue und seltene parasitische Süsswasser-Asseln der Familie
Cymothoidae aus dem Río Mogi Guassu, Brasilien (Isopoda). - An. Acad. Brasil. Cien.
32(1): 107-124.
TABERNER, R. (1976): Un nuevo isópodo de la familia Cymothoidae, Paracymothoa parva sp. nov.
parásito de Hyphessobrycon callistus (COULENGER, 1860) EIGENMANN, 1910 (Pisces, Characidae). -
Physis, B 35(91): 163-169.
TABERNER, R. (1977): Nebula maculatus gen. nov. sp. nov. (Crustacea, Isopoda, Cymothoidae),
ectoparásito de Metynnis maculatus (KNER, 1869) BERG, 1897 (Pisces, Characidae). - Physis, B
35(92): 141-146.
TABERNER, R. (1979): Aclaración sobre el nombre Nebula maculatus TABERNER, 1977 (Crustacea,
Isopoda, Cymothoidae). - Physis, B 38(94): 55.
THATCHER, V.E. (1988): Asotana magnifica n. sp. (Isopoda, Cymothoidae) an unusual parasite
(commensal?) of the buccal cavities of piranhas (Serrasalmus sp.) from Roraima, Brazil. -
Amazoniana 10(3): 239-248.
THATCHER, V.E. (1994a): Vanamea gen. nov. (Isopoda; Cymothoidae) for Livoneca symmetrica VAN
NAME, 1925, and a redescription of the species based on specimens from Brazilian piranhas.
- Acta Amazonica 23(2-3): 287-296.
THATCHER, V.E. (1994b): Anphira branchialis gen. et sp. nov. (Isopoda, Cymothoidae) a gill cavity parasite
of piranhas (Serrasalmus spp.) in the Brazilian Amazon. - Acta Amazonica 23(2-3): 297-307.
THATCHER, V.E. (1995a): Anphira xinguensis sp. nov. a gill chamber parasite of an Amazonian
serrasalmid fish Ossubtus xinguense JÉGU, 1992. - Amazoniana. 13(3/4) 293-303.
THATCHER, V.E. (1995b): Comparative pleopod morphology of eleven species of parasitic isopods
from Brazilian fish. - Amazoniana 13(3/4): 305-314.
THATCHER, V.E. (1996): Braga amapaensis n. sp. (Isopoda, Cymothoidae) a mouth cavity parasite of
the Amazonian fish, Acestrorhynchus guyanensis MENEZES. - Amazoniana 14(1/2): 121-129.
THATCHER, V.E. (1997): Mouthpart morphology of six freshwater species of Cymothoidae (Isopoda)
compared to that of three marine forms. - Amazoniana 14(3/4): 311-322.
THATCHER, V.E. (2000a): The isopod parasites of South American Fishes.- In: SALGADO-MALDONA-
DO, G., GARCÍA ALDRETE, A.N. & V.M. VIDAL-MARTÍNEZ: Metazoan parasites in the Neotro-
pics: 193-226. Universidad Nacional Autónoma de México Press, México, D.F.: 310 pp.
THATCHER, V.E. (2001): Some unusual features of Amazonian fish parasitic isopods (Cymothoidae).
- In: KENSLEY, B. & R.C. BRUSCA (eds.): Isopod systematics and evolution: 337-342. Balkema
Publ., Rotterdam: 357 pp.
THATCHER, V.E. (2002a): Anphira guianensis sp. nov. (Isopoda, Cymothoidae) from the gills of
Acnodon oligacanthus EIGENMANN (Pisces, Serrasalmidae) of French Guiana. - Rev. Bras. Zool.
19, Suppl. 2: 53-59.
THATCHER, V.E. (2004): The isopods of South American fishes. - Imprensa Ministério de Ciência e
Tecnologia Press, Museu Paraense Emílio Goeldi, Belém: 131 pp., 209 figs.
THATCHER, V.E. & M.L. CARVALHO (1988): Artystone minima n. sp. (Isopoda, Cymothoidae) a body
cavity parasite of the pencil fish (Nannostomus beckfordi GÜNTHER) from the Brazilian Amazon.
- Amazoniana 10(3): 255-265.
THATCHER, V.E. & C. LOBOS-BLUMENFELDT (2001): Anilocra montti sp. n. (Isopoda, Cymothoidae) a
parasite of caged salmon and trout in Chile. - Revta. bras. Zool. 18 (Supl 1): 269-276.
THATCHER, V.E. & I. SCHINDLER (1999): Artystone bolivianensis sp. n. (Isopoda, Cymothoidae) from a
loricariid catfish of the Bolivian Amazon. - Amazoniana 15(3/4): 183-191.
THATCHER, V.E., LOPES, L.P.C. & O. FROEHLICH (2002): Riggia acuticaudata sp. n. (Isopoda,
Cymothoidae) from the body cavity of a freshwater fish of Mato Grosso do Sul State, Brasil.
- Rev. Brasil. Zool. 19 (Supl.2): 195-201.
THATCHER, V.E., SOUZA-CONCEIÇÃO, J.M. & G.F. JOST (2003): Lironeca desterroensis sp. n. (Isopoda,
Cymothoidae) from the gills of a marine fish, Cetengraulis edentulus CUVIER, of Santa Catarina
Island, Brazil. - Rev. Brasil. Zool. 20: 251-255.
THATCHER, V.E., LOPES, L.P.C. & O. FROEHLICH (2003): Riggia cryptocularis sp. n. (Isopoda,
Cymothoidae) from the body cavity of a freshwater fish of Mato Grosso do Sul State, Brasil.
- Rev. Brasil. Zool. 20: 285-289.
THATCHER, V.E., LOYOLA E SILVA J. DE, JOST G.F. & J.M. SOUZA-CONCEIÇÃO (2003): Comparative
morphology of Cymothoa spp. (Isopoda, Cymothoidae) from Brazilian fishes, with the
description of Cymothoa catarinensis sp. nov. and redescriptions of C. excise PERTY and C. oestrum
(LINNAEUS). - Rev. Bras. Zool. 20: 541-552.
TRILLES, J.P. (1973): Notes documentaries sur les isopodes cymothodiens parasites de poissons d’eau
douce del ‘Amérique du Sud. - Bull. Mus. Natl. Hist. Nat. Paris, sér. 3 Zool. 88: 239-272, pls. 1-2.
VAN NAME, W.G. (1925): The isopods of Kartabo, Bartica District, British Guiana. - Zoologica 6(5):
461-503, pls. A-B.
WEIBEZAHN, F.H. & M.V. RAMÍREZ (1957): Mortandad de peces de agua dulce causada por un
crustáceo parásito, Artystone trysibia SCHIÖDTE & MEINERT, 1866 (Isopoda, Cymothoidae). -
Bol. Soc. Venezol. Cienc. Nat. 18(89): 153-156.
11.
HIRUDINEA, PENTASTOMIDA AND
OTHER PATHOGENS
HIRUDINEA
The Hirudinea, or leeches, comprise a highly specialized class of the phylum Annelida.
Most occur in freshwater but many are also found in the sea and some are terrestrial in the
humid tropics. They are thought to have arisen from the class Oligochaeta, to which the
earthworm belongs. Leeches have segmented bodies, as do the other members of the
phylum, but unlike the earthworm, the segments are not defined by internal septa.
Leeches can generally be recognized by the anterior and posterior suckers that they
possess. Those that attack fish do so as temporary ectoparasites seeking a blood meal.
The bodies of leeches are frequently spindle-shaped being narrower anteriorly
than posteriorly. In cross-section they are either circular or oval. In size, they range from
5 mm to about 45 cm. Surface features include pigmentation (red, brown, green and
yellow in spots bands or longitudinal lines), one or more pairs of eyes, annuli and
sometimes dorsal papillae.
The digestive system of leeches consists of an anterior mouth, followed by a
muscular pharynx, an esophagus and an intestine. The mouth is either provided with
teeth or it has a protrusible proboscis. The intestine shows segmentation in that there
are paired lateral ceca. The intestine opens through the anus that is located dorsally over
the posterior sucker.
All leeches are hermaphroditic. The male reproductive system consists of paired
testes (or testisacs) five to ten in number or they may be more numerous. On each side
there are vasa efferentia joining the testes to the vas deferens that, in turn, leads into the
atrium. The atrium is a three-chambered structure with lateral horns. The female system
is comprised of paired ovaries (or ovisacs) terminating in ducts which join to form a
vagina. In most species, the male and female genital pores are separate and located on
the 11th and 12th body segments, respectively.
Most leeches are predatory and feed on small invertebrates, but a few species attack fish
and other vertebrates. Predatory leeches swim actively but those that parasitize fish usually
hide on aquatic vegetation and wait until a fish passes near.
Reproduction in leeches is similar to that of earthworms in that there is copulation
and a mutual exchange of sperm bundles (spermatophores). In some species there is a
penis for depositing sperm in or near the vagina, but in others, spermatophores are simply
implanted on the surface. In such species, the sperm penetrate the body wall by a local
histolysis and make their way to the ovisac where they fertilize the ova. The fertilized ova
(eggs) are released through the female genital pore and are there encapsulated by
secretions from a band of glandular cells (clitellum). Several eggs are usually placed in
each capsule (cocoon) and these are normally fastened to a substrate or buried in the mud.
In Glossiphoniidae, however, true cocoons are not formed, but the fertilized eggs are
carried on the ventral surface of the body in membranous capsules. After the eggs hatch,
the young leeches remain attached to the parent for some time.
III. Pathology
Other than localized hemorrhaging at the feeding sites, little specific pathology has been
associated with Neotropical leeches. In heavy infestations, leeches can produce anemia
and death from blood loss. Furthermore, after feeding they drop from the host leaving
small wounds that may facilitate secondary invasions by bacteria and viruses. Leeches also
serve as intermediate hosts for Trypanosoma and Cryptobia, which are flagellate protozoan
parasites of the circulatory system.
Leeches are most often introduced into aquaria on aquatic vegetation. Plants collected in
nature should be maintained in aquaria for several weeks before being placed with fish.
Leeches are relatively sensitive to chemicals in the water, so most of the common dip
treatments will cause them to leave the fish. One of the easiest and cheapest is the 1:4000
formalin solution treatment described in Chapter 3.Contaminated ponds should be
drained and treated with quick lime.
Living leeches should be killed by placing them in 15 % alcohol for 15 minutes. They can
then be fixed by arranging them between glass microscopes slides in a Petri dish and
pouring AFA over them. Permanent whole-mount slides can be made by the method
described for trematodes in Chapter 4.
VI. Identification and key
Some important characters used in identifying leeches are: body shape, size and color;
number and position of eyes; and position of genital pores. So far, only three genera and
four species of Hirudinea have been reported from Neotropical freshwater fish.
Key to leeches from Neotropical freshwater fishes
I. Body dorso-ventrally flattened; tapering towards anterior extremity; one, 2, 3 or 4

pairs of eyes present on head end; eyespots lacking on posterior sucker and on
body; three annuli present for each body segment; oral sucker small, continuous
with neck ............................................................... Glossiphoniidae (Figs. 11-3 to 11-5)
A. One pair of eyes present .............................................................................. Placobdella
B. Two to four pairs of eyes present ............................................................ Theromyzon
II. Body cylindrical, slender; none, 1 or 2 pairs of eyes present on head end; eyespots may
be present on posterior sucker or lateral margins of body; seven or more annuli
present for each body segment; oral sucker expanded, distinct from neck ..............
..................................................................... Piscicolidae ..................................... Myzobdella
Clave para Sanguijuelas de peces Neotropicales de agua dulce
I. Cuerpo aplanado dorso-ventralmente, estrechándose hacia la extremidad anterior; un,

2, 3 o 4 pares de ojos presentes en la cabeza; manchas ocelares faltan en la ventosa
posterior y en el cuerpo; tres anillos presentes para cada segmento del cuerpo.
Ventosa oral pequeña, contínua con el cuello .................................................................
................................................................................... Glossiphoniidae (Figs. 11-3 a 11-5)
A. Un par de ojos presentes .............................................................................. Placobdella
B. Dos a 4 pares de ojos presentes ............................................................... Theromyzon
II. Cuerpo cilíndrico, delgado; ninguno, 1 o 2 pares de ojos presentes en la cabeza;
manchas ocelares pueden estar presentes en la ventosa posterior o en los márgenes
laterales del cuerpo; siete o más anillos presentes en cada segmento del cuerpo;
ventosa oral expandida, diferenciada del cuello .............................................................
..................................................................... Piscicolidae ..................................... Myzobdella
VII. Checklist of Hirudinea from Neotropical fishes
Glossiphoniidae
Body flattened, wider posteriorly and tapering anteriorly. Oral sucker
fused with body; mouth a small pore in oral sucker through which a
muscular proboscis can be protruded; jaws and teeth absent. One to 4
pairs of eyes present; eyespots absent on body and posterior sucker.
Three annuli present per segment, in midbody; clitellum not distinct.

Eggs in membranous sacs on ventral surface of adult; young cling to
ventral surface of adult for some time. Free-living and parasitic on
invertebrates and vertebrates.
Placobdella BLANCHARD, 1893
Glossiphoniidae. Body heavily pigmented, usually greenish or brownish
with spots, stripes or blotches of yellow or orange; dorsum frequently
papillate; size 10 – 65 mm. Anterior end with or without expanded head-
like structure. One pair of eyes present; accessory eyes rarely present.
Free-living or parasitic on fish or turtles.
P. molesta CORDERO, 1934: turtles and fishes: Uruguay. Description unavailable.
P. parasitica (SAY, 1924): turtles and fishes: United States & Mexico. Body =
38-65 mm long; Coloration variable, dorsum usually provided with
median, longitudinal stripe; Ventral surface with 8-12 bluish, greenish or
brownish longitudinal stripes.
Theromyzon PHILLIPPI, 1867
Glossiphoniidae. Body gelatinous, not heavily pigmented, pale amber or
green in color; spots or lines sometimes present; shape variable. Four pairs
of eyes present. Gonopores separated by 2-4 annuli. Free-living or
parasitic on fish or in nasal fossae or eyes of aquatic birds.
T. propinquum RINGUELET, 1947: Percichthys trucha: Argentina. Body smooth,
except for small sensilla; 14-15.5 mm long. Three annuli present between
male and female gonopores.
Piscicolidae JOHNSTON, 1865
Body cylindrical, narrow; sometimes divided into narrow neck (trach-
elosome) and thicker posterior region (urosome); with or without
lateral projections (pulsatile vesicles) on urosome. Oral sucker usually
distinct from neck. None, 1 or 2 pairs of eyes present; with or
without eyespots (ocelli) on caudal sucker and lateral margins of
urosome. Cocoons attached to substrates, not held on ventral side of
adults. Parasitic on fishes.
Myzobdella LEIDY, 1851
Psiscicolidae. Body cylindrical, divided into trachelosome and urosome;
without dorsal papillae and without lateral extensions of urosome; 12-14
annuli per somite in midbody. Suckers weakly developed; not of greater
diameter than body. Five pairs of testes present. Parasites on marine and
freshwater fishes.
M.platense (CORDERO, 1933): Hoplias malabaricus: Uruguay. Description un-
available.
M.uruguayensis MAÑÉ-GARZÓN & MONTERO, 1977: Rhamdia sapo: Uruguay.
Body = 8.7-12.7 x 0.83-1.04 mm; Pharynx = 1.35-1.68 x 0.18-0.30 mm;
Testes = 0.45-0.53 mm in diameter.
PENTASTOMIDA
Pentastomids are worm-like endoparasitic arthropods with obscure affinities. Formerly the
group was considered a subphylum of Arthropoda, but in more recent years, the tendency
among most zoologists has been to recognize Pentastomida as a separate phylum. Adult
pentastomids are found mainly in the lungs of reptiles and the young (nymphs) occur
encysted in the internal organs of many vertebrates including fish and men. Adults range
from 1 to 20 cm in length, and have cylindrical or flattened bodies that are expanded
anteriorly. The sexes are separate and fertilization is internal. The name pentastomid means
“five-mouthed” and comes from the fact that on each side of the mouth slit there are two
other depressions each containing a retractable claw (making a total of 5 mouth-like slits).
The pentastomids so far reported in fish have all been young, or nymphal stages.
They are ordinarily encysted and they can be recognized by their cylindrical, annulated
bodies and their peculiar claws.
The pentastomid life-cycle involves an intermediate and a definitive host, both of which
are vertebrates. Fish act as intermediate hosts for those that mature in crocodilians, and
most of these belong to the genus Sebekia. The adults live in the air passages of the lungs
where they copulate. Females produce eggs that pass to the external environment in the
host’s feces. When the eggs are ingested by a fish, they hatch in the intestinal tract and
four-legged larvae, somewhat resembling mites emerge. These larvae make their way
through the wall of the intestine aided by the legs that are tipped with small claws. Once
within the tissues of the fish, the larvae metamorphose into nymphs, molting several
times as they grow. After an infected fish is eaten by a crocodilian, the larvae make their
way to the lungs where they mature.
III. Pathology
The passage of larval pentastomids through the intestinal wall of a fish could probably
provoke a localized inflammation, but this has not been documented. The presence of nymphs
in the mesenteries and other organs causes localized inflammatory reactions leading to fibrotic
encapsulation. After complete encapsulation, no further histopathology is observed.
Crocodilians should not be permitted to inhabit ponds where fish are grown. Care should
also be taken not to introduce mud, or other material that might be contaminated with
crocodilian feces, into fish ponds and aquaria. No specific treatment is possible or
necessary for fish already having encysted nymphs.
Pentastomid nymphs (Fig. 11-1) can be studied in temporary phenol preparations after
first dehydrating them partially in 95 % alcohol. Permanent whole mounts can be made by
the methods described in Chapter 6 for nematodes. Claws can be dissected out with
needles and mounted separately for comparative studies.
VI. Identification
It is not presently possible to identify pentastomid nymphs from fish to species. It is

presumed that all are Sebekia, but since there are several species in the genus, life-cycle
studies would need to be performed in order to identify specific differences at the
nymphal level.
VII. Checklist for Pentastomida
TRAVASSOS et al. (1928) called pentastomid nymphs found in fish, Porocephalus gracile
(DIESING, 1836). The genus Porocephalus, as presently defined by specialists, occurs as an
adult in the lungs of snakes and as a nymph in the organs of mammals. Porocephalus
nymphs probably do not occur in fish. The pentastomid nymphs from fish are all thought
to represent species of Sebekia, and these have been reported from the following hosts:
Astronotus ocellatus Pimelodus vituga

Astyanax mexicanus Pinirampus pirinampu
Bagrus pemecus Potamotrygon motoro
Electrophorus electricus Pseudoplatystoma tigrinum
Gymnotus carapo Rhaphiodon vulpinus
Hemisorubim platyrhynchus Salminus brevidens
Hoplias malabaricus Serrasalmus piraya
Pellona castelnaeana Stenarchus albifrons
Phractocephalus hemiopterus Symbranchus marmoratus
Pimelodus megacephalus Tachysurus hertzbergii
OTHER PATHOGENS
Tropical Gill-Rot:
One of the most common disease conditions found in pond reared fish in the tropics is
gill rot. By the time the problem is noted, the disease has frequently advanced beyond the
point of treatment. Affected fish are usually observed to swim sluggishly or erratically and
spend much time at the surface gasping for breath. A cursory examination of the gills of
such fish will show them to be ragged, whitish, with localized hemorrhaging areas and
much mucus. Closer inspection may reveal parasites, such as Myxozoa, Ciliophora and
Monogenoidea. Localized or generalized fungal growths may also be visible. Many kinds
of bacteria will also be present.
The immediate cause of death in these cases is asphyxia, and the moment of death
will be advanced, of course, if there should be any drop in the level of dissolved oxygen
in the water. In natural, or outdoor, ponds there is a normal slump in oxygen level every
night because at sundown photosynthesis stops abruptly. Typically, a pisciculturist, whose
fish are “doing fine”, arrives at his pond some morning and finds about half of them dead
and floating around on the surface. He gathers some and dead fish in hand goes to consult
an ichthypathologist or a veterinarian to find out how to save the survivors. However, it is
too late. Treatment at this point will ordinarily hasten rather than prevent death.
Although the immediate cause of death in gill rot infections is asphyxia caused by gill
inefficiency, the root cause is malnutrition. The condition described above is symptomatic
of deficiencies in vitamins A and C and proteins. The root cause boils down to a matter
of simple economics or human greed. The pisciculturist, like any businessman, wants to
maximize profit and minimize expense. Since vitamins and proteins cost more than
carbohydrates, he puts a minimal amount of these in the ration he feeds his fish. The
result is frequently gill rot.
Encysted parasites and foreign bodies:

An inspection of the interior of any wild-caught tropical fish will usually reveal the
presence of numerous “white spots” of various sizes and shapes. These represent
invaders that the fish has been able to immobilize in fibrous capsules. Most invaders reach
the body cavity by passing through the intestinal wall, but many of them are stopped
within the wall itself . Those that reach the other side often end up in the mesentery that
supports the intestine (Fig. 11-2). Herbivorous and omnivorous fish frequently have their
stomachs and intestines pierced by plant spines, which also become encapsulated. The
skin barrier is more effective, but is also breached at times. Occasional oddities turn up in
the intestinal walls of fish, such as larval mites and insects (Figs. 11-6 & 11-7). In order to
find out what is inside the “white spots”, it is best to excise them, partially dehydrate them
in 95 % alcohol and clear them in pure phenol.
VIII. Plates of Hirudinea and miscellaneous pathological conditions

(Figs. 11-1 to 11-7)
11-1
11-2
11-1. Pentastomid nymph encysted in the mesentery of Colossoma macropomum. 11-2. Cestode larvae
(spherical structure) and nematode larva (elongate form) in the intestinal wall of Chaetobranchus semifasciatus.
11-3
11-5 11-4
Morphology of glosiphoniid leeches: 11-3. and 11-5. Branching intestinal tract; 11-4. Anterior end,
showing eyes, annuli and proboscis.
11-6
11-7
11-6. & 11-7. Insects in the intestinal wall of Semaprochilodus insignis.

Hirudinea
CABALLERO & C., E. (1940): Sobre la presencia de la Placobdella rugosa (Hirudinea: Glossiphoniidae) en
las aguas del Lago de Xochimilco. XIII. - An. Instituto Biológico de México 12(1/2): 255-260.
CABALLERO & C., E. (1941): Hirudineos de México. XVI. Nuevos huespedes y localidades para
algunas sanguijuelas ya conocidas y descripción de una nueva especie. - An. Instituto
Biológico de México 12(2): 752-753.
CABALLERO & C., E. (1960): Hirudineos de México. XXII. Taxa y nomenclatura de la clase
Hirudinea hasta géneros (nueva edición). - An. Instituto Biológico de México 30: 227-242.
MAÑE-GARZÓN, F. & R. MONTERO (1977): Myzobdella uruguayensis n. sp. (Hirudinea, Piscicolidae)
parasita de las branquias del bagre amarilla, Rhambdia sapo (VALL.). - Rev. Biol. Uruguay 5: 59-65.
RINGUELET, R.A. (1944): Sinopsis sistemática y zoogeográfica de los hirudíneos de la Argentina,
Brasil, Chile, Paraguay y Uruguay. - Rev. Mus. La Plata, n.s. 3 (Zool.) 3(22): 163-232.
RINGUELET, R.A. (1947): Notas sobre hirudíneos neotropicales III. Theromyzon propinquus nov. sp. de
la Argentina. - Notas Mus. La Plata 12(100): 217-222.
RINGUELET, R.A. (1976): Clave par las familias y géneros de sanguijuelas (Hirudinea) de aguas dulces
y terrestres de Mesoamérica y Sudamérica. - Limnobios 1(1): 9-19.
Pentastomida
THATCHER, V.E. (1981): Patologia de peixes da Amazônia brasileira, 1. Aspectos gerais. - Acta
Amazônica 11(1): 125-140.
TRAVASSOS, L., ARTIGAS, P. & C. PEREIRA (1928): Fauna helminthológica dos peixes de água doce do
Brasil. - Arch. Inst. Biol. São Paulo, Brasil 1: 5-68, 155 figs.
12.
ADDENDUM: HOST-PARASITE TABLE
In the following table, the scientific names of host fish are listed alphabetically without
regard to phylogeny. Beside each fish name, the parasites reported from that host are
listed. Trypanosoma (Protozoa) and Sebekia (Pentastomida) are omitted here because their
hosts are listed in Chapters 1 and 11 respectively.
Host fish Parasites

Phylum or Class Genera and species
Acanthicus hystrix Monogenoidea Trinigyrus acuminatus
Acestrorhampus (= Oligosarcus) macrolepis Nematoda Procamallanus (S.) hilarii
Acestrorhampus sp. (= Oligosarcus) Myxozoa Myxidium gurgeli
Ciliophora Nyctotherus piscicola
Trematoda Halipegus genarchella
Halipegus parvus
Prosorhynchus costai
Prosthenhystera obesa
Nematoda Paracapillaria piscicola
Contracaecum sp. (larvae)
Eustrongylides sp. (larvae)
Acestrorhynchus falcatus Trematoda Bellumcorpus major
Acanthocephala Palliolisentis polyonca
Procamallanus (S.) inopinatus
Copepoda Ergasilus turucuyus
Acestrorhynchus falcirostris Copepoda Ergasilus turucuyus
Acestrorhynchus guyanensis Isopoda Braga amapaensis
Acestrorhynchus lacustris Nematoda Procamallanus (S.) paraensis
Travassosnema travassosi
Achirus lineatus Acanthocephala Neoechinorhynchus variabilis
Achirus mazatlanus Acanthocephala Neoechinorhynchus roseum
Acnodon normani Monogenoidea Notozothecium euzeti
Acnodon oligacanthus Isopoda Anphira guianensis
Aequidens maroni Monogenoidea Sciadicleithrum aequidens
Sciadicleithrum cavanaughi
Aequidens pulcher Trematoda Crassicutis chuscoi

Acanthocephala Pandosentis iracundus
Branchiura Dolops geayi
Ageneiosus brevifilis Cestoda Anthobothrium mandube
Endorchis mandube
Gibsoniela mandube
Copepoda Gamispinus diabolicus
Isopoda Excorallana berbicensis
Ageneiousus valenciennesi Nematoda Cucullanus pinnai
Ancistrus cirrhosus Nematoda Rhaphidascaris (S.) hypostomi
Isopoda Braga bachmanni
Ancistrus dolicopterus Nematoda Capillostrongyloides ancistri
Ancistrus sp. Monogenoidea Nothogyrodactylus amazonicus
Nothogyrodactylus clavatus
Nothogyrodactylus plaesiophallus
Nematoda Guyanema ancistri
Isopoda Riggia cryptocularis
Riggia acuticaudata
Anodus elongatus Copepoda Brasergasilus anodus
Brasergasilus oranus
Arapaima gigas Monogenoidea Dawestrema cycloancistrioides
Dawestrema cycloancistrium
Dawestrema punctata
Trematoda Caballerotrema arapaimense
Caballerotrema brasiliense
Goezia spinulosa
Cestodaria Nesolecithus janicki
Schizochoerus liguloideus
Nematoda Camallanus tridentatus
Goezia spinulosa
Philometra senticosa
Porrocaecum draschei
Acanthocephala Polyacanthorhynchus macrorhynchus
Polyacanthorhynchus rhopalorhynchus
Copepoda Ergasilus sp.
Branchiura Argulus sp.
Dolops discoidalis
Ariopsis seemanni Acanthocephala Pseudogorgorhynchus arii
Arius comersonii Monogenoidea Fridericianella ovicola
Astronotus ocellatus Monogenoidea Gusseiva asota
Gusseiva astronoti
Gusseiva rogersi
Goezia spinulosa
Astronotus ocellatus Branchiura Argulus sp.

Dolops bidentata
Dolops discoidalis
Dolops geayi
Astyanax bimaculatus Monogenoidea Cycloplectanum americanum
Urocleidoides costaricensis
Palombitrema heteroancistrium
Urocleidoides trinidadensis
Trematoda Magnivitellinum simplex
Nematoda Cosmozynemoides aguirrei
Paraseuratum albidum
Procamallanus (S.) cearensis
Procamallanus (S.) hilarii
Rhabdochona australis
Rhabdochona acuminate
Spiniteatus rudolphiheringi
Travnema travnema
Acanthocephala Quadrigyrus torquatus
Branchiura Argulus juparanaensis
Isopoda Paracymothoa astyanactis
Astyanax fasciatus Myxozoa Henneguya wenyoni
Monogenoidea Anacanthocotyle anacanthocotyle
Diaphorocleidus kabatai
Gyrodactylus neotropicalis
Jainus hexops
Urocleidoides astyanacis
Urocleidoides kabatai
Urocleidoides strombicirrus
Urocleidoides trinidadensis
Trematoda Antorchis lintoni
Chalcinotrema ruedasueltensis
Dadaytremoides grandistomis
Halipegus dubius
Halipegus tropicus
Saccocoelioides octavus
Nematoda Capillostrongyloides sentinosa
Paraseuratum albidum
Rhabdochona fasciata
Astyanax fasciatus Procamallanus (S.) hilarii
Astyanax fasciatus Acanthocephala Quadrigyrus torquatus

Astyanax scabripinnis Monogenoidea Urocleidoides astyanacis
Astyanax sp. Trematoda Phyllodistomoides duncani
Nematoda Pseudocapillaria sentinosa
Rhabdochona fasciata
Procamallanus (S.) wrighti
Copepoda Minilernaea floricapitella
Atherina bleekeri Copepoda Ergasilus orientalis
Atherinichthys Isopoda Braga patagonica
Atractosteus tristoechus Cestoda Proteocephalus manjuariphilus
Auchenipterus nuchalis Nematoda Cucullanus brevispiculus
Basilichthys microlepidotus Cestoda Proteocephalus macdonaghi
Belonesox belizanus Monogenoidea Salsuginus neotropicalis
Bergiaria westermanni Nematoda Procamallanus (S.) freitasi
Boulengerella lucia Trematoda Genelopa magnacirrus
Paraproctotrema delicata
Brachyplatystoma filamentosum Cestoda Endorchis piraeeba
Nominoscolex piraeeba
Brachyplatystoma rousseauxii Cestoda Amphoteromorphus peniculus
Nominoscolex dorad
Nominoscolex sudobim
Brachyplatystoma sp. Isopoda Telotha henselii
Vanamea symmetrica
Brachyplatystoma vaillanti Cestoda Anabothrium piramutab
Monticellia piramutab
Brevoortia pectinata Copepoda Ergasilus euripedesi
Brycon brevicaudata Nematoda Procamallanus (S.) inopinatus
Brycon cephalus Monogenoidea Annulotrematoides bryconi
Copepoda Amplexibranchius bryconis
Brycon cryptopterus Copepoda Ergasilus bryconis
Brycon erythropterum Branchiura Argulus chicomendesi
Brycon falcatus Nematoda Rhabdochona acuminata
Brycon hilarii Nematoda Procamallanus (S.) inopinatus
Acanthocephala Echinorhynchus briconi
Nematoda Procamallanus (S.) hilarii
Brycon lundi Trematoda Dendrorchis neivai
Brycon melanopterus (=erythropterus) Monogenoidea Anacanthorus brevis
Anacanthorus elegans
Anacanthorus kruidenieri
Anacanthorus spiralocirrus
Jainus amazonensis
Tereancistrum kerri
Tereancistrum ornatus
Trinibaculus brazilensis
Brycon melanopterus (=erythropterus) Nematoda Procamallanus (S.) inopinatus

Copepoda Ergasilus bryconis
Brycon sp. Monogenoidea Rhinoxenus anaclaudiae
Bryconops alburnoides Nematoda Procamallanus (D.) dentatus
Bunocephalus coracoideus Myxozoa Myxobolus braziliensis
Callophysus macropterus Monogenoidea Pavanelliella pavanellii
Nematoda Philometra amazonica
Copepoda Ergasilus callophysus
Carapus fasciatus Myxozoa Myxidium fonsecai
Carassius auratus Monogenoidea Gyrodactylus elegans
Branchiura Argulus ernsti
Carnegiella strigata Isopoda Vanamea symmetrica
Catoprion mento Monogenoidea Amphithecium prodotum
Anacanthorus catoprioni
Heterothecium dicrophallum
Odothecium raphidiophallum
Pithanothecium piranhus
Centropomus nigriscens Monogenoidea Mexicotrema bychowskyi
Cephalosilurus (=Pseudopimelodus) zungaro Monogenoidea Ameloblastella mamaevi
Phanerothecium caballeroi
Urocleidoides mamaevi
Cetopsis caecutiens Cestoda Monticellia siluri
Chaetobranchus semifasciatus Trematoda Ascocotyle sp. (metacercariae)
Cestoda Proteocephalidae (plerocercoids)
Chaestostomus leucomelas Trematoda Dadaytremoides grandistomis
Chalceus macrolepidotus Monogenoidea Jainus jainus
Chalcinus nematurus Myxozoa Myxidium cruzi
Charax gibbosus Nematoda Procamallanus (S.) inopinatus
Characidium caucanum Monogenoidea Urocleidoides anops
Characidium lanei Monogenoidea Cacatuocotyle paranaensis
Characidium pterostictum Monogenoidea Cacatuocotyle paranaensis
Chilodus punctatus Acanthocephala Octospiniferoides australis
Chloroscombrus chysurus Isopoda Cymothoa liannae
Chromis sp. Branchiura Argulus chromidis
Cichla monoculus Cestoda Proteocephalus macrophallus
Copepoda Amazolernaea sannerae
Ergasilus coatiarus
Isopoda Vanamea symmetrica
Cichla ocellaris Sporozoa Calyptospora tucunarensis
Monogenoidea Gussevia arilla
Gussevia longihaptor
Gussevia tucunarense
Gussevia undulata
Cichla ocellaris Monogenoidea Sciadicleithrum ergensi

Sciadicleithrum umbilicum
Sciadicleithrum uncinatum
Cestoda Proteocephalus macrophallus
Proteocephalus microscopicus
Sciadocephalus megalodiscus
Nematoda Goezia intermedia
Copepoda Acusicola tucunarense
Ergasilus sp.
Isopoda Braga cichlae
Vanamea symmetrica
Nerocila armata
Cichla sp. Isopoda Braga cichlae
Cichla temensis Copepoda Amazolernaea sannerae
Branchiura Argulus multicolor
Cichlasoma aureum Acanthocephala Neoechinorhynchus golvani
Cichlasoma bimaculatum Monogenoidea Gussevia alii
Gusevia cichlosomatis
Gusevia dobosi
Trinidactylus cichlasomatis
Cichlasoma facetum Nematoda Procamallanus (P.) peraccuratus
Cichlasoma fenestratum Ciliophora Nyctotherus dilleri
Cichlasoma festivum Nematoda Ichthyouris ro
Cichlasoma mayorum Trematoda Crassicutis cichlasomae
Cichlassoma pearsei Monogenoidea Sciadicleithrum bravohollisae
Cichlasoma severum (=Heros severus) Monogenoidea Gussevia alioides
Gussevia dispar
Gussevia disparoides
Trematoda Pronamphistoma cichlasomae
Cichlasoma synspilum Monogenoidea Sciadicleithrum bravohollisae
Cichlasoma tetracanthus Branchiura Argulus cubensis
Cichlasoma urophthalmus Monogenoidea Sciadicleithrum mexicanum
Cichlasoma sp. Trematoda Crassicutis opisthoseminis
Nematoda Ichthyouris ovifilamentosa
Copepoda Ergasilus pitalicus
Isopoda Braga fluviatilis
Cichlidae Nematoda Touzeta ecuadoris
Isopoda Livoneca orinoco
Cnesterodon decemmaculatus Branchiura Argulus vierai
Cochliodon cochliodon Nematoda Procamallanus (S.) annipetterae
Raphidascaris (S.) hypostomi
Colomesus asellus Copepoda Ergasilus colomesus
Colossoma bidens(=Piaractus brachypomus) Myxozoa Myxobolus sp.

Ciliophora Nyctotherus piscicola
Monogenoidea Anacanthorus spathulatus
Trematoda Dadaytrema oxycephala
Denticauda quadrangulata
Pseudocladorchis cylindricus
Pseudoparabaris parabaris
Travassosinia dilatata
Nematoda Klossinemella iheringi
Rondonia rondoni
Spectatus spectatus
Acanthocephala Echinorhynchus jucundum
Branchiura Dolops bidentata
Dolops carvalhoi
Colossoma macropomum Monogenoidea Anacanthorus spathulatus
Linguadactyloides brinkmanni
Mymarothecium boegeri
Notozothecium janauachensis
Nematoda Chabaudinema americana
Cucullanus colossomi
Acanthocephala Neoechinorhynchus buttnerae
Copepoda Perulernaea gamitanae
Branchiura Argulus chicomendesi
Argulus multicolor
Dolops carvalhoi
Colossoma mitre(=Piaractus mesopotamicus) Nematoda Rondonia rondoni
Corydoras aeneus Monogenoidea Urocleidoides corydori
Urocleidoides margolisi
Nematoda Procamallanus (S.) pintoi
Corydoras ehrhardti Monogenoidea Gyrodactylus samirae
Gyrodactylus anisopharynx
Gyrodactylus superbus
Copepoda Minilernaea floricapitella
Corydoras lepidata Nematoda Procamallanus peraccuratus
Corydoras paleatus Monogenoidea Gyrodactylus anisopharynx
Gyrodactylus samirae
Gyrodactylus superbus
Paragyrodactyloides superbus
Philocorydoras platensis
Nematoda Procamallanus (S.) pintoi
Creatochanes affinis Monogenoidea Diaphorocleidus affinis
Jainus robustus
Urocleidoides affinis
Crenicichla geayi Trematoda Crassicutis wallini
Crenicichla geayi Acanthocephala Pandosentis iracundus

Quadrigyrus torquatus
Branchiura Dolops geayi
Crenicichla johanna Trematoda Sphericomonorchis spinulosus
Crenicichla lacustris Isopoda Artystone trysibia
Crenicichla lepidota Sporozoa Calyptospora spinosa
Nematoda Procamallanus (S.) peraccuratus
Crenicichla saxatilis Isopoda Nerocila armata
Crenicichla sp. Branchiura Dolops geayi
Curimata (=Steindachnerina) argentea Monogenoidea Curvianchoratus hexacleidus
Urocleidoides curimatae
Curimata (=Psectrogaster) ciliata Branchiura Argulus sp.
Curimata cyprinoides Copepoda Miracetyma etimaruya
Curimata (=Steindachnerina) elegans Trematoda Zonocotyle bicaecata
Nematoda Cosmoxynemoides aguirrei
Travnema travnema
Acanthocephala Gorytocephalus spectabilis
Monogenoidea Rhinoxenus guianensis
Curimata (=Cyphocharax) gilberti Monogenoidea Androspira chascomusensis
Androspira triangula
Notodiplocerus singularis
Trematoda Zonocotyle bicaecata
Zonocotyloides haroltravossosi
Nematoda Cosmoxynema viannai
Cosmoxynemoides aguirrei
Travnema araujoi
Curimata (=Potamorhina) laticeps Branchiura Argulus sp.
Curimata (=Cyphocharax) platana Isopoda Riggia paranensis
Curimata plumbea Nematoda Travnema travnema
Curimata vittata Trematoda Curimatrema microscopica
Cynopotamus humeralis Trematoda Halipegus dubius
Philostomella cigarra
Cynopotamus kneri Nematoda Cystidicoloides fischeri
Cucullanus pinnai
Paracapillaria piscicola
Cyphocharax gilberti Isopoda Riggia paranensis
Cyphocharax platana Isopoda Riggia paranensis
Doras brunnescens Nematoda Rondonia rondoni
Doras (=Platydoras) costatus Ciliophora Rynchodinium paradoxum
Doras (=Platydoras) costatus Trematoda Pseudocladorchis ferrumequinum

Pseudodiplodiscus cornu
Doras (=Lithodoras) dorsalis Trematoda Pseudocladorchis ferrumequinum
Pseudodiplodiscus cornu
Eigenmannia virescens Myxozoa Henneguya theca
Myxobolus inaequus
Monogenoidea Urocleidoides virescens
Electrophorus electricus Trematoda Echinostoma annulatum
Elipesurus spinicauda Cestoda Eutetrarhynchus araya
Rhinebothroides scorzai
Erythrininus erythrinus Nematoda Paraseuratum albidum
Erythrinidae Nematoda Procamallanus (S.) paraensis
Farlowella amazona Monogenoidea Oogyrodactylus farlowellae
Fitzroyia lineate Acanthocephala Wolffhugelia matercula
Galaxias maculatus Myxozoa Myxobolus galaxii
Myxobolus magellanicus
Monogenoidea Phylureter trygonopsis
Gasterostomus (=Potamorhina) latior Branchiura Dolops sp.
Geophagus brasiliensis Monogenoidea Gyrodactylus geophagensis
Nematoda Cosmoxynemoides aguirrei
Procamallanus peraccuratus
Acanthocephala Neoechinorhynchus paraguayensis
Isopoda Artystone trysibia
Geophagus (=Satanoperca) jurupari Branchiura Argulus multicolor
Geophagus steindachneri Nematoda Cosmoxynemoides aguirrei
Ichthyouris ro
Geophagus surinamensis Monogenoidea Sciadicleithrum geophagi
Geophagus sp. Isopoda Telotha henselii
Gephyrocharax valenciae Acanthocephala Quadrigyrus torquatus
Gerres brasiliani Isopoda Cymothoa gerris
Glanidium melanopterum Monogenoidea Scleroductus spp.
Glanidium neivai Nematoda Rhabdochona acuminata
Glanidium sp. Cestoda Brayela karuatayi
Gymnocorymbus ternetzi Monogenoidea Diaphorocleidus armillatus
Gymnorhamphichthys hypostomus Trematoda Amazonadistoma negrensis
Gymnotus carapo Monogenoidea Urocleidoides carapus
Urocleidoides gymnotus
Hemiancistris sp. Monogenoidea Unilatus dissimilis
Hemiancistrus scaphirhynchae Monogenoidea Unilatus scaphirhynchae
Hemigrammus microstomus Monogenoidea Urocleidoides microstomus
Hemidoras carinatus Isopoda Vanamea symmetrica
Hemiodus microlepis Trematoda Rondotrema microvitellarium
Hemiodus orthonops Nematoda Procamallanus (S.) paraguayensis
Hemiodus (=Hemiodopsis) semitaeniatus Monogenoidea Cleidodiscus microcirrus
Hemiodus (=Hemiodopsis) semitaeniatus Monogenoidea Monocleithrium lavergneage

Hemisorubim platyrhynchus Monogenoidea Vancleavius platyrhynchi
Hemisorubim sp. Branchiura Dolops discoidalis
Holobrycon pesu Copepoda Ergasilus holobryconis
Hoplerythrinus unitaeniatus Nematoda Guyanema seriei
Guyanema seriei paraguayensis
Procamallanus (S.) krameri
Acanthocephala Quadrigyrus brasiliensis
Quadrigyrus nickoli
Branchiura Dolops discoidalis
Hoplias lacerdae Nematoda Procamallanus (S.) inopinatus
Hoplias macrophthalmus Isopoda Paracymothoa tholoceps
Hoplias malabaricus Myxozoa Henneguya malabarica
Monogenoidea Gyrodactylus trairae
Urocleidoides eremitus
Nematoda Capillaria zederi
Capillostromgyloides sentinosa
Guyanema baudi
Klossinemella iheringi
Paraseuratum soaresi
Procamallanus (S.) iheringi
Acanthocephala Grasilisentis variabilis
Neoechinorhynchus paraguayensis
Quadrigyrus brasiliensis
Quadrigyrus torquatus
Hirudinea Myzobdella platense
Copepoda Bedsylernaea collaris
Ergasilus iheringi
Pindapixara tarira
Taurocherus tarangophilus
Branchiura Argulus carteri
Argulus spinulosus
Argulus violaceus
Dolops discoidalis
Dolops geayi
Dolops reperta
Dolops striata
Isopoda Braga patagonica
Telotha henselii
Hydrolycus pectoralis Copepoda Brasergasilus mamorensis
Hydrolycus scomberoides Copepoda Ergasilus hydrolycus

Hydrolycus sp. Trematoda Glandulorhynchus turgidus
Hyphessobrycon callistus Isopoda Paracymothoa parva
Hyphessobrycon strictus Monogenoidea Urocleidoides strictus
Hypophthalmus edentatus Nematoda Paracamallanus amazonensis
Copepoda Ergasilus hypophthalmi
Prehendorastrus bidentatus
Prehendorastrus monodontus
Hypophthalmus fimbriatus Copepoda Prehendorastrus bidentatus
Prehendorastrus monodontus
Hypopomus sp. Trematoda Micramphistoma ministoma
Hypoptopoma thoracathum Monogenoidea Trinigyrus tentaculoides
Hypostomus albopunctatus Nematoda Procamallanus (S.) annipetteae
Raphidascaris (S.) mahnerti
Hypostomus auroguttatus Acanthocephala Gracilisentis variabilis
Hypostomus bolivianus Monogenoidea Unilatus anoculus
Hypostomus carinatus Acanthocepala Gorytocephalus elongorchis
Hypostomus commersonii Trematoda Gonocercella magnifica
Nematoda Raphidascaris (S.) mahnerti
Branchiura Argulus violaceus
Hypostomus derbyi Nematoda Raphidascaris (S.) mahnerti
Hypostomus lituratus Acanthocephala Gracilisentis variabilis
Hypostomus marginatus Monogenoidea Trinigyrus mourei
Hypostomus melanopterus Acanthocephala Gracilisentis variabilis
Hypostomus plecostomus Trematoda Eocreadium intermedium
Gonocercella magnifica
Megacoelium plecostomi
Acanthocephala Gorytocephalus plecostomorum
Gracilisentis variabilis
Hypostomus punctatus Monogenoidea Phanerothecium spinatus
Trematoda Kalitrema kalitrema
Hypostomus robinii Monogenoidea Trinigyrus hypostomatis
Unilatus unilatus
Hypostomus sp. Monogenoidea Heterotylus heterotylus
Unilatus brittani
Nematoda Raphidascaris (S.) hypostomi
Isopoda Braga nasuta
Braga fluviatilis
Iheringichthys labrosus Monogenoidea Demidospermus labrosis
Demidospermus mandi
Pseudovancleaveus paranaensis
Pseudovancleaveus platensis
Iheringichthys labrosus Trematoda Crepidostomum platense

Sanguinicola coelomicola
Isopoda Telotha silurii
Iheringichthys westermanni Monogenoidea Demidospermus cornicinus
Demidospermus leptosynophallus
Laemolyta taeniata Copepoda Ergasilus triangularis
Lahilliela (=Schizodon) kneri Trematoda Paralecithobotrys brasiliensis
Lebiasina bimaculata Monogenoidea Gyrodactylus bimaculatus
Gyrodactylus lebiasinus
Gyrodactylus slendrus
Lebiasina multimaculata Nematoda Procamallanus (S.) inopinatus
Leiarius marmoratus Branchiura Dolops discoidalis
Lepidosiren paradoxa Myxozoa Agarella gracilis
Trematoda Kalipharynx piramboae
Leporellus vittatus Isopoda Riggia brasiliensis
Leporinodus vittatus Trematoda Saccocoelioides leporinodus
Nematoda Acyracanthus schubarti
Rhabdochona acuminata
Liporinus agassizii Monogenoidea Rhinoxenus euryxenus
Leporinus copelandi Trematoda Creptotrema lynchi
Nematoda Acyracanthus schubarti
Cucullanus mogi
Cucullanus pinnai
Isopoda Riggia brasiliensis
Leporinus elongatus Trematoda Creptotrema creptotrema
Paralecithobotrys brasiliensis
Procamallanus (S.) amarali
Leporinus fasciatus Monogenoidea Tereancistrum parvus
Trematoda Prosthenhystera obesa
Copepoda Ergasilus leporinidis
Brasergasilus guaporensis
Branchiura Dolops striata
Leporinus fasciatus Isopoda Livoneca guianensis

Nerocila armata
Leporinus friderici Nematoda Dichelyne leporine
Leporinus lacustris Monogenoidea Kritskyia eirasi
Nematoda Procamallanus (S.) inopinatus
Leporinus mormyrops Myxozoa Henneguya leporini
Myxobolus associatus
Leporinus muyscorum Trematoda Chalcinotrema lucieni
Leporinus obtusidens Trematoda Saccocoelioides magniovatus
Leporinus octofasciatus Trematoda Creptotrema lynchi
Nematoda Ancyracanthus schubarti
Isopoda Riggia brasiliensis
Leporinus piau Nematoda Procamallanus (S.) inopinatus
Leporinus reinhardti Nematoda Procamallanus (S.) inopinatus
Leporinus striatus Isopoda Riggia nana
Leporinus taeniatus Nematoda Procamallanus (S.) inopinatus
Leporinus sp. Trematoda Creptotrema creptotrema
Nematoda Capillaria minima
Capillostrongylus sentinosa
Cucullanus mogi
Procamallanus (S.) amarali
Loricaria anus Monogenoidea Demidospermus anus
Trematoda Saccocoelioides quintus
Loricaria laticeps Nematoda Raphidascaris (S.) mahnerti
Loricaria sp. Myxozoa Henneguya occulta
Trematoda Halipegus tropicus
Loricaria vetula Trematoda Procaudotestis uruguayensis
Loricariichthys brunneus Nematoda Guyanema longispiculum
Raphidascaris (S.) mahnerti
Loricariichthys platymetopon Nematoda Sprentascarus mahnerti
Loricariidae Isopoda Asotana splendida
Luciopimelodus (=Pimelodus) pati Nematoda Cucullanus pinnai
Cucullanus zungaro
Luciopimelodus (=Pimelodus) pati Nematoda Dichelyne moraveci

Philometra alii
Rondonia rondoni
Branchiura Dipteropeltis hirundo
Lycengraulis grossidens Copepoda Acusicola lycengraulidis
Ergasilus euripedesi
Isopoda Excorallana berbicensis
Lycengraulis sp. Acanthocephala Neoechinorhynchus macronucleatus
Markiana geayi Nematoda Procamallanus (S.) inopinatus
Megalancistrus brasiliensis Nematoda Ichthyouris brasilienis
Megalodoras irwini Trematoda Doradamphistoma bacuensis
Megalodoras sp. Branchiura Dolops geayi
Megalonema platinum Nematoda Cucullanus pinnai
Dichelyne moraveci
Menticirrhus litoralis Isopoda Cymothoa catarinensis
Metynis maculates Isopoda Isonebula maculatus
Monochir maculipennis (=Achirus achirus) Acanthocephala Gracilisentis variabilis
Mylesinus paraschomburgkii Monogenoidea Notozothecium bethae
Trematoda Alphamphistoma canoeforma
Betamphistoma jariense
Deltamphistoma pitingaense
Gammamphistoma collaris
Saccocoelioides rotundus
Zetamphistoma compacta
Rondonia rondoni
Mylesinus paucisquamatus Monogenoidea Notozothecium bethae
Myletes edulus Nematoda Rondonia rondoni
Myleus (Myloplus)asterias Trematoda Annelamphistoma elegans
Dadayius pacuensis
Inpamphistoma papillatum
Myleus pacu Monogenoidea Notozothecium bethae
Myleus rhomboidalis Monogenoidea Notozothecium bethae
Myleus (Myloplus) rubripinnis Monogenoidea Anacanthorus hoplophallus
Anacanthorus pedanophallus
Anacanthorus spinatus
Anacanthorus stagmophallus
Notothecioides llewellyni
Trematoda Annelamphistoma elegans
Dadayius pacuensis
Myleus sp. Trematoda Curumai curumai
Dadaytrema elongata
Myleus sp. Nematoda Klossinemella iheringi

Myleusnema brasiliense
Rondonia rondoni
Myleus ternetzi Nematoda Myleusnema bicornis
Spinoxyuris annulata
Myleus torquatus Monogenoidea Notothecioides llewellyni
Nematoda Rondonia rondoni
Mylossoma aureum Trematoda Dadayius marenzelleri
Microrchis megacotyle
Pseudocladorchis nephrodorchis
Mylossoma paraguayense Acanthocephala Echinorhynchus salobrense
Mylossoma duriventris Monogenoidea Anacanthorus paraspathulatus
Nannostomus beckfordi Isopoda Artystone minima
Odonthestes (= Basilichthys) bonariensis Branchiura Argulus violaceus
Odontostilbe paraguayensis Isopoda Riggia cryptocularis
Onchorhynchus mykiss Isopoda Anilocra montti
Ossubtus xinguense Isopoda Anphira xinguensis
Osteoglossum bicirrhosum Monogenoidea Gonocleithrum aruanae
Gonocleithrum coenoideum
Gonocleithrum cursitans
Gonocleithrum planacroideum
Gonocleithrum planacrus
Telethecium nasalis
Trematoda Caballerotrema aruanense
Nematoda Camallanus acaudatus
Otocinclus vestitus Isopoda Artystone bolivianensis
Oxydoras kneri Nematoda Spinoxyuris oxydoras
Oxydoras niger (=Pseudodoras niger) Monogenoidea Cosmetocleithrum confusus
Cosmetocleithrum gussevi
Cosmetocleithrum parvum
Cosmetocleithrum rarum
Cosmetocleithrum sobrinus
Cestoda Proteocephalus kuyukuyu
Nematoda Cucullanus grandistomis
Acanthocephala Paracavisoma impudica
Branchiura Dolops longicauda
Pachyurus bonariensis Nematoda Spinitectus pachyuri
Pachyurus junki Monogenoidea Euryhalotrema dontycoleos
Pacu nigricans Acanthocephala Echinorhynchus gomezi
Parapimelodus valenciennis Monogenoidea Demidospermus valenciennesi
Parauchenipterus galeatus Monogenoidea Demidospermus uncusvalidus
Parauchenipterus striatulus Monogenoidea Scleroductus spp.

Paulicea lütkeni Monogenoidea Unibarra paranoplatensis
Nematoda Cucullanus paulicea
Cucullanus pinnai
Cucullanus schubarti
Cucullanus zungaro
Pellona castelnaeana Trematoda Bacciger pellonae
Copepoda Acusicola pellonidis
Branchiura Dolops carvalhoi
Pellona flavipinnis Monogenoidea Telethecium paniculum
Percichthys melanops Acanthocephala Pomphorhynchus yamagutii
Percichthys trucha Monogenoidea Acolpenteron australe
Duplaaccessorius andinus
Branchiura Argulus patagonicus
Hirudinea Theromyzon propinquum
Petenia splendida Monogenoidea Sciadicleithrum bravohollisae
Sciadicleithrum splendidae
Phractocephalus hemiliopterus (=Pirarara bicolor) Monogenoidea Urocleidoides amazonensis
Urocleidoides catus
Vancleaveus cicinnus
Cestoda Ephedrocephalus microcephalus
Myzophorus pirarara
Zygobothrium megacephalum
Branchiura Dolops carvalhoi
Dolops discoidalis
Piabucina sp. Nematoda Rhabdochona uruyeni
Piaractus brachypomus (=Colossoma bidens) Ciliophora Nyctotherus piscicola
Monogenoidea Mymarothecium viatorum
Rondonia rondoni
Spectatus spectatus
Acanthocephala Echinorhynchus jucundum
Copepoda Perulernaea pirapitingae
Piaractus mesopotamicus Monogenoidea Anacanthorus penilabiatus
Pimelodella albicans Nematoda Cucullanus pinnai
Pimelodella gracilis Nematoda Cucullanus pinnai
Pimelodella lateristriga Nematoda Brasilnema pimelodellae
Cucullanus pimelodellae
Procamallanus (S.) pimelodus
Procamallanus (S.) rarus
Spinitectus rudolphiheringi
Spinitectus yorkei
Pimelodella laticeps Monogenoidea Aphanoblastella travassosi

Pimelodella maculates Nematoda Cucullanus pinnai
Pimelodella sp. Myxozoa Myxobolus stokesi
Monogenoidea Scleroductus spp.
Nematoda Spinitectus multipapillata
Pimelodella yuncensis Monogenoidea Gyrodactylus pimelodellus
Scleroductus yuncensi
Pimelodidae Acanthocephala Acanthodelta scorzai
Pimelodus albicans Myxozoa Myxobolus sp.
Monogenoidea Cosmetocleithrum longivaginatum
Demidospermus armostatus
Demidospermus bidiverticulatum
Demidospermus idolus
Demidospermus majusculus
Vancleaveus cicinnus
Pimelodus clarias Myxozoa Myxobolus cunhai
Myxobolus inaequalis
Ciliophora Balantidium piscicola
Nyctotherus piscicola
Zelleriella piscicola
Monogenoidea Demidospermus armostus
Demidospermus paravalenciennesi
Demidospermus uncusvalidus
Trematoda Crepidostomum platense
Halipegus tropicus
Parspina argentinensis
Nematoda Cucullanus pinnai
Philometra baylisi
Rondonia rondoni
Procamallanus (S.) rarus
Procamallanus (S.) pimelodus
Isopoda Livoneca guianensis
Telotha henselii
Pimelodus grosskopfii Monogenoidea Urocleidoides lebedevi
Pimelodus maculatus Monogenoidea Ameloblastella platensis
Paramphocleithrium bidiverticulatum
Pseudovancleaveus platensis
Unibarra paranoplatensis
Nematoda Cucullanus debacoi
Cucullanus fabregasi
Pimelodus maculatus Cucullanus patoi
Pimelodus maculatus Nematoda Cucullanus riograndensis

Procamallanus (S.) freitasi
Spinitectus sternopygi
Raphidascaris (S.) pimelodi
Pimelodus megacephalus Trematoda Dadaytrema oxycephala
Pseudoclardorchis cylindricus
Pimelodus ornatus Trematoda Pseudocladorchis cylindricus
Pimelodus ortmanni Nematoda Procamallanus (S.) pimelodus
Pimelodus (=Luciopimelodus) pati Cestoda Monticellia piracatinga
Nominoscolex piracatinga
Proteocephalus fossatus
Rudolphiella lobosa
Pimelodus sp. Myxozoa Myxobolus stokesi
Pinirampus pirinampu Monogenoidea Demidospermus luckyi
Demidospermus pinirampi
Cestoda Myzophorus admonticellia
Rudolphiella myoides
Rudolphiella piranabu
Nematoda Philometra alii
Pirarara bicolor (=Phractocephalus hemioliopterus) Cestoda Zygobothrium megacephalum
Plagioscion sp. Monogenoidea Diplectanum hilum
Diplectanum pescadae
Plagioscion squamosissiumus Monogenoidea Diplectanum decorum
Diplectanum gymnopeus
Diplectanum pescadae
Diplectanum piscinarius
Euryhaliotrema chaoi
Euryhaliotrema lovejoyi
Euryhaliotrema monacanthus
Euryhaliotrema potamocetes
Euryhaliotrema succedaneus
Euryhaliotrema thatcheri
Trematoda Brasicystis bennetti
Acanthocephala Rhadinorhynchus plagioscionis
Copepoda Therodamas elongatus
Therodamas tamarae
Branchiura Dolops sp.
Platydoras costatus Cestoda Proteocephalus renaudi
Proteocephalus soniae
Platystoma sp. (=Sorubim lima) Cestoda Manaosia bracodemoca
Platystomatichthys sturio Cestoda Monticellia megacephala
Platystomatichthys sturio Cestoda Nominoscolex lenha

Ornithoscolex lenha
Peltidocotyle lenha
Spasskyellina lenha
Woodlandiella myzophora
Plecostomus bolivianus Monogenoidea Unilatus anoculus
Plecostomus plecostomus Monogenoidea Phanerothecium harrisi
Plecostomus sp. Monogenoidea Unilatus brittani
Unilatus unilatus
Pleuronectes sp. Acanthocephala Gracilisentis variabilis
Poecilia caucana Monogenoidea Gyrodactylus poeciliae
Gyrodactylus milleri
Poecilia reticulata Monogenoidea Gyrodactylus bullatarudis
Gyrodactylus tumbulli
Urocleidoides minuta
Urocleidoides reticulatus
Poecilia sphenops Monogenoidea Gyrodactylus bullatarudis
Gyrodactylus costaricensis
Poecilia vivipara Myxozoa Myxobolus lutzi
Pogonias chromis Isopoda Braga fluviatilis
Pogonias sp. Isopoda Braga patagonica
Potamorhina latior Copepoda Miracetyma etimaruya
Potamotrygon circularis Monogenoidea Paraheteronchocotyle amazonensis
Potamotrygonocotyle tsalickisi
Cestoda Acanthobothrium amazonensis
Potamotrygonocestus amazonensis
Rhinebothroides circularisi
Nematoda Echinocephalus daileyi
Potamotrygon falkneri Cestoda Eutetrarhynchus araya
Rhinebothrium paratrygoni
Potamotrygon hystrix Cestoda Acanthobothrium regoi
Eutetrarhynchus araya
Rhinebothrium paratrygoni
Rhinebothroides freitasi
Rhinebothroides glandularis
Rhinebothroides venezuelensis
Nematoda Echinocephalus daileyi
Terranova diazungriai
Acanthocephala Megapriapus ungriai
Potamotrygon magdalenae Cestoda Acanthobothrium quinonesi
Potamotrygonocestus magdalenensis
Rhinebothroides moralarai
Potamotrygon motoro Cestoda Acanthobothrium terezae
Potamotrygon motoro Cestoda Eutetrarhynchus araya

Nematoda Brevimulticaecum regoi
Potamotrygon reticulatus Cestoda Eutetrarhynchus araya
Potamotrygonocestus amazonensis
Potamotrygonocestus orinocoensis
Rhinebothrium paratygoni
Potamotrygon yepezi Cestoda Potamotrygonocestus amazonensis
Rhinebothroides venezuelensis
Pristis perotteti Cestoda Anthobothrium pristis
Pristobrycon eigenmanni Monogenoidea Amphithecium minutum
Amphithecium muricatum
Amphithecium verecundum
Anacanthorus beleophallus
Anacanthorus gravihamulatus
Anacanthorus jegui
Anacanthorus mastigophallus
Anacanthorus mesocondylus
Anacanthorus xaniophallus
Enallothecium comutum
Mymarothecium galeolum
Notozothecium teinodendrum
Pristobrycon sp. Monogenoidea Amphithecium diclonophallum
Amphithecium falcatum
Amphithecium minutum
Amphithecium pretiosum
Anacanthorus jegui
Anacanthorus palamophallus
Anacanthorus serrasalmi
Anacanthorus xaniophallus
Enallothecium segidatum
Mymarothecium dactylotum
Mymarothecium goleolum
Notozothecium foliolum
Pristobrycon striolatus Monogenoidea Amphithecium prodotum
Anacanthorus cinctus
Anacanthorus crytocaulus
Anacanthorus lasiophallus
Enallothecium variabilum
Pristobrycon striolatus Monogenoidea Notozothecium robustum

Trematoda Anavilhanatrema robusta
Prochilodus lineatus Monogenoidea Kritskyia boegeri
Rhinoxenus curimbatae
Trematoda Saccocoelioides nanii
Prochilodus nigricans Monogenoidea Rhinonastes pseudocapsaloideum
Trematoda Lecithobotrioides elongatus
Copepoda Ergasilus urupaensis
Dolops bidentata
Prochilodus platensis Trematoda Sanguinicola argentinensis
Prochilodus reticulatus Monogenoidea Anacanthoroides mizellei
Tereancistrum omatus
Trematoda Colocladorchis ventrastomis
Lecithobotrioides mediacanoensis
Unicoelium prochilodorum
Nematoda Spinitectus asperus
Acanthocephala Neoechinorhynchus prochilodorum
Prochilodus platensis Nematoda Spinitectus asperus
Prochilodus scrofa Nematoda Spinitectus asperus
Psectrogaster essequibensis Copepoda Miracetyma etimaruya
Psectrogaster rutiloides Monogenoidea Anacanthorus amazonicus
Annulotrematoides amazonicus
Pseudauchenipterus nodosus Isopoda Nerocila armata
Pseudocurimata (=Cyphocharax) plumbea Nematoda Travnema travnema
Pseudocurimata gilberti Monogenoidea Androspira chascomusensis
Androspira triangula
Curvianchoratus singularis
Pseudodoras niger Nematoda Cucullanus grandistomis
Acanthocephala Paracavisoma impudica
Pseudopimelodus roosevelti Trematoda Iheringtrema iheringi
Pseudopimelodus (=Cephalosilurus) zungaro Myxozoa Henneguya lutzi
Monogenoidea Phanerothecium caballeroi
Urocleidoides mamaevi
Trematoda Prosthenhystera obesa
Cestoda Amphoteromorphus parkarmoo
Nematoda Cucullanus oswaldocruzi
Cucullanus pinnai
Cucullanus zungaro
Rondonia rondoni
Pseudoplatisma sp. Monogenoidea Amphocleithrium paraguayensis
Pavanelliella pavanelli
Pseudoplatystoma corruscans Branchiura Argulus silvestrii

Nematoda Cucullanus pseudoplatystomae
Goezia brasiliensis
Pseudoplatystoma fasciatum Myxozoa Henneguya linearis
Monogenoidea Vancleaveus fungulus
Cestoda Monticellia rugosa
Myzophorus sorobim
Nominoscolex sorobim
Nominoscolex woodlandi
Spasskyellina spinulifera
Nematoda Dichelyne moraveci
Argulus pestifer
Dolops carvalhoi
Dolops discoidalis
Pseudoplatystoma tigrinum Monogenoidea Vancleavius fungulus
Trematoda Witenbergia witenbergia
Cestoda Monticellia surubim
Nominoscolex kaparari
Peltidocotyle rugosa
Argulus pestifer
Dolops carvalhoi
Dolops discoidalis
Pseudoplatystoma sp. Monogenoidea Amphocleithrium paraguayensis
Cestoda Monticellia surubim
Proteocephalus platystomi
Nematoda Cucullanus pinnae
Contracaecum sp. (larvae)
Eustrongylides sp. (larvae)
Branchiura Argulus nattereri
Pseudotylosurus angusticeps Copepoda Acusicola cunula
Pterodoras granulosus Monogenoidea Cosmetocleithrum bulbocirrus
Vancleavius janauacaensis
Pseudocladorchis ferrumequinum
Neoparaseuratum travassosi
Paracamallanus brasiliensis
Raphidascaroides brasiliensis
Rondonia rondoni
Pterodoras granulosus Acanthocephala Neoechinorhynchus pterodoridis

Branchiura Dolops longicauda
Pterophyllum scalare Monogenoidea Gussevia spiralocirrus
Sciadicleithrum iphthimum
Pterygoplichthys aculeatus Nematoda Ichthyyouris brasiliensis
Pterygoplichthys multiradiatus Monogenoidea Unilatus brevispinus
Unilatus longispinus
Pterygoplichthys pardalis Trematoda Megacoelium spinicavum
Pterygoplichthys sp. Trematoda Megacoelium spinispecum
Pygidium (=Trichomycterus) brasiliensis Nematoda Procamallanus (S.) pexatus
Pygidium (=Trichomycterus) punctulatum Nematoda Procamallanus (S.) chimusensis
Procamallanus (S.) incarocai
Pygocentrus nattereri Monogenoidea Amphithecium brachycirrum
Amphithecium calycinum
Amphithecium camelum
Amphithecium catalaoensis
Amphithecium junki
Amphithecium microphallum
Anacanthorus anacanthorus
Anacanthorus brazilensis
Anacanthorus maltai
Anacanthorus neotropicalis
Anacanthorus reginae
Anacanthorus rondonensis
Anacanthorus stachophallus
Anacanthorus thatcheri
Calpidothecioides orthus
Calpidothecium crescentis
Calpidothecium serrasalmus
Cleidodiscus serrasalmus
Enallothecium aegidatum
Notothecium mizellei
. Notozothecium minor
Notozothecium penetrarum
Pithanothecium amazonensis
Rhinoxenus piranhus
Copepoda Rhinergasilus piranhas
Miracetyma piraya
Pygocentrus sp. Nematoda Procamallanus (S.) inopinatus
Pygopristis denticulata Monogenoidea Calpidothecioides pygopristi

Calpidothecium crescentis
Calpidothecium serrasalmus
Pithanothecium amazonensis
Rhamdia guatemalensis Monogenoidea Ameloblastella chavarriai
Aphanoblastella travassosi
Rhamdia quelen Myxozoa Henneguya sp.
Myxobolus sp.
Monogenoidea Ameloblastella chavarriai
Kritskyia moraveci
Scleroductus spp.
Trematoda Acanthostomum gnerii
Nematoda Procamallanus (S.) hilarii
Copepoda Ergasilus thatcheri
Rhamdia rogersi Monogenoidea Ameloblastella chavarriai
Trematoda Acanthostomum gnerii
Rhamdia sapo Monogenoidea Aphanoblastella mastigatus
Nematoda Hysterothylacium rhamdiae
Hirudinea Myzobdella uruguayensis
Rhamdia sebae Myxozoa Henneguya linearis
Monogenoidea Ameloblastella chavarriai
Rhamdia sp. Monogenoidea Aphanoblastella robustus
Cestoda Megathylacus jandia
Proteocephalus jandia
Branchiura Argulus chromidis
Rhaphiodon vulpinus Nematoda Guyanema raphiodoni
Copepoda Miracetyma kawa
Dolops carvalhoi
Rhamphichthys rostratus Nematoda Cucullanus rhampichthydis
Copepoda Miracetyma kawa
Dolops carvalhoi
Rhinelepis aspera Nematoda Parasynodontisia petterae
Rhineloricaria sp. Monogenoidea Hyperopletes malmbergi
Rhinodoras d’orbignyi Trematoda Paleocryptogonimus claviformis
Rhinodoras d’orbignyi Nematoda Procamallanus (S.) rarus
Rhytiodus argenteofuscus Monogenoidea Rhinoxenus arietinus

Rhytiodus microlepis Monogenoidea Urocleidoides paradoxus
Dolops bidentata
Rivulus harti Monogenoidea Cycloplectanum americanum
Pseudorhabdosynochus sp.
Roeboides bonariensis Trematoda Crepidostomum macrorchis
Halipegus genarchella
Roeboides myersi Monogenoidea Anacanthorus dipelecinus
Saccodon caucae Trematoda Saccocoelioides magnorchis
Saccocoelioides saccodontis
Salminus affinis Monogenoidea Anacanthorus colombianus
Anacanthorus cuticulovaginus
Salminus brevidens Trematoda Prosthenystera obesa
Nematoda Agamonema sp. larvae
Copepoda Taurocheros salminisii
Branchiura Argulus nattereri
Argulus salmini
Dipteropeltis hirundo
Dolops longicauda
Salminus hilarii Trematoda Bellumcorpus major
Cladocystis intestinalis
Pararhipidocotyle jeffersoni
Nematoda Cystidicoloides fischeri
Neocucullanus neocucullanus
Philometroides maplestonei
Isopoda Braga patagonica
Salminus maxillosus Monogenoidea Rhinoxenus bulbovaginatus
Trematoda Cladocystis intestinalis
Dadaytrema oxycephala
Halipegus dubius
Pararhipidocotyle jeffersoni
Prosorhynchus schubarti
Nematoda Freitascapillaria maxillosa
Cucullanus interrogativus
Cystidicoloides fischeri
Salminus maxillosus Nematoda Neocucullanus neocucullanus

Philometra paraguayensis
Branchiura Argulus paranensis
Argulus paulensis
Argulus pestifer
Argulus salmini
Dipteropeltis hirundo
Dolops discoidalis
Dolops geayi
Dolops longicauda
Dolops nana
Salminus spp. Isopoda Braga fluviatilis
Braga patagonicus
Sargus sp. Isopoda Cymothoa brasiliensis
Scatophagus argus Monogenoidea Metahaliotrema scatophagi
Metahaliotrema yamagutii
Schizodon borelli Nematoda Cucullanus pinnai
Schizodon fasciatus Monogenoidea Rhinoxenus arietinus
Rhinoxenus nyttus
Trematoda Saccocoelioides szidati
Nematoda Dichelyne leporini
Acanthocephala Octospiniferoides incognita
Copepoda Gamispatulus schizodontis
Dolops striata
Schizodon knerii Nematoda Procamallanus (S.) inopinatus
Schizodon nasutum Isopoda Riggia brasiliensis
Semaprochilodus insignis Copepoda Brasergasilus jaraquensis
Ergasilus jaraquensis
Gamidactylus jaraquensis
Semaprochilodus taeniurus Monogenoidea Gyrodactylus gemini
Serrasalmus compressus Monogenoidea Amphithecium diclonophallum
Enallothecium cornutum
Enallothecium umbelliferum
Serrasalmus elongatus Monogenoidea Amphithecium diclonophallum
Serrasalmus elongatus Monogenoidea Amphithecium falcatum

Amphithecium speirocamarotum
Anacanthorus lepyrophallus
Anacanthorus prodigiosus
Anacanthorus ramosissimus
Anacanthorus sciponophallus
Notothecium deleastum
Serrasalmus gouldingi Monogenoidea Amphithecium diclonophallum
Amphithecium minutum
Heterothecium globatum
Notothecium circellum
Rhinoxenus euryxenus
Serrasalmus manuelli Nematoda Procamallanus (S.) inopinatus
Monogenoidea Amphithecium falcatum
Serrasalmus marginatus Monogenoidea Kritskyia annakohnae
Serrasalmus (=Pygocentrus) nattereri Monogenoidea Amphithecium brachycirrum
Amphithecium calycinum
Amphithecium camelum
Amphithecium catalaoensis
Amphithecium junki
Anacanthorus anacanthorus
Anacanthorus brazilensis
Anacanthorus maltai
Anacanthorus neotropicalis
Anacanthorus reginae
Anacanthorus rondonensis
Serrasalmus (=Pygocentrus) nattereri Monogenoidea Anacanthorus thatcheri

Cleidodiscus amazonensis
Notothecium aegidatum
Notothecium mizellei
Notozothecium penetrarum
Notozothecium minor
Copepoda Rhinergasilus piranhus
Argulus sp.
Dolops bidentata
Dolops carvalhoi
Dolops longicauda
Isopoda Amphira branchialis
Serrasalmus (=Pygocentrus) piraya Myxozoa Myxobolus cunhai
Myzobolus pygocentris
Serrasalmus rhombeus Myxozoa Myxobolus serrasalmi
Monogenoidea Amphithecium diclonophallum
Amphithecium junki
Anacanthorus amazonicus
Anacanthorus jegui
Enallothecium cornutum
Mymarothetcium whittingtoni
Notothecium cyphophallum
Notothecium phyleticum
Trematoda Prosorhynchus piranhus
Serrasalmus spilopleura Myxozoa Henneguya iheringi
Myxobolus noguchii
Monogenoidea Amphithecium falcatum
Serrasalmus spilopleura Monogenoidea Amphithecium minutum

Amphithecium unguiculum
Anacanthorus cladophallus
Anacanthorus jegui
Anacanthorus scapanus
Kritskyia annakohnae
Mymarothecium perplanum
Notothecium modestum
Vanamea symmetrica
Serrasalmus sp. Monogenoidea Amphithecium diclonophallum
Amphithecium microphallum
Amphithecium verecundum
Anacanthorus amazonicus
Anacanthorus jegui
Anacanthorus lepyrophallus
Anacanthorus periphallus
Mymarothecium whittingtoni
Notothecium deleastoideum
Notothecium reduvium
Asotana magnifica
Braga patagonica
Vanamea symmetrica
Silurus palmito (=Ageneiosus sp.) Trematoda Microrchis megacotyle
Silurus sp. Cestoda Choanoscolex abscisus
Silurus sp. Cestoda Monticellia coryphicephala

Sorubim lima (see Platystoma sp.) Monogenoidea Urocleidoides megorchis
Sphaeroides testudineus Ciliophora Trichodina fariai
Steindachneridion prahybae Nematoda Cucullanus pinnai
Stenarchus (=Apteronotus) brasiliensis Isopoda Telotha lunaris
Sternopygus macrurus Nematoda Cystidicoloides dlouhyi
Strongylura fluviatilis Copepoda Ergasilus argulus
Strongylura scapularis Copepoda Ergasilus argulus
Srongylura incisa Copepoda Ergasilus orientalis
Strongylura sp. Copepoda Acusicola tenax
Symphysodon discus Monogenoidea Sciadicleithrum variabilum
Synbranchus marmoratus Nematoda Agamonema sp. (larvae)
Acanthocephala Quadrigyrus torquatus
Tetragonopterus argenteus Nematoda Paraseuratum albidum
Tetragonopterus chalceus Nematoda Procamallanus (S.) saofranciscencis
Tetragonopterus sp. Nematoda Rhabdochona elegans
Thoracocharax sternicla Nematoda Procamallanus (S.) inopinatus
Tilapia mossambica Monogenoidea Cichlidogyrus sclerosus
Cichlidogyrus tilapiae
Trachydoras insignis Nematoda Procamallanus (S.) penneri
Trachydoras paraguayensis Nematoda Ichthyouris laterifilamenta
Neoparaseuratum travassosi
Rondonia rondoni
Triportheus albus Monogenoidea Anacanthorus acuminatus
Anacanthorus alatus
Anacanthorus bellus
Anacanthorus euryphallus
Anacanthorus quinqueramis
Anacanthorus ramulosus
Ancistrohaptor falciferum
Ancistrohaptor falcunculum
Isopoda Anphira junki
Triportheus angulatus Monogenoidea Anacanthorus acuminatus
Anacanthorus andersoni
Anacanthorus carinatus
Anacanthorus chaunophallus
Anacanthorus chelophorus
Anacanthorus cornutus
Triportheus angulatus Monogenoidea Anacanthorus glyptophallus

Anacanthorus lygophallus
Anacanthorus nanus
Anacanthorus pithophallus
Anacanthorus tricornis
Acanthocephala Palliolisentis quinqueungulis
Triportheus elongatus Monogenoidea Anacanthorus acuminatus
Anacanthorus alatus
Anacanthorus bellus
Anacanthorus calophallus
Anacanthorus formosus
Anacanthorus furculus
Anacanthorus pelorophallus
Anacanthorus quinqueramis
Anacanthorus ramulosus
Anacanthorus strongylophallus
Anacanthorus tricornis
Ancistrohaptor falcatum
Branchiura Dolops sp.
Triportheus flavus Isopoda Anphira junki
Triportheus cf. nematurus Monogenoidea Rhinoxenus anaclaudiae
Triportheus paranensis Trematoda Chalcinotrema salobrensis
Creptotrema dispar
Nematoda Spinitectus sternopygi
Acanthocephala Echinorhynchus paranense
Palliolisentis ornatus
Palliolisentis quinqueungulis
Triportheus sp. Monogenoidea Ancistrohaptor falciferum
Nematoda Procamallanus (S.) barroslimai
Typhlias pearsei Nematoda Rhabdochona kidderi
Uaru amphiacanthoides Monogenoidea Gusseva elephus
Gusseva obtusa
Sciadicleithrum tortrix
Vandelia cirrhosa Isopoda Vanamea symmetrica
Xiphophorus helleri Monogenoidea Gyrodactylus rasini
Urocleidoides vaginoclaustrum
Xiphophorus hybrids Monogenoidea Gyrodactylus rasini

Xiphostomum (=Boulengerella) cuvieri Trematoda Prosthenhystera obesa
Other hosts Phylum or class Parasite

Plankton Copepoda Anklobrachius marajoensis
Vaigamus retrobarbatus
Vaigamus spinicephalus
13.
SUBJECT INDEX 1
Abergasilinae, 337 Alphamphistoma, 124, 128, 145, 185, 195, 478

Acanthicus hystrix, 80, 465 Amazolernaea, 334, 336, 343, 377, 469, 470
Acanthobothrium, 209, 211, 219, 222, 483, 484 Amazonadistoma, 126, 130, 138, 155, 473
Acanthocephala, 5, 7, 15, 16, 20, 21, 269, 299, 302, Amazonicopeus, 329, 330
303, 305, 311, 324 Ameloblastella, 55, 59, 64, 96, 465, 481, 488
Acanthocheilidae, 239, 244 Amphilinidae, 209, 211, 213
Acanthodelta, 302, 303, 305, 315, 481 Amphilinidea, 206, 213
Acanthogyridae, 302, 303, 305 Amphilophus alfari, 86
Acanthostomidae, 118, 119, 126, 130, 131 Amphithecium, 43, 47, 53, 58, 65, 96, 469, 484,
Acanthostomum, 126, 130, 154, 488 487, 490, 491, 492, 493
Accessorius, 51,56,61,93 Amphoteromorphus, 209, 211, 213, 223, 224, 468, 485
Acestrorhamphus, 30, 134, 135, 139, 264 Amplexibranchius, 328, 338, 367, 368, 468
Acestrorhynchus 154, 308, 387, 423, 465 Amplexibranchus, 333, 335
falcatus, 308, 465 Anacanthocotyle, 51, 55, 61, 62, 94, 467
guyanensis, 423, 465 Anacanthoroides, 51, 56, 66, 96, 485
lacustris, 254, 465 Anacanthorus, 43, 51, 56, 66, 67, 96,
Achiurus mazatlanus, 307 Anavilhanatrema, 124, 128, 145, 146, 195, 485
Acnodon 82, 422, 465 Ancistrohaptor, 54, 59, 69, 98, 494, 495
normani, 82, 465 Ancistrus, 63, 254, 423, 425, 452, 466
oligacanthus, 422, 465 cirrhosus, 423, 466
Acolpenteron, 47, 480 sp., 68, 254, 425, 466
Acusicola, 328, 331, 333, 335, 338, 353, 354, 355, Angiodictyidae, 118, 123, 127, 132
356, 385, 470, 478, 480, 486, 494 Anisakidae, 238, 243, 254, 255
Acusicolinae, 327, 333, 335, 337, 338, 340 Annelamphistoma, 124, 128, 146, 189, 193, 195, 478
Aequidens, 86, 131, 307, 399, 465, 466 Annelida, 20, 454
maroni, 86, 465 Annulotrematoides, 55, 59, 70, 98, 468, 485
pulcher, 131, 307, 399, 466 Anodus, 337, 359, 360, 466
Agarella, 28, 29, 34, 476 elongatus, 337, 466
Ageneiosus brevifilis, 214, 215, 342, 426, 466 Anphira, 420, 421, 422, 429, 434, 441, 442, 446,
Alaria, 120 465, 479, 494, 495
Allocreadiidae, 118, 126, 130, 131 Anthobothrium, 209, 211, 219, 222, 227, 466, 483, 484
1
Figures in bold
Antorchis, 125, 129, 137, 160, 467 Boulengerella, 135, 143, 144, 468, 496
Aphanoblastella, 43, 53, 54, 57, 58, 70, 481, 488 lucia, 143, 144, 468
Arapaima, 73, 136, 206, 208, 213, 235, 236, 249, Brachyplatystoma, 214, 215, 216, 424, 425, 426
255, 259, 309, 310, 399, 466 filamentosum, 214, 216, 468
gigas, 73, 136, 206, 213, 235, 236, 249, 255, rousseauxii, 216, 468,
259, 309, 310, 399, 466 sp., 424, 425, 426, 468
Archocentrus nigrofasciatus, 86 vaillanti, 215, 468
Argulidae, 397 Braga, 418, 420, 421, 423, 429, 431, 432, 433, 437,
Argulus, 339, 357, 390, 391, 392, 394, 397, 401, 438, 442, 465, 466, 468, 470, 472,
402, 403, 404, 405, 406, 411, 466, 474, 475, 477, 482, 483, 489, 490, 493
467, 468, 469, 470, 471, 472, 473, Branchiura, 5, 20, 21, 390, 392, 394, 397, 401, 466,
474, 475, 477, 479, 480, 481, 485, 467, 468, 469, 470, 471, 472, 473,
486, 487, 488, 489, 490, 492, 494 474, 475, 475, 477, 478, 479, 480,
Ariopsis seemanni, 310, 466 481, 482, 485, 486, 487, 488, 489,
Arthropoda, 20, 458 490, 492, 494, 495
Artystone, 418, 420, 421, 422, 427, 430, 445, 472, Brasergasilus, 328, 331, 333, 335, 337, 358, 360,
473, 479 369, 383, 466, 475, 476, 490
Ascaridoidea, 238, 242, 243 Brasicystis, 119, 123, 127, 135, 156, 180, 482
Ascaroidea, 235, 254 Brasilnema, 239, 244, 257, 292, 480
Ascocotyle, 120, 180, 469 Brayela, 209, 211, 214, 227, 473
Asotana, 418, 420, 421, 430, 433, 435, 436, 441, Brevimulticaecum, 239, 484
477, 493 Brevoortia pectinata, 339, 468
Astronotus ocellatus, 77, 235, 255, 399, 459, 466, 467 Brycon, 34, 37, 67, 68, 70, 78, 85, 87, 138, 235, 251,
Astyanax, 24, 62, 75, 78, 91, 132, 134, 135, 137, 261, 305, 330, 338, 339, 342, 468, 469
138, 139, 140, 142, 148, 250, 252, 261, brevicaudatus, 251
263, 309, 344, 397, 424, 459, 467, 468 cephalus, 70, 338, 468
bimaculatus, 91, 132, 250, 261, 309, 397, erythropterus, 235, 339
424, 467 falcatus, 261, 468
fasciatus, 24, 30, 62, 75, 78, 91, 112, 134, hilarii, 305, 468
137, 140, 148, 261, 263, 467, 468 lundi, 138, 468
mexicanus, 459 melanopterus, 34, 37, 69, 78, 87, 468, 469
scabripinnis, 91, 468 pellegrini, 342
sp., 138, 252, 468 Bryconops alburnoides, 249, 468
spp., 344 Bucephalidae, 118, 123, 127, 133, 197
Atractidae, 234, 240, 245, 248 Bunocephalus, 30, 469
Atractosteus tristoechus, 217, 468 Caballerotrema, 126, 130, 136, 161, 466, 479
Auchenipterus nuchalis, 252, 468 Cacatuocotyle, 51, 56, 71, 98, 469
Bacciger, 125, 129, 137, 201, 480 Callodistomidae, 126, 130, 134
Bagrus pemecus, 459 Callophysus macropterus, 82, 260, 339, 388, 469
Basilichthys microlepidotus, 271, 468 Calpidothecioides, 54, 59, 71, 99, 487, 488
Bedsylernaea, 334, 336, 343, 378, 474 Calpidothecium, 47, 53, 58, 71, 99, 487, 488
Bellumcorpus, 123, 127, 133, 157, 465, 489 Calyptospora, 23, 25, 31, 35, 39, 40, 41, 469, 472
Belonesox belizanus, 91, 468 Camallanidae, 234, 239, 244, 248, 277
Betamphistoma, 124, 128, 146, 182, 478 Camallaninae, 239, 240, 244
Camallanoidea, 238, 239, 243, 244, 248 festivum, 258, 470

Camallanus, 240, 244, 248, 276, 278, 279 maculicauda, 86
Capillaria, 241, 246, 263, 474, 477 mayorum, 131, 470
Capillariidae, 241, 246, 263, 264 severum, 77, 151, 470
Capillostrongyloides, 241, 246, 263, 466, 467, 476, 490 sp., 131, 340, 470
Carapus fasciatus, 30, 469 tetracanthus, 397, 470
Carassius auratus, 469 pearsei, 86
Carnegiella strigata, 426, 469, synspilum, 86, 470
Catoprion mento, 66, 67, 78, 82, 83, 469 urophthalmus, 86, 470
Cephalosilurus zungaro, 63, 65, 485 Cichlidae, 43, 62, 77, 85, 88, 261, 424, 470
Ceratophrys cornuta, 216, Ciliophora, 20, 21, 23, 25, 26, 28, 31, 33, 460,
Cestoda, 5, 7, 20, 21, 206, 209, 211, 213, 222, 466, 465, 470, 471, 473, 480, 481, 494
468, 469, 470, 473, 479, 480, 482, Cladocystis, 126, 130, 144, 159, 489,
483, 484, 485, 486, 488, 493, 496 Cladorchiidae, 123, 145
Cestodaria, 20, 206, 209, 211, 213, 222, 466 Cleidodiscus, 47, 64, 70, 71, 83, 90, 474, 487, 492
Cetopsis caecutiens, 215, 469, Clinostomidae, 119
Chabaudinema, 240, 245, 256, 272, 471 Clinostomum, 119
Chaetobranchus, 120, 469 Coccidida, 23, 25, 26
semifasciatus, 120, 469 Collection, 05, 07, 22, 26, 45, 46, 47, 48, 121, 208,
Chalceus macrolepidotus, 78, 469 236, 301, 331, 391, 419, 455, 459
Chalcinotrema, 125, 129, 139, 167, 170, 467, 477, 495 Colocladorchis, 124, 128, 147, 176, 485
Chalcinus nematurus, 30, 469 Colomesus asellus, 471,
Characidae, 61, 62, 67, 68, 69, 70, 71, 75, 78, 80, Colossoma 31, 33, 44, 45, 69, 79, 80, 82, 133, 148,
85, 87, 89, 91, 253 152, 248, 252, 256, 261, 305, 306, 329,
Characidium, 71, 89, 469 334, 336, 344, 398, 399, 461, 471, 480
caucanum, 89, 469 bidens, 33, 69, 133, 148, 152, 305, 399,
lanei, 71, 469 471, 480
pterostictum, 71, 469 brachypomum 248, 256
Characinidae, 27 macropomum, 44, 45, 79, 80, 82, 252, 256,
Charax, 139, 469 261, 306, 329, 334, 336, 344, 398,
Chilodus punctatus, 307, 469 399, 461, 471
Choanoscolex, 210, 212, 214, 223, 493 metrei, 248
Chromis sp., 397, 469 Copepoda, 5, 7, 9, 20, 21, 326, 337, 346, 465, 466,
Cichla, 32, 77, 86, 217, 218, 236, 338, 339, 343, 468, 469, 470, 471, 472, 474, 475,
398, 423, 424, 426, 437, 469, 470 476, 478, 480, 482, 483, 485, 486,
monoculus, 339, 343, 426, 469 487, 488, 489, 490, 492, 494, 496
ocellaris, 32, 77, 89, 217, 218, 236, 338, 423, Corydoras, 62, 83, 91, 251, 344, 471
424, 469, 470 aeneus, 91, 471,
temensis, 398, 470 ehrhardti, 62, 344, 471
Cichlasoma, 31, 77, 86, 88, 131, 151, 250, 258, 306, paleatus, 62, 83, 251, 471
340, 397, 470 Cosmetocleithrum, 53, 57, 58, 72, 99, 479, 481, 486
bimaculatum, 77, 88, 470 Cosmocercoidea, 238, 240, 243, 245, 255
facetum, 250, 470 Cosmoxynema, 239, 244, 257, 267, 472
fenestratum, 31, 470 Cosmoxynemoides, 239, 244, 257, 267, 472, 473
Cotyloda, 213 Dendrorchis, 126, 130, 138, 163

Crassicutis, 126, 130, 131, 162, 466, 470, 472 Denticamallanus, 240, 245, 249, 290, 468
Creatochanes affinis, 75, 78, 471 Denticauda, 123, 127, 133, 179, 471
Crenicichla, 32, 131, 144, 307, 309, 399, 422, 424, 472 Diaphorocleidus, 54, 59, 74, 100, 467, 471, 473
geayi, 131, 307, 309, 399, 472 Dichelyne, 241, 246, 477, 478, 486, 490
johanna, 144, 472 Didymozoidae, 117, 118, 119, 123, 127, 135
lacustris, 422, 472 Diplectanidae, 43, 64
lepidota, 32, 472 Diplectanum, 47, 52, 57, 64, 95, 482
Crepidostomum, 126, 130, 131, 150, 476, 481, 489, Diplostomum, 120
Creptotrema, 126, 130, 132, 159, 476, 477, 495 Dipteropeltis, 390, 392, 394, 400, 406, 408, 412,
Cryptobia, 455 478, 482, 489, 490, 492
Cryptogonimidae, 124, 125, 128, 129, 135 Dolops, 390, 391, 392, 393, 394, 395, 399, 406,
Cucullanidae, 234, 241, 246, 252, 253 407, 408, 409, 410, 466, 467, 471,
Cucullanus, 241, 246, 252, 253, 271, 280, 285, 291, 472, 473, 474, 476, 477, 478, 479,
466, 468, 471, 472, 476, 477, 478, 480, 482, 485, 486, 487, 488, 489,
479, 480, 481, 482, 485, 486, 488, 490, 492, 494, 495
489, 490, 494 Doradamphistoma, 124, 149, 194, 202, 478
Curimata, 73, 85, 89, 91, 147, 153, 257, 258, 308, Doradidae, 72, 90, 239, 244, 330
340, 425, 472 Doras dorsalis, 152, 473
argentea, 73, 84, 91, 472 Dracunculoidea, 245, 253, 259
cyprinoides, 85, 340, 472 Dranculoidea, 238, 243
elegans, 153, 159, 309 Dujardinascaris, 235
gilberti, 91, 153, 257, 259 Echinocephalus, 235, 242, 246, 253, 269, 483
platana, 425, Echinorhynchidae, 302, 304, 305, 306
vittata, 147, 472 Echinorhynchus, 305, 318, 323, 468, 471, 479, 480, 495
Curimatrema, 124, 128, 147, 186, 442 Echinostomatidae, 126, 130, 136
Curumai, 123, 127, 132, 133, 179, 471 Eigenmannia virescens, 30, 91, 473
Curvianchoratus, 46, 52, 57, 73, 99, 472, 485 Electrophorus electricus, 459, 473
Cyclops, 326 Elipesurus, 220, 221, 473
Cymothoidae, 4, 416, 419, 420, 421, 422, 423, sp., 221
424, 425 spinicauda, 220, 473
Cynopotamus humeralis, 139, 423, 425, 472 Enallothecium, 53, 58, 75, 100, 484, 485, 487, 490,
Cystidicolidae, 261, 262 491, 492, 493
Cystidicoloides, 242, 247, 262, 266, 472, 489, 490, 494 Encysted parasites and foreign bodies, 460
Dactylogyridae, 43, 44, 64, 90, Endorchis, 210, 212, 214, 222, 223, 466, 468
Dadayius, 123, 127, 147, 168, 187, 192, 194, 478, 479 Eocreadium, 126, 130, 159, 475
Dadaytrema, 148, 174, 195, 471, 479, 480, 482, Ephedrocephalus, 210, 212, 214, 223, 480
486, 489 Ergasilidae, 326, 328, 329, 333, 335, 337, 338, 341
Dawestrema, 54, 59, 73, 100, 466 Ergasilinae, 333, 338, 339, 340, 341
Definition, 7, 19, 22, 23, 42, 84, 117, 206, 234, Ergasilus, 326, 327, 328, 330, 331, 334, 335, 339,
299, 326, 390, 416, 454, 458 346, 347, 348, 349, 350, 351, 352,
Deltamphistoma, 124, 128, 148, 182, 478 356, 357, 384, 385, 465, 466, 468,
Demidospermus, 43, 53, 54, 57, 58, 59, 73, 100, 475, 469, 470, 471, 474, 475, 476, 478,
476, 477, 479, 480, 481, 482 485, 488, 490,494, 496
Erythrinus erythrinus, 261, 473 Guyanemidae, 240, 245, 253, 254

Eucestoda, 206, 208, 213 Gymnorhamphichthys hypostomus, 183, 473
Euryhaliotrema, 47, 52, 57, 75, 100, 482 Gymnotus carapo, 91, 459, 473
Eustrongylides, 235, 236, 465, 486 Gyrodactylidae, 43, 44, 49, 50, 51, 55, 61
Eutetrarhynchidae, 209, 211, 221 Gyrodactylus, 47, 51, 56, 62, 94, 467, 469, 471, 473,
Eutetrarhynchus, 209, 211, 221, 222, 473, 483, 484 474, 476, 481, 483, 490, 495, 496
Excorallana, 426, 444, 466, 478 Habronematoidea, 238, 242, 243, 247, 261
Farlowella amazona, 63, 473 Halipegidae, 125, 129, 138
Fellodistomidae, 118, 125, 126, 129, 130, 136 Halipegus, 125, 129, 139, 163, 465, 467, 472, 481, 489
Fitzroyia lineate, 308, 473 Haploporidae, 118, 125, 129, 139, 197
Flabellifera, 426 Heliconema, 242, 247, 264, 292
Franciscodoras marmoratus, 27 Hemiancistris, 89, 473
Freitascapillaria, 241, 246, 263, 489 scaphirhynchae, 89, 473
Galeocharax, 135 Hemidoras carinatus, 426, 473
Gamidactylus, 333, 335, 342, 363, 366, 383, 490 Hemiodus, 80, 90, 141, 473, 474
Gamispatulus, 333, 335, 342, 365, 366, 369, 490 microlepis, 141, 473
Gamispinus, 333, 335, 342, 364, 366, 466 semitaeniatus, 80, 90
Gammamphistoma, 124, 128, 478 Hemisorubim, 90, 399, 459, 474
Genolopa, 125, 129, 143, 191 platyrhynchus, 459, 474
Geophagus, 62, 86, 250, 257, 258, 307, 398, 422, sp., 399, 474
425, 445, 473 Henneguya, 23, 24, 28, 29, 34, 37, 38, 467, 473,
brasiliensis, 62, 250, 257, 307, 422, 445, 473 474, 477, 485, 486, 488, 492
jurupari, 398 Herotilapia, 77
sp., 425, 473 Heteronchoinea, 50, 55, 60
steindachneri, 258, 473 Heterophyidae, 118, 119, 126, 130, 142
surinamensis, 86, 473 Heterothecium, 53, 58, 77, 101, 469, 491
Gephyrocharax valenciae, 309, 473 Heterotylus, 53, 57, 78, 101, 475
Gibsoniela, 210, 212, 214, 225, 446 Hexabothriidae, 43, 51, 55, 60
Glandulorhynchus, 123, 127, 133, 197, 475 Hirudinea, 5, 20, 21, 24, 455, 456, 461, 474, 480, 488
Glanidium, 64, 214, 473 Holobrycon pesu, 339, 474
melanopterum, 64, 473 Hoplerythrinus unitaeniatus, 254, 309, 399, 474
neivai, 473 Hoplias, 27, 29, 62, 90, 119, 248, 252, 254, 261,
sp., 214, 473 263, 264, 307, 309, 339, 340, 343,
Glossiphoniidae, 455, 456, 457 344, 397, 398, 399, 400, 424, 425,
Gnathostomatidae, 242, 246, 253 457, 459, 474
Gnathostomatoidea, 238, 242, 243, 246, 253 macrophthalmus, 424, 474
Goezia, 235, 239, 243, 254, 273, 274, 466, 470, 486 malabaricus, 27, 29, 62, 90, 19, 248, 252,
Gonocercella, 125, 129, 138, 475 254, 261, 263, 264, 307, 309, 339,
Gonocleithrum, 42, 54, 58, 59, 76, 101, 479 340, 343, 344, 397, 398, 399, 400,
Gorgoderidae, 126, 130, 137 424, 425, 457, 459, 474
Gorytocephalus, 303, 304, 306, 314, 317, 322, 472, 475 Hydrolycus, 133, 339, 398, 475
Gracilisentis, 303, 304, 307, 475, 478, 483 scomberoides, 339, 398, 475
Gussevia, 43, 52, 54, 57, 58, 59, 76, 101, 469, 470, 487 sp., 133, 475
Guyanema, 240, 245, 253, 272, 466, 474, 477, 488 Hyperopletes, 51, 55, 63, 94, 488
Hyphessobrycon callistus, 424, 475 Leporinus, 29, 30, 79, 85, 87, 132, 135, 139, 140,
Hypophthalmus edentatus, 249, 339, 341, 475 141, 142, 248, 250, 252, 263, 337,
Hypopomus sp., 150, 475 340, 400, 424, 425, 476, 477
Hypoptopoma thoracathum, 88, 475 copelandi, 248, 425, 476
Hypostomus, 27, 28, 61, 63, 78, 88, 89, 139, 140, 143, elongatus, 132, 140, 141, 476
150, 250, 255, 306, 307, 399, 424, 475 fasciatus, 87, 248, 337, 340, 400, 424, 476, 477
albopunctatus, 27, 250, 475 lacustris, 79, 477
aurogutatus, 27 mormyrops, 29, 30, 477
bolivianus, 89, 475 obtusidens, 142, 477
carinatus, 306, 475 octofasciatus, 425, 477
commersoni, 27, 399, 475 Lernaea, 334, 336, 343, 380
marginatus, 88, 475 Lernaeidae, 329, 334, 336, 342, 343, 344, 381
plecostomus, 139, 140, 143, 306, 475 Life-cycle, 22, 24, 117, 207, 234, 300, 330, 390,
punctatus, 27, 63, 150, 475 416, 455, 458
regani, 27, 61 Linguadactyla, 44
robinii, 88, 89, 475 Linguadactyloides, 44, 52, 56, 79, 108, 109, 471
sp., 27, 61, 78, 89, 255, 424, 475 Livoneca, 420, 421, 424, 428, 470, 477, 481
strigaticeps, 28 Longihaptor, 76
Ichthyophthirius, 25, 26, 28, 29, 35 Loricaria, 27, 29, 74, 132, 139, 142, 399, 477
Ichthyouris, 239, 244, 257, 258, 283, 470, 473, 478, 494 anus, 74, 142, 399, 477
Iheringichthys labrosus, 132, 153, 425, 475, 476 piracicabae, 27
Iheringtrema, 124, 128, 135, 164, 485 sp., 27, 29, 477
Inpamphistoma, 124, 128, 149, 188, 478 vetula, 132, 477
Isopoda, 5, 7, 9, 20, 21, 416, 427, 465, 466, 467, Loricariichthys, 27, 254, 255, 477
468, 469, 470, 472, 473, 474, 475, anus, 27
476, 477, 478, 479, 481, 482, 483, brunneus, 254, 477
485, 489, 490, 491, 492, 493, 494, 495 platymetopon, 255, 477
Jainus, 52, 54, 57, 59, 78, 102, 467, 468, 469, 471 Loricariidae, 61, 63, 74, 78, 88, 89, 239, 244,
Kalipharynx, 126, 130, 137, 161, 476 330, 477
Kalitrema, 124, 128, 150, 175, 475 Luciopimelodus, 39, 248, 400, 477, 478, 482
Kathlaniidae, 240, 245, 255, 256 pati, 248, 400, 482
Klossinemella, 240, 245, 248, 267, 282, 471, 474, Lycengraulis grossidens, 338, 339, 426, 478
476, 478, 479, 480, 486, 489 Magnivitellinum, 126, 130, 132, 467
Kritskyia, 51, 56, 79, 102, 477, 485 Manaosia, 209, 211, 215, 223, 482
Lahilliela kneri, 141 Megapriapus, 301, 302, 304, 315, 323, 483
Lamproglena, 334, 336 Megacoelium, 125, 129, 140, 165, 475, 487
Lebiasina bimaculata, 61, 62, 476 Megalancistrus aculeatus, 258
Lebistes reticulata, 90 Megalodoras irwini, 149, 478
Lecithobotrioides, 125, 129, 140, 169, 197, 485 Megapriapus, 301, 302, 309, 315, 323, 483
Leiarius marmoratus, 399, 476 Megathylacus, 209, 211, 215, 225, 478
Lepidosiren paradoxa, 29, 137, 476 Micramphistoma, 124, 128, 150, 194, 198, 475
Lepocreadiidae, 126, 130, 143 Microcotylidae, 51, 55, 60
Leporellus vitattus, 425 Micropogonias furnieri, 339
Leporinodus vittatus, 141, 476 Minilernaea, 334, 336, 343, 344, 379, 468, 471
Miracetyma, 333, 335, 340, 382, 472, 483, 485, Myxosporida, 20, 23, 25, 26, 28, 29
487, 488 Myxozoa, 5, 20, 29, 460, 465, 467, 469, 473,
Moenklausia, 139 474, 476, 477, 481, 482, 483, 485,
Monhysterides, 234, 235 486, 488, 492
Monocleithrium, 52, 56, 79, 103, 474 Myzobdella, 456, 457, 474, 488
Monocotylidae, 43, 50, 55, 61 Myzophorus, 210, 212, 215, 223, 480, 482, 486
Monogenoidea, 5, 7, 9, 11, 20, 21, 42, 43, 44, 45, Nannostomus beckfordi, 423, 479
46, 47, 48, 50, 55, 60, 92, 460, 465, Nematoda, 5, 7, 9, 20, 21, 238, 242, 248, 265, 465
466, 467, 468, 469, 470, 471, 472, 473, Neocucullanus, 241, 246, 252, 253, 270, 489, 490
474, 475, 476, 478, 479, 480, 481, 482, Neoechinorhynchidae, 303, 304, 306, 307, 333,
483, 484, 485, 486, 487, 488, 489, 490, 335, 340, 382, 472, 483, 485, 487, 488
491, 492, 493, 494, 495, 496 Neoechinorhynchus, 303, 304, 306, 314, 316, 317, 465,
Monorchiidae, 125, 129, 143 470, 471, 473, 474, 478, 485, 487
Monticellia, 210, 212, 215, 223, 225, 468, 469, 482, Neoparaseuratum, 241, 246, 260, 292, 486, 494
485, 486, 494 Nerocila, 4, 417, 420, 421, 424, 433, 447, 470, 472,
Morphology, 7 477, 485
Mugilicola, 329 Nesolecithus, 206, 209, 211, 213, 229, 230, 466
Mylesinus, 82, 142, 145, 146, 149, 152, 248, 478 Nilonema, 241, 245, 259, 270
paraschomburgkii, 82, 142, 145, 146, 149, Nominoscolex, 210, 212, 216, 222, 223, 468, 482,
152, 248, 478 483, 486
paucisquamatus, 82, 478 Nothogyrodactylus, 51, 55, 63, 94, 466
Myletes, 248, 478 Nothozothecium, 43
torquatus, 248, 478 Notothecioides, 53, 58, 81, 103, 478, 479
Myleus, 68, 69, 81, 82, 133, 146, 147, 148, 150, 151, Notothecium, 53, 57, 58, 75, 80, 103, 487, 491,
152, 248, 256, 259, 426, 478, 479 492, 493
(Myloplus) asterias, 146, 147, 150, 248, 478 Notozothecium, 54, 55, 59, 81, 103, 465, 471, 478,
(Myloplus) rubripinnis, 146, 147, 478 484, 485, 487, 491, 492, 493
pacu, 82, 151, 478 Nyctotherus, 25, 28, 29, 31, 35, 465, 470, 471, 480, 481
rhomboidalis, 82, 478 Octospiniferoides, 303, 304, 307, 315, 319, 469, 490
rubripinnis, 68, 69, 81 Odhneriotrema, 119
sp., 133, 148, 152, 256, 478, 479 Odontostilbe sp., 425
spp., 248 Odothecium, 54, 59, 82, 104, 469
ternetzi, 151, 256, 259, 479 Oligonchoinea, 50, 55, 60
torquatus, 81, 479 Onchobothriidae, 218
Myleusnema, 150, 240, 245, 246, 286, 287, 288, 479 Oncobothriidae, 209, 211
Myleustrema, 124, 125, 150, 183 Oogyrodactylus, 44, 51, 55, 63, 95, 473
Mylosoma, 305 Opisthorchiidae, 118, 126, 130, 144
paraguayensis, 305 Ornithoscolex, 209, 211, 216, 224, 483
Mymarothecium, 43, 53, 54, 55, 58, 59, 80, 103, Ossubtus xinguensis, 422
471, 480, 484, 487, 491, 492, 493 Osteoglossum biccirhosum, 76
Myxidium, 28, 30, 35, 465, 469 Ostospiniferoides, 320
Myxobolus, 23, 24, 25, 28, 31, 33, 37, 38, 39, 469, Otocinclus, 27, 423, 479
471, 473, 477, 481, 482, 483, 488, 492 francirochai, 27
Myxosoma, 24, 25 vestitus, 429, 479
Oxydoras, 72, 259, 262, 400, 479 Perulernaea, 329, 331, 334, 336, 344, 375, 376, 381,
kneri, 259, 262, 479 471, 480
niger, 72, 400, 479 Petenia splendida, 86, 480
Oxyuroidea, 238, 239, 242, 244, 248, 257 Phanerothecium, 51, 55, 63, 95, 469, 475, 483, 485
Pachyurus, 76, 262, 397, 479 Pharyngodonidae, 239, 244, 257, 258, 259
bonariensis, 76, 479 Phillostomella, 420, 421
junki, 262, 397, 479 Philocorydoras, 53, 58, 83, 104, 471
Pacu nigricans, 305, 479 Philometra, 241, 245, 260, 466, 469, 478, 481,
Pacudistoma, 124, 128, 151, 184, 194 482, 490
Paeonodes, 329 Philometridae, 234, 240, 245, 259, 260
Palaeocryptogonimus, 125, 129, 135 Philometroides, 240, 245, 489
Palliolisentis, 301, 302, 303, 308, 465, 495 Philostomella, 425, 431, 472
Pandosentis, 303, 304, 307, 315, 466, 472 Phractocephalus hemiliopterus, 90, 91, 214, 218,
Paracamallanus, 239, 244, 249, 277, 281, 475, 486 399, 480
Paracapillaria, 241, 246, 264, 265, 465, 472, 477, Phyllobothriidae, 209, 21, 219
478, 489, 490 Phyllodistomoides, 126, 130, 138, 468
Paracavisoma, 302, 304, 306, 318, 479, 485 Physalopteridae, 242, 247, 264
Paracymothoa, 420, 421, 424, 428, 431, 467, 474, 475 Physalopteroidea, 238, 242, 243, 247, 264
Paragyrodactyloides, 62, 471 Piabucina sp., 261, 480
Paragyrodactylus, 62 Piaractus brachypomus, 80, 248, 334, 336, 345,
Paraheteronchocotyle, 51, 55, 60, 93, 483 471, 480
Paralecithobotrys, 125, 129, 141, 173, 476 Pimelodella, 62, 64, 71, 251, 252, 257, 261, 262,
Paramphistomata, 123, 127, 145 263, 480, 481
Paramphistomidae, 118 lateristriga, 251, 252, 257, 261, 262, 263,
Paranaella, 51, 55, 60, 61, 93 480, 481
Paraproctotrema, 125, 129, 144, 190, 191, 468 sp., 64, 262, 481
Pararhipidocotyle, 123, 127, 134, 489 yuncensis, 62, 64, 481
Paraseuratum, 241, 246, 260, 271, 467, 473, 474, 494 Pimelodidae, 43, 62, 63, 64, 65, 70, 72, 74, 79,
Parasynodontisia, 239, 244, 258, 488 82, 84, 88, 90, 91, 239, 244, 305, 481
Parauchenipterus striatulus, 64, 480 Pimelodus, 30, 31, 65, 72, 74, 84, 88, 90, 91, 132,
Parspina, 126, 130, 142, 163, 481 135, 139, 143, 152, 215, 216, 217,
Pathology, 7, 19, 22, 25, 44, 119, 207, 235, 301, 218, 248, 251, 252, 255, 260, 399,
333, 391, 471, 455, 458 424, 425, 459, 477, 478, 481, 482
Paulicea luetkeni, 88, 217, 252, 253 albicans, 30, 72, 74, 399, 480, 481
Paurorhynchus, 123, 127, 134, 157 clarias maculatus, 74
Pavanelliella, 51, 56, 82, 104, 469, 486 clarias, 30, 31, 74, 84, 88, 132, 143, 248,
Pellona, 86, 137, 338, 399, 459, 480 251, 252, 260, 424, 425, 481
castelnaeana, 137, 338, 399, 459, 480 grosskopfi, 91
flavipinnis, 86, 480 maculatus, 255, 481, 482
Peltidocotyle, 209, 211, 216, 225, 483, 486 megacephalus, 459, 482
Pentastomida, 5, 20, 458, 459, 464, 465 ornatus, 152, 482
Percichthys, 308, 398, 457, 480 pati, 215, 216, 217
melanops, 308, 480 Pimelodus, 30, 31, 65, 72, 74, 84, 88, 90, 91, 132,
trucha, 398, 457, 480 135, 139, 143, 152, 215, 216, 217,
218, 248, 251, 252, 255, 260, 399, Potamotrygonocestus, 209, 211, 219, 227, 483, 484
424, 425, 459, 477, 478, 481, 482 Potamotrygonocotyle, 50, 55, 61, 93, 483
sp., 30, 482 Prehendorastrus, 328, 333, 335, 341, 369, 475
spp., 90 Prevention, 5, 7, 22, 26, 45, 120, 208, 236, 301,
vituga, 459 331, 391, 441, 455, 458
Pindapixara, 334, 336, 340, 382, 474 Pristis perottetti, 220
Pinirampus, 74, 215, 459, 482 Pristobrycon eigenmanni, 66, 67, 68, 69, 75, 80, 81,
pirinampu, 74, 459, 482 82, 484
sp., 215 Pristobrycon sp., 65, 66, 68, 69, 75, 80, 81, 484
piraeeba, 214, 216, 222, 468 Pristobrycon striolatus, 66, 67, 68, 75, 82, 83, 146,
Pirarara bicolor, 218, 480, 482 484, 485
Piscicolidae, 456, 457, 464 Procamallaninae, 240, 244
Pithanothecium, 53, 58, 83, 104, 469, 485, 487, 488 Procamallanus, 235, 240, 244, 245, 249, 250, 251,
Placobdella, 455, 457 252, 275, 276, 277, 289, 290, 465,
Plagioscion, 64, 76, 119, 136, 309, 345, 482 466, 467, 468, 469, 470, 471, 472,
sp., 64, 482 473, 474, 475, 476, 477, 478, 479,
squamosissimus, 64, 76, 19, 136, 309 480, 481, 482, 483, 484, 485, 486,
Platydoras costatus, 217, 473, 482 487, 488, 489, 490, 491, 492, 493,
Platyhelminthes, 20, 48 494, 495
Platystoma sp., 215, 494 Procaudotestis, 126, 130, 132, 160, 477
Platystomatichthys sturio, 215, 216, 218, 483 Prochilodus, 27, 62, 66, 79, 84, 85, 87, 140, 142,
Plecostomus, 63, 89, 139, 140, 143, 306, 307, 475, 483 147, 153, 262, 307, 340, 485
plecostomus, 63, 483 lineatus, 79, 84, 85, 142, 485
sp., 89, 483 nigricans, 84, 140, 340, 479, 485
Poecilia, 62, 483 platensis, 153, 485
caucana, 62, 483 reticulatus, 66, 87, 140, 142, 262, 307, 485
reticulata, 62, 483 scrofa, 262, 485
sphenops, 62, 483 sp., 27
Poecilostomatoida, 337, 341, 345 Pronamphistoma, 124, 128, 151, 195, 470
Polyacanthorhynchus, 302, 304, 309, 322, 466 Prosorhynchus, 123, 127, 134, 157, 196, 465, 467,
Polyonchoinea, 50, 55, 61 489, 492
Pomoxis annularis, 338 Prosthenhystera, 126, 130, 134, 158, 465, 467, 476,
Pomphorhynchidae, 302, 304, 308 477, 485, 489, 496
Pomphorhynchus, 302, 304, 308, 318, 480 Proteocephalidae, 209, 211, 213, 469
Porrocaecum, 235, 466 Proteocephalidea, 208, 213
Potamotrygon, 60, 61, 219, 220, 221, 253, 255, 306, Proteocephalus, 210, 212, 217, 226, 227, 228, 468,
459, 483, 484 469, 470, 479, 482, 486, 488
circularis, 60, 61, 219, 220, 253, 255, 483 Protorhinoxenus, 52, 57, 83, 104
falkneri, 220, 221, 483 Protozoa, 5, 20, 23, 33, 455, 456
hystrix, 219, 220, 221, 253, 255, 306, 483 Psectrogaster, 70, 340, 472, 485
magdalenae, 219, 220, 483 essequibensis, 346, 485
motoro, 219, 220, 221, 459, 484 rutiloides, 70, 485
reticulatus, 220, 221, 484 Pseudocladorchis, 124, 128, 151, 168, 185, 471, 473,
yepsi, 220 478, 479, 482
Pseudodiplodiscus, 123, 127, 152, 168, 473 Rhadinorhynchidae, 302, 303, 309, 310
Pseudodoras niger, 252, 306, 479, 485 Rhadinorhynchus, 302, 304, 309, 311, 312, 313, 482
Pseudogorgorhynchus, 302, 304, 310, 321, 466 Rhamdia, 27, 29, 64, 65, 71, 79, 82, 131, 215, 340,
Pseudoparabaris, 123, 127, 133, 179, 471 397, 398, 399, 457, 488
Pseudopimelodus, 28, 29, 135, 213, 248, 252, 253, guatemalensis, 82, 398, 488
469, 485 quelen, 27, 64, 65, 71, 79, 131, 340, 399, 488
roosevelti, 135, 485 sapo, 27, 71, 457, 488
zungaro, 28, 29, 213, 243, 252, 253, 469 sebae, 29, 65, 71, 488
Pseudoplatystoma, 29, 82, 90, 145, 215, 216, 217, sp., 71, 215, 397, 488
218, 252, 391, 398, 399, 459, 486 Rhamphichthys rostratus, 253, 488
coruscans, 82, 90, 398, 486 Rhaphiodon, 340, 398, 399, 459, 488
fasciatum, 29, 90, 215, 216, 218, 398, 399, 486 vulpinus, 340, 398, 399, 459, 488
sp., 90, 217, 252 Rhinebothrium, 209, 21, 220, 483, 484
tigrinum, 215, 216, 217, 459, 486 Rhinebothroides, 209, 21, 220, 222, 473, 483, 484
Pseudotylosurus angusticeps, 338, 486 Rhinelepis aspera, 61, 258, 488
Pseudovancleaveus, 54, 58, 84, 104, 475, 476, 481 Rhineloricaria sp, 63, 488
Pterodoras granulosus, 31, 72, 90, 152, 248, 255, 260, Rhinergasilus, 334, 336, 341, 356, 362, 487, 492
307, 400, 486, 487 Rhinodoras dorbignyi, 135
Pterophyllum scalare, 77, 86, 487 Rhinonastes, 52, 56, 84, 105, 485
Pterygoplichthys, 140, 141, 258, 487 Rhinoxenus, 52, 56, 85, 105, 469, 472, 476, 485,
aculeatus, 258, 487 487, 489, 490, 491, 492, 495
pardalis, 140, 487 Rhynchodinium, 28, 29, 31, 36
multiradiatus, 487 Rhytiodus, 85, 90, 399, 489
sp., 141, 487 argenteofuscus, 85, 489
Pygidium, 250, 251, 487 microlepis, 90, 399, 489
brasiliensis, 251 Riggia, 418, 420, 421, 425, 430, 433, 439, 440, 448,
punctatum, 250 449, 466, 472, 476, 477, 479, 490
Pygocentrus, 43, 65, 66, 67, 68, 69, 71, 72, 75, Roeboides, 67, 132, 139, 489
80, 81, 82, 83, 85, 252, 340, 487, bonariensis, 132, 489
488, 491, 492 myersi, 67, 489
nattereri, 43, 65, 66, 67, 68, 69, 71, 72, 75, Rondonia, 234, 235, 240, 245, 248, 267, 275, 471,
80, 81, 82, 83, 85, 340, 487, 491, 492 472, 478, 479, 480, 481, 485, 487, 494
sp., 252, 487 Rondotrema, 125, 129, 141, 197, 473
Pygopristis denticulata, 71, 72, 83, 488 Rudolphiella, 217, 227, 482
Quadrigyridae, 302, 303, 308 Rumai, 241, 245
Quadrigyrus, 302, 303, 308, 314, 467, 468, 472, Saccocoelioides, 125, 129, 141, 171, 172, 178, 182,
473, 474, 494 467, 476, 477, 478, 485, 489, 490
Quimperiidae, 241, 246, 260, 261 Saccodon caucae, 142, 489
Raphidascaris, 239, 243, 255, 273, 470, 475, 477, 482 Salminus, 67, 85, 133, 134, 135, 139, 145, 250,
Raphidascaroides, 239, 243, 255, 285, 486 252, 260, 262, 263, 344, 398, 399,
Raphiodon vulpinus, 254 400, 424, 459, 489, 490
Rhabdochona, 242, 247, 261, 266, 467, 468, 473, affinis, 67, 489
476, 480 brevidens, 344, 386, 398, 400, 459, 489
Rhabdochonidae, 242, 247, 261 hilarii, 133, 134, 250, 260, 262, 424, 489
maxillosus, 85, 134, 139, 145, 252, 262, 263, Spirocamallanus, 235, 240, 244, 250, 275, 276,
398, 399, 400, 489, 490 277, 285
sp., 400 Sprentascaris, 239, 243, 255, 273
Sanguinicola, 123, 127, 153, 169, 476, 485 Stenarchorhamphus muelleri, 443
Sanguinicolidae, 118, 119, 123, 127, 152 Stenarchus brasiliensis, 425
Schizochoerus, 206, 209, 21, 213, 229, 466 Sternopygus macrurus, 262, 494
Schizodon, 29, 85, 142, 248, 307, 342, 399, 400, Strigeoidea, 120, 180
425, 476, 490 Strongylura, 338, 339, 494
fasciatus, 29, 142, 490 study methods, 22, 26, 45, 121, 208, 236, 301,
nasutus, 248, 425 331, 391, 419, 455, 459
Sciadicleithrum, 43, 52, 57, 85, 105, 465, 470, 473, Symbranchus marmoratus, 309, 459
480, 487, 494, 495, Symphysodon discus, 258, 494
Sciadocephalus, 210, 212, 218, 226, 470 Tachysurus hertzbergii, 459
Scleroductus, 51, 56, 63, 95, 473, 480, 481, 488 Taurocheros, 344, 373, 489
Semaprochilodus, 62, 119, 337, 339, 342, 490 Telethecium, 51, 56, 86, 106, 479, 480
insignis, 19, 337, 339, 342, 490 Telotha, 420, 421, 425, 429, 431, 450, 468, 473,
taeniurus, 62, 490 474, 476, 481, 494
Serrasalmus, 29, 30, 31, 65, 66, 67, 68, 69, 75, 78, 79, Tereancistrium, 54, 58, 59, 86, 106
80, 81, 82, 85, 134, 341, 398, 399, 400, Terranova, 235, 239, 243, 255, 270, 292, 483
422, 423, 426, 459, 490, 491, 492, 493 Tetragonopterus argenteus, 261, 494
compressus, 65, 66, 75, 81, 490 Tetraphyllidae, 218
elongatus, 65, 66, 68, 69, 75, 81, 82, 491 Tetraphyllidea, 208, 209, 21
gouldingi, 65, 66, 75, 78, 80, 81, 82, 491 Thelazioidea, 238, 242, 243, 247, 261
manuelli, 66, 82, 491 Therodamas, 329, 330, 331, 345, 370, 371, 372,
marginatus, 79, 85, 491 381, 482
nattereri, 341, 398, 399, 400 Therodamasidae, 329, 330, 345, 381
piraya, 30, 400, 459 Theromyzon, 456, 457, 480
rhombeus, 30, 65, 66, 67, 68, 69, 75, 80, 81, Touzeta, 241, 246, 261, 274, 470
82, 134, 492 Trachydoras paraguayensis, 248, 258, 260, 494
sp., 31, 65, 66, 67, 68, 69, 75, 80, 81, 82, Transmission, 7, 19, 21, 22, 24, 25, 26, 44, 117,
423, 493 118, 207, 234, 236, 300, 330, 390,
spilopleura, 66, 67, 68, 69, 75, 79, 80, 81, 416, 455, 458
422, 426, 492, 493 Travassosinia, 124, 128, 152, 168, 471
Seuratoidea, 238, 241, 243, 246, 252, 260 Travassosnema, 240, 245, 254, 292, 465
Silurus sp., 214, 215, 493, 494 Travnema, 239, 244, 259, 267, 467, 472, 485
Sorubim lima, 91, 482, 494 Treatment, 5, 7, 21, 22, 26, 45, 120, 121, 208,
Spasskyellina, 210, 212, 218, 483, 486 214, 236, 301, 331, 391, 419, 455,
Spectatus, 240, 245, 256, 267, 471, 480 458, 459, 460
Sphaeroides testudineus, 31, 494 Trematoda, 5, 9, 20, 21, 131, 154, 465, 466, 467,
Sphericomonorchis, 125, 129, 144, 190, 472 468, 469, 470, 471, 472, 473, 474,
Sphericomonorchis spinulosus, 125, 129, 144, 190 475, 476, 477, 478, 479, 480, 481,
Spinitectus, 235, 242, 247, 262, 268, 280, 479, 480, 482, 485, 486, 487, 488, 489, 490,
481, 472, 482, 485, 489, 494, 495 493, 493, 495, 496
Spinoxyuris, 239, 244, 259, 284, 479 Trichinelloidea, 238, 241, 242, 246, 263,
Trichodina, 28, 29, 31, 36, 494 Unilatus, 53, 57, 89, 107, 473, 475, 483, 487
Trinibaculum, 52, 57, 87, 106 Urocleidoides, 42, 47, 54, 58, 64, 70, 74, 89, 91, 467,
Trinidactylus, 52, 56, 87, 106, 470 468, 469, 471, 472, 473, 474, 475,
Trinigyrus, 52, 56, 88, 106, 465, 475 480, 481, 483, 485, 489, 494, 495
Triportheus 24, 67, 68, 69, 70, 85, 132, 250, 301, Urocleidus, 47, 71, 83
305, 308, 422, 494, 495 Vaigamidae, 329, 33, 335, 341, 342
albus, 67, 69, 70, 422, 494 Vaigamus, 328, 33, 335, 341, 361, 496
angulatus, 67, 68, 69, 70, 494, 495 Vanamea, 418, 420, 421, 425, 428, 433, 468, 469,
elongatus, 24, 67, 68, 69, 70, 321, 495 470, 473, 478, 491, 492, 493, 495
paranensis, 132, 305, 308, 495 Vancleaveus, 43, 54, 59, 64, 84, 90, 107, 480, 481, 486
sp., 67, 70, 85, 250, 495 Vandelia cirrhosa, 426, 495
Tropical Gill-Rot, 460 Witenbergia, 126, 130, 145, 158, 486
Trypanorhyncha, 208, 209, 21, 221 Wolffhugelia, 303, 304, 307, 314, 473
Trypanosoma, 23, 27, 36, 455, 465 Woodlandiella, 210, 212, 218, 225, 493
Tubifex, 24, 26 Xiphophorus helleri, 90, 495
Uaru amphiacanthoides, 77, 86, 495 Zetamphistoma, 124, 128, 152, 185, 194, 478,
Unibarra, 52, 56, 88, 107, 480, 481 Zonocotyle, 123, 127, 153, 177, 472
Unicoelium, 125, 129, 142, 485 Zonocotylidae, 118, 123, 127, 153
unidentified catfish, 251, 423 Zonocotyloides, 123, 127, 153, 177, 472
unidentified fish, 30, 251 Zygobothrium, 210, 212, 218, 224, 480, 482

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ABLA Vol.

1 – Thatcher: Amazon Fish Parasites 1

Aquatic Biodiversity in Latin America

Biodiversidad Acuática en América Latina

BIODIVERSIDAD ACUÁTICA EN AMÉRICA LATINA

Lector Científico a los Editores:

AQUATIC BIODIVERSITY IN LATIN AMERICA

Scientific Reader to the Editors:

AQUATIC BIODIVERSITY IN LATIN AMERICA

Series Editors/Editores de la Serie:

Aquatic Biodiversity in Latin America Vol. 1

First published 2006

Printed in Bulgaria, March 2006

AQUATIC BIODIVERSITY IN LATIN AMERICA

BIODIVERSIDAD ACUÁTICA EN AMÉRICA LATINA

Series Editors: Joachim Adis, Jorge R. Arias,

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

DR. JORGE R. ARIAS

M.Sc. GUILLERMO RUEDA-DELGADO

Amazon Fish Parasites by Vernon E. Thatcher

a variety of studies including taxonomic, phylogenetic and ecologic investigations of the

DELANE C. KRITSKY, Ph.D.

PREFACE TO THE SECOND EDITION

PREFACE TO THE FIRST EDITION

3 MI. Definition and morphology

Key to genera of freshwater Amazonian Gyrodactylidae 51

4 T I. Definition (Dand morphology

5 C I. Definition and morphology

Key to Cestodaria and Cestoda of Neotropical

6 NI. Definition and morphology

Key to the Habronematoidea of Amazonian freshwater fishes 242

7 A I. Definition and morphology

V. Collection and study methods 301

8 C I. Definition and morphology

9 B I. Definition and morphology

III. Pathology 391

10 II. Definition and morphology

IV. Prevention and treatment 455

12 A DDENDUM: HOST–PARASITE TABLE 465

13 S UBJECT INDEX 497

concerned with parasites, at least in Northern temperate climes. The ichthyopathologist,

Cited and general references

I. Definition and morphology

II. Life-cycle and transmission

IV. Prevention and treatment

V. Collection and study methods

VI. Identification and keys

TRYPANOSOMA HOST FISH

strigaticeps Hypostomus strigaticeps

Key to the genera of Neotropical Myxosporida

I. Polar capsules located in one extremity of the spore only.

Clave para los géneros de Myxosporida Neotropicales

I. Cápsulas polares localizadas en solo una de las extremidades de la espora.

Key to Neotropical fish ciliophorans

Clave para los cilioforos de peces Neotropicales

I. Macronúcleo en forma herradura.

VII. Checklist of Myxosporida from Neotropical freshwater fishes

Measurements are given in micrometers (µm).

Myxobolus sp. WALLIKER, 1969: kidney of Serrasalmus sp.: Brazil. Spore = 9 x 11

Checklist of Ciliophora from Neotropical freshwater fishes

Measurements are given in micrometers (µm).

posterior end. Sporozoite developing in invertebrate host. Parasites of

VIII. Plates of Protozoa and Ciliophora (Figs. 2–1 to 2–44)