parasites of offshore big game fishes of puerto rico - Uprm

PARASITES OF OFFSHORE BIG GAME 

FISHES OF PUERTO RICO AND 

THE WESTERN ATLANTIC

PARASITES OF OFFSHORE BIG 

GAME FISHES OF PUERTO RICO 

AND THE WESTERN ATLANTIC 

Ernest H. Williams, Jr. 

and 

Lucy Bunkley-Williams 

Sportfish Disease Project 

Department of Marine Sciences 

and 

Department of Biology 

University of Puerto Rico 

P.O. Box 5000 

Mayagüez, PR 00681-5000 

1996

2 

Cover drawing: Atlantic blue marlin with magnifications of parasites found 

on this host in Puerto Rico. Cover by Ms. Gladys Otero 

Published cooperatively by the Puerto Rico Department of Natural and 

Environmental Resources , P.O. Box 5887 , Puerta de Tierra, San Juan, PR 

00906; and the University of Puerto Rico, Mayaguez, PR 00681 

Funding provided by the Department of Natural and Environmental Resources 

with Sportfish Restoration Funds, Federal Aid Projects F-28 and F-35 (U.S. 

Fish and Wildlife Service) and the University of Puerto Rico, Mayaguez. 

The content of this book is the sole responsibility of the authors and does not 

necessarily reflect policy of the Department of Natural and Environmental 

Resources or the U.S. Fish and Wildlife Service. 

Printed by Antillean College Press, Mayaguez, PR 00681 

First printing 1.5 M, 1996 

Library of Congress Catalog Card Number 96-86468 

ISBN 0-9633418-2-0 

Any part of this book may be copied for individual use. Whole copies may be 

requested from: The Department of Natural and Environmental Resources or 

Ernest H. Williams, Department of Marine Sciences, University of Puerto Rico, 

P.O. Box 908, Lajas, PR 00667 

Suggested citation: 

Williams, E. H., Jr. and L. Bunkley-Williams 1996. Parasites of offshore big 

game fishes of Puerto Rico and the western Atlantic. Puerto Rico 

Department of Natural and Environmental Resources, San Juan, PR, 

and the University of Puerto Rico, Mayaguez, PR, 382 p., 320 

drawings. 

Key Words: Fish parasites, big game fishes, parasite ecology, parasite 

evolution, fish diseases.

CONTENTS 

REVIEWERS ............................................................................................. 4 

INTRODUCTION ..................................................................................... 5 

Definitions and conventions................................................................. 7 

PROTOZOA (PROTOZOANS)................................................................. 11 

MYXOZOA (MYXOZOANS) ................................................................... 11 

FUNGI (FUNGUS) .................................................................................... 19 

PLATYHELMINTHES (FLATWORMS) ................................................ 22 

Udonellidea (copepod worm) ............................................................... 22 

Digenea (flukes)................................................................................... 24 

Didymozoidea (tissue flukes) ......... .................................................... 64 

Monogenea (gillworms) ................................. .................................... 79 

Cestoda (tapeworms).......................................................................... 103 

NEMATODA (ROUNDWORMS)................................ ........................... 133 

ACANTHOCEPHALA (SPINY-HEADED WORMS)............................. 156 

CRUSTACEA (CRUSTACEANS) ..................... ...................................... 163 

Ostracoda (seed shrimp) .............................. ........................................ 163 

Copepoda (copepods) .................................. ....................................... 165 

Branchiura (fish lice) ...................................... .................................... 223 

Cirripedia (barnacles) ............................... .......................................... 224 

Isopoda (isopods) ................................................................................. 228 

PISCES (FISHES) ..................................................................................... 239 

Petromyzontiformes (lampreys) ......................................... 241 

Squaliformes (cookiecutter sharks) ....................................................... 242 

Perciformes (remoras and pilotfish) .............. ....................................... 247 

OTHER DISEASES AND CONDITIONS .............. .................................. 260 

DISCUSSION .............................................. .............................................. 267 

Harm caused by parasites ........................ ............................................. 267 

Transmission of parasites .............................. ...................................... 268 

Usefulness of parasites .................................... .................................... 270 

HOST SUMMARIES AND HOST-DISEASE CHECKLISTS ................. 275 

ACKNOWLEDGMENTS 

Assistance ................................................ ............................................ 341 

Illustration sources .................................. ............................................. 342 

BIBLIOGRAPHY ............................................. ......................................... 344 

INDEX .................................................................... .................................. 363 

ABOUT THE AUTHORS ...................................... .................................. 381 

3

INTRODUCTION 

The purpose of this book is to serve as a fish parasite guide for sport 

fishermen, commercial and tour-guide fishermen, fishery biologists, ecologists, 

scientists, and anyone interested in the health and welfare of big game fishes. 

We hope it will encourage the study of the interrelationships between fishes and 

their intimate parasite partners around Puerto Rico and throughout the Atlantic. 

Big game fishes are important sportfishing and food resources. Many of 

their fish parasites attract the attention of fishermen because they are large, 

abundant and always on particular host species. The health of these fishes, and 

the humans who enjoy catching and eating them, is of great concern. The 

environments of many coastal areas have deteriorated and there is little 

agreement about how much of this environmental damage has spilled over into 

the open ocean. The abundance and diversity of big game fish parasites might 

be used as an indicator of environmental changes. Many parasites are useful as 

biological tags for tracing stock movements, mixing, migrations and other 

aspects of the biology of big game fishes. They can also provide readily 

available examples of many invertebrate phyla which can be used for classroom 

examinations. 

The present knowledge of parasites of big game fishes is at the most basic 

level, describing species, but the interrelationships between big game fish 

parasites and their intermediate and final hosts are as complex and intricate as 

food webs described by ecologists. It is extremely difficult to study the parasites 

and diseases of live oceanic, fast-moving fishes. Hook and line fishing selects 

for healthy fishes because sick fishes seldom bite lures or baits. Debilitated 

fishes in the open sea are quickly eaten, or sink, and seldom wash ashore. Still, 

we can assume that parasites and diseases cause as many problems for these big 

game fishes as they do for fishes in more easily examined habitats. A study 

found that the presence of one species of parasite reduced the yield of a big 

game fishery by about one fifth. We estimate that somewhere between one 

third to one half of all big game fish resources are lost due to disease. We need 

to understand the workings of diseases throughout the ecosystem if we are to 

have any hope of recuperating losses due to diseases. Unfortunately, we lack 

complete knowledge of these processes in a single fish, or even of a single 

disease organism! Most big game fishes are either already overexploited, or are 

soon to be over fished. As dolphin, greater amberjack and other big game 

fishes are raised in captivity, we are beginning to discover their deadly 

parasitic, bacterial and viral diseases. We cannot afford to ignore manageable 

problems that have the potential to double the stocks of big game fishes We 

hope that this book will serve as a beginning to better understand these forces 

in the ecology of big game fishes. 

No one before has had the opportunity to make an overview of an entire 

ecological mix of parasites of the dominant predators of an ocean system. We 

discuss some trends and relationships, but too little information exists for a 

5

6 

PARASITES OF OFFSHORE BIG GAME FISHES OF PUERTO RICO 

complete analysis of this system. Even though many big game fishing 

tournaments are held every year, there is an incredible lack of published 

information and basic knowledge about parasites of big game fishes. Much 

confusion has been caused by the difficulties in identifying these parasites. We 

hope this book will solve that problem and will allow fisherman and amateur 

scientists to make contributions to our knowledge of big game fishes. It is a 

world of great to little mysteries--all worthy of exploration. 

The information included in this book comes from our long-term, original 

observations on big game fishes in Puerto Rico, the West Indies, Gulf of 

Mexico, Atlantic coast of North America, and from published records in a 

scattered, often obscure and contradictory literature. Fishes examined in this 

work were collected by hook and line in fishing tournaments in Puerto Rico 

from 1974 to 1996 and in Dauphin Island, Alabama, from 1967 to 1974; or by 

individual sport fishermen and scientists (see Acknowledgments). Some of the 

barracuda, mackerels and jacks were speared by the authors using skin and 

scuba diving equipment; or collected with nets by Department of Natural and 

Environmental Resources (DNER) personnel. There was no attempt to make 

systematic collections. Parasites were either removed from freshly caught hosts 

and preserved immediately, or organs and tissue samples removed and placed 

in plastic bags surrounded by ice for laboratory examinations within 24 hours. 

We recognize 39 species of offshore big game fishes in the western Atlantic. 

Three (Atlantic bonito, chub mackerel, frigate tuna) have not been reported off 

Puerto Rico and 3 others only occur further south (serra Spanish mackerel) or 

north (Atlantic mackerel, Spanish mackerel) than our island. They were 

included to provide a more complete analysis of these otherwise rather 

homogenous, and wide-ranging, groups of fishes and parasites. 

We identify and define 273 species on/in these fishes from the western 

Atlantic. Each parasite species is illustrated and descriptions of its diagnostic 

characters, records, geographic range, location in host and length are always 

presented; and, usually to occasionally, name, life history, ecology, 

associations, host specificity, damage to host, detection, harm to humans, 

preparation for study, treatment and significance to sport fishing are discussed. 

The damage parasites cause in big game fishes has rarely been documented 

histologically (in tissue sections), therefore we have estimated potential damage 

based on number and size of parasites, known pathological changes caused by 

similar infections, and our own experience. No genius is required to predict 

that superinfections will harm a host, but our other interpretations may be less 

certain. Most families and genera of parasites are not discussed as groups 

except for a few that have a series of similar species. In these cases it was more 

efficient to discuss their similar characters rather than repeating this 

information with every species. 

Methods are suggested to avoid the spread of diseases. Diseases of big 

game fishes (other than parasites) are noted. The importance of parasites, their 

use as environmental indicators, controlling and avoiding fish parasites and the 

effects of fish parasites on humans are discussed. Damaging and dangerous 

parasites are described, and methods to protect human health are explained. The

INTRODUCTION 

Host Summaries and Host-Disease Checklists include the complete classification of 

these fishes and a list of the diseases we found on each host from the western 

Atlantic and parasites on these fishes worldwide. 

We hope you will use this book as your guide to the parasites of big game 

fishes. Within the drama of big game fishing lies a play-within-a-play of the 

parasites and associates of these great fishes. We look forward to receiving 

reader input concerning these fascinating creatures. - - - Good fishing! 

DEFINITIONS AND CONVENTIONS 

We tried to avoid the use of scientific jargon, but some scientific terms must 

be used for the sake of conciseness, clarity or continuity. These terms are either 

defined where they appear, in each 

section explanation or below. 

Levels of infections for microscopic 

parasites (protista) were estimated from 

skin or fin scrapings or gill clippings 

viewed with a compound microscope. 

Five medium power (100X) fields were 

averaged to establish parasite levels 

shown in the box at right. The very 

light category can be less than 1 because 

the value was taken from 5 fields. 

These categories were also used to 

express the number of metazoan 

parasites per host (or average of 5 hosts when possible). 

Frequency of Infection 

always = 100-99% 

almost always = 98-90% 

usually = 89-70% 

frequently = 69-50% 

commonly = 49-30% 

often = 29-10% 

occasionally = 9-1% 

rarely =

8 


dependable indicators of the evolutionary relationships (phylogeny) and population 

parameters (biological tags) of the host than more casual parasites. 

Characteristic and primary parasites are usually specific to a species, genus or 

family of host(s). Approximately 100 specimens of a host (from different 

times and localities)must be examined for parasites to confirm its characteristic 

or primary parasites. Few hosts and parasites in this book are so well known. 

In many cases, we have speculated and used these terms with insufficient data. 

Further studies will be necessary to confirm or refute these projections. Less 

important categories are "secondary" parasites occurring occasionally to 

commonly in the host; and "accidental" parasites occurring rarely to almost 

never. Any of these terms can be used to describe multiple parasite species on 

a host (for example: characteristic parasite fauna). 

The specificity of parasites has been variously defined. We prefer: 

Host specific - The parasite is confined to one species of host, or occurs no 

more than accidentally in other hosts or in false or transfer hosts. 

Almost host specific - The parasite is confined to one species of host, but 

rarely to very rarely occurs in others. 

Dominant host - The parasite is characteristic of one host, and occurs no more 

than commonly in others. 

Preferred host - The parasite occurs most often and/or in higher numbers in 

one host species. 

Genus specific, Family specific, etc. - These terms can be used with similar 

definitions and gradations as above. 

Habitat specific - The parasite usually to always occurs in hosts associated 

with a particular ecological niche or set of ecological conditions, and 

demonstrates no other host specificity. 

No specificity - The parasite occurs in all available hosts with no discernable 

pattern of preference or abundance. 

A number of different kinds of hosts have been defined and we coin a few 

new ones which seem to be important in the realm of big game fishes: 

Accidental host - Sometimes called Incidental host. The parasite very rarely 

to almost never occurs in very light infections. Usually these parasites do not 

mature on or in the accidental host. 

False host - Sometimes called Temporary host. The parasite occasionally to 

almost never occurs in very light to moderate infections in the stomach or 

intestine of the host. The host is a predator which has eaten a prey fish infected 

with this parasite. The parasite remains after the prey fish has been digested. 

It probably does not survive long in this inappropriate host. Another indication 

of prey transfer is parasites, that only occur in the intestine of prey fishes, but 

are only found in the stomach of predators. 

Intermediate host - A different larval stage of a parasite infects each 

intermediate host. One or more intermediate hosts may be necessary to 

complete a life cycle. 

Intermediary host - Many parasites and associates rest and feed on/in a host 

that is not appropriate as their final host. This mechanism allows them to sur-

DEFINITIONS AND CONVENTIONS 

vive longer to locate a final host. It is particularly important in oceanic habitats. 

This host is not an intermediate host because the parasites are not larval stages; 

and it is not a transport host because the parasites are not encysted. 

Decoy host - An intermediary host on/in which parasites lie in wait to either 

ambush attach or penetrate the gut of predators that prey on their host. This 

mechanism appears to be well established in big game fishes. Possibly an 

abundance of intermediary parasites (or the reputation for having them) could 

protect prey organism from predation. 

Transport host - Sometimes called Reservoir host or Paratenic host. An 

encysted parasite in an intermediate host is eaten by a predator that is not the 

correct final host. The parasite re-encysts in the inappropriate host and will 

develop into an adult only when this host is eaten by the proper final host. The 

ability to transfer among succeeding hosts increases the parasite's chances of 

reaching the correct host. 

Describing the size of a parasite 

is useful. Too often terms such as 

"large" and "small" have been used 

without explanation. We define a 

scale of relative size from the 

perspective of fishermen or casual 

observers. If the unaided eye cannot 

see it, then it is "microscopic"; if it 

can be barely seen it is "minuscule"; 

a fingernail length is "moderate". 

We also define size in metric scale 

as shown in the box at right 

We provide a classification for 

each group of parasites. Classifica- 

Size of a Parasite 

microscopic = 40.0 mm 

tion groups organisms at different levels and is a method of sorting similar 

things close together and different things further apart. It allows us to guess 

what an unknown organism must be, just by knowing another organism in its 

group or by the general characters of the group. Most parasite classifications 

have been stable for decades to centuries, but advances in modern biology have 

brought about many exciting changes in the ways we classify parasites. We 

have attempted to choose the most recent and stable of these systems. 

Some species of big game fish parasites have been named more than once 

because the characters used to separate species are sometimes difficult to interpret. 

As parasites become better studied, two or more similar forms that had 

been described as different species may be found to overlap in characteristics. 

Thus one species may be found to have more than one species name. Since a 

species can only have one name, the first name used (first description in the 

literature) is valid and the names published later are called synonyms. We only 

explain synonyms which have either been used until recently, or are important 

in understanding the biology or history of a parasite. Other synonyms can be 

found in the technical guides that we list. 

9

10 


Proper describing and naming of an organism allows us to correctly 

identify it time after time and to build up biological information about it. 

Otherwise we would be overcome with uncertainty, identifying organisms over 

and over, and starting from scratch each time. Indifferent or incomplete 

classification or taxonomy of big game fish parasites has drastically repressed 

knowledge of these important organisms. Certainty in absolutely identifying a 

parasite is the difference between knowing if a worm you noticed in your last 

piece of sushi at your boss's party can kill you; and in saving $100,000 worth 

of thawed filets because you can quickly prove the contaminating worms are 

harmless. 

American Fisheries Society (AFS) approved common names (Robins et 

al. 1991) of western Atlantic big game fishes are used in the text with their 

scientific names listed in the Host Summaries. Fishes not occurring in the 

western Atlantic are not in the Checklists, but their AFS approved common 

names are used in the text and their scientific names appear with the first use of 

their common names. Organisms other than fishes are similarly identified in 

the text. Any common name can be matched with its scientific name by 

looking in the Index for the page number in bold face. Departures from AFS 

names are discussed in the Host Summaries. Parasite common names are not 

used except for a few that are generally accepted. 

Samples of most of the parasites were deposited in the U.S. National 

Museum (USNM) and U.S. National Parasite Collection (USNPC) (see 

Acknowledgments). They are indicated in the text by one of those acronyms 

followed by their deposition number, or by the acronym alone where a 

deposition number was not received prior to publication of this book. 

Collection numbers for specimens deposited in the British Museum of Natural 

History (BMNH) and Museum of the Canadian Department of Fisheries and 

Oceans, Atlantic Reference Collection (ARC) are also listed.

PROTOZOA (PROTOZOANS) 

MYXOZOA (MYXOZOANS) 

Protozoa were once considered a single phylum in the Animal Kingdom. 

Later, they were classed as a subkingdom, containing a number of phyla, in the 

Kingdom Protista. Today they are classified among several phyla in 2 

kingdoms. In this section, we examine protozoan parasites including members 

of the Phyla Sarcomastigophora (amoebas and flagellates) and Apicomplexa 

(coccidians) in the Kingdom Protista, and the Myxozoa (myxozoans) in the 

Phylum Cnidaria in the Animal Kingdom (Siddall et al. 1995). We list these 

forms adjacent to each other as they were traditionally presented. 

As a group, protozoa are essentially complex, unicellular, microscopic 

organisms. They differ from the basic animal or plant cell by having additional 

morphological and physiological characteristics. Protozoa have one to several 

nuclei; multiple nuclei can be either identical or different. Flagellates usually 

have few, relatively long flagella for locomotion and one nucleus. Ciliates 

usually have many, relatively short cilia for locomotion and 2 types of nuclei. 

Coccidians have a group of specialized organelles forming the apical complex 

that is visible only by using the electron microscope. Reproduction may be 

asexual by binary fission, multiple fission, external budding or internal 

budding; or sexual by fusion, conjugation or autogamy. Myxozoans are characterized 

by producing multicellular spores with 2-6 external valves containing 

1-6 (usually 2) polar capsules and 1-2 infective units. Polar capsules are 

essentially identical to the nematocysts (stinging cells) of cnidarians. 

Myxozoans have complex life cycles involving an intermediate host where 

sexual reproduction takes place. The spores produced from the intermediate 

host are infective to the fish host. Protozoan lifestyles range from free-living 

through various forms of commensalism to parasitism inhabiting animals, 

plants, and even other protozoans. Amoebas, flagellates and ciliates live in a 

range of habitats from freeliving to parasitic, while all coccidians and 

myxozoans are parasites. Most consume solid food (holozoic) or fluid 

(saprozoic), but a few photosynthesize and make their own energy (holophytic 

or photo-autotrophic).Parasitic protozoa kill, mutilate and debilitate more 

people in the world than any other group of disease organisms. Protozoan 

parasites of fishes, however, are not known to infect humans but can directly 

transmit microbial diseases to other fishes. Protozoans sometimes kill marine 

or freshwater fishes in Puerto Rico (Bunkley-Williams and Williams 1994, 

1995). Many free-living species occur abundantly in salt water here, consuming 

dead material in all habitats. Some of these can occur in fishes under certain 

conditions, but others are specific pathogens of fishes. In Puerto Rico, we have 

found protozoa to rarely parasitize the skin, fins, gills, and blood of marine 

fishes. 

There are more than 65,000 described species of protozoa with half being 

fossil (useful in identifying oil deposits) and about 8800 parasitic species, 

including 2500 ciliates and 1800 flagellates. More than 1300 parasitize fishes. 

11

12 


In addition to the protozoans, there are almost 1200 species of myxozoans which 

parasitize fish. Protozoans and myxozoans vary in length from 1 Φm to 7 cm 

or more, but most are 5-250 Φm. Many thousands of species remain to be 

described and about one new species is described every day! 

Popular Reference - "How to Know the Protozoa" (Jahn, Bovee and Jahn 

1979). 

Reference - Lom and Dyková (1992), Lee, Hutner and Bovee (1985). 

Classification and Contents 

Kingdom Protista - one-celled organisms Page 

Subkingdom Protozoa 

Phylum Sarcomastigophora - amoebas and flagellates 

Class Zoomastigophorea 

Order Kinetoplastida 

Family Trypanosomatidae 

Trypanosoma sp. ................................................................................... 13 

Phylum Apicomplexa - coccidians and gregarines 

Class Sporozoasida 

Subclass Coccidiasina 

Order Eucoccidiorida (suborder Adeleorina) 

Family Haemogregarinidae 

Haemogregarina bigemina ................................................................ 13 

Order Eucoccidiorida (suborder Eimeriorina) 

Family Eimeriidae 

Goussia clupearum ............................................................................ 15 

Subclass Piroplasmasina 

Order Piroplasmida 

Family Haemohormidiidae 

Haematractidium scombri ............................................................... 15 

Kingdom Animalia 

Phylum Cnidaria - jellyfish, anemones, corals 

Class Myxozoa - myxozoans 

Order Bivalvulida 

Family Kudoidae 

Genus Kudoa .................................................................................. 16 

Kudoa clupeidae ....................................................................... 16 

Kudoa crumena ........................................................................ 17 

Kudoa histolytica ...................................................................... 17 

Kudoa nova .............................................................................. 18


Trypanosoma sp. of Saunders (1958) 

This West Indian blood flagellate infects the young 

of a variety of marine fishes, including the offshore big 

game fishes that spend sufficient time inshore. 

Name - This parasite remains undescribed but we 

assume that what we found is the same as that 

observed by Saunders (1958). It is possible that these 

records represent more than 1 species. 

Diagnostic Characters - This flagellate has a single 

flagellum at one end, and swims freely in the blood 

(not inside blood cells). 

Records - We found light infections in 1 of 8 crevalle 

jack, 2 of 10 great barracuda and in a variety of coral 

reef fishes from various localities around Puerto Rico. 

A light infection occurred in 1 of 137 great barracuda in the Florida Keys. 

Geographic Range - West Indies. 

Life History - Abundant in young fishes, but few remain in adult hosts. 

Ecology - This blood parasite is limited to inshore fishes because it is 

transmitted by leeches, predominantly Trachelobdella lubrica(Grube), which 

do not occur on offshore fishes. 

Location in Host - Free swimming in blood. 

Length - Approximately 50.0 Φm. 

Host Specificity - There is a great variation in fishes infected in different 

localities which seems to be related to the availability of vectors. It can 

apparently infect any inshore fish. Crevalle jack is a new host for this parasite. 

Damage to Host - More abundant in and probably more damaging to young 

fishes. Fishes infected by this trypanosome may not grow as quickly or as 

large. 

Detection - Free swimming flagellates are easy to see in saline-diluted blood 

mounts viewed with a compound microscope. 

Preparation for Study - A thin blood smear must be air dried on a microscope 

slide, fixed with 100% methyl alcohol, stained with Giemsa, and examined 

with the oil immersion lens of a compound microscope. Stained blood smears 

can be permanently mounted with commercial mounting medium and coverslips. 

Haemogregarina bigemina Laveran and Mesnil 

This cosmopolitan parasite of red blood cells infects a 

variety of fishes, including offshore big game fishes. 

Name - The marine members of this genus, producing 2 

or more elongate structures called gamonts per red blood 

cell, are called "schizohaemogregarines", and may repre- 

sent a new genus. The wide geographic distribution and 

broad host choices of this parasite suggest that it could 

represent several species. However, its occurrence in 

13

14 


many wide-ranging big game fishes could provide an avenue for spreading a 

single species to the reported localities. 

Diagnostic Characters - This parasite lives inside the cytoplasm of red blood 

cells. In each infected red blood cell, there are 2 gamonts that are slender at 

one end and clubbed at the other. Infected red blood cells are usually disfigured 

and the nucleus displaced by the parasites. 

Records - Light infections occurred in 1 of 10 crevalle jack, 1 of 1 frigate tuna 

and 1 of 9 yellow jack in Puerto Rico; blue runner and great barracuda from 

Bimini, Bahamas; 2 of 46 bar jack, 1 of 3 dolphin and 2 of 16 great barracuda 

in Bermuda; and 2 of 17 Atlantic sailfish, 3 of 63 blue runner, 1 of 83 cero, 1 

of 3 crevalle jack, 6 of 137 great barracuda, 8 of 37 greater amberjack and 1 of 

62 king mackerel in the Florida Keys, USA. It possibly occurred in Atlantic 

mackerel from the northwest Atlantic (RTLA 4752), and has been reported in 

other fishes from Canada, Europe and the Pacific. 

Geographic Range - Worldwide. 

Life History - Most fish-blood protozoans are thought to be transmitted by 

leeches, but this haemogregarine is apparently transmitted by parasitic isopods. 

Slender free gamonts, and infective stages called oocysts and sporozoites were 

found in the gut of Gnathia maxillaris Montagu in Europe. Probably Gnathia 

sp. and other isopods parasitizing offshore game fishes act as vectors. In other 

species in the genus Haemogregarina, sporocysts arise in the oocyst. In H. 

bigemina, oocysts produce sporoblasts, which later segment into sporozoites 

around a residual body. In New Zealand fishes, a division also occurs in white 

blood cells. 

Ecology - This haemogregarine is most numerous in younger fishes. It is 

found in both inshore and offshore fishes as are its isopod vectors. 

Location in Host - It only infects red blood cells (except for the New Zealand 

form which has a stage in white blood cells). 

Length - Gamonts 10.0 Φm. 

Host Specificity - This parasite occurs in a very great variety of inshore and 

offshore fishes. It appears to have little if any host preference. 

Damage to Host - It is more abundant in and probably more damaging to 

young fishes. Fishes infected by this haemogregarine may not grow as quickly 

or as large. 

Detection - Damaged red blood cells are difficult to see in saline-diluted blood 

mounts. A thin blood smear must be air dried on a microscope slide, fixed with 

100% methyl alcohol, stained with Giemsa, and examined with the oil 

immersion lens of a compound microscope. 

Preparation for Study - Stained blood smears can be permanently mounted 

with commercial mounting medium and coverslips. 

Significance to Sport Fishing - Balao, Hemiramphus balao (Lesueur), and 

ballyhoo, Hemiramphus brasiliensis(Linnaeus), are used as live or fresh bait 

for billfishes and swordfishes. These fishes are often infected with H. bigemina, 

but this parasite cannot be transmitted directly to big game fishes.


Goussia clupearum (Thélohan) 

This parasite lightly infects Atlantic mackerel. 

Name - Goussia cruciata (Thélohan) may be a 

synonym. 

Diagnostic Characters - This parasite is found 

in the liver of infected hosts. The spherical 

oocyst contains 4 ellipsoidal sporocysts, each 

with 2 sporozoites. 

Records - It occurred in an Atlantic mackerel 

(RTLA 4422) from Maryland, USA; in Atlantic 

mackerel from France (RTLA 4689, 4699) and sardines from Europe, the 

Mediterranean and Pacific. A similar parasite G. auxidis (Dogiel) was found 

in the livers and spleen of almost all of 400 albacore from Australia, the Coral 

Sea, New Zealand, Tonga and the central south Pacific. 


Life History - Older, larger fishes are more often infected (up to 90% of some 

herrings and sardines). 

Associations - In specimens from France, it occurred with a bacteria 

Mycobacterium sp. (RTLA 4689, 4699). 

Location in Host - The oocysts occur in liver parenchyma. (In a French 

specimen it was found in hepatic and renal granulomas.) 

Length - Oocyst 18.0-25.0 Φm; sporocyst 8.0-12.0 X 4.0-10.0 Φm. 

Host Specificity - It is almost family specific to herrings and sardines 

(Clupeidae) but occasionally occurs on Atlantic mackerel. 

Damage to Host - Pathological changes include infiltration of macrophages 

(specialized white blood cells) around oocysts, necrosis (tissue death) and 

inflammation around oocysts and eventual encapsulation. In heavy infections 

up to 14% of the liver may be replaced by these parasites. 

Significance to Sport Fishing - It appears to injure herrings and sardines 

more often than Atlantic mackerel, but heavy infections could limit stocks of 

these forage fishes, thus reducing the food available for big game fishes. 

Haematractidium scombri Henry 

This is a common but relatively harmless parasite in 

Atlantic mackerel. 

Diagnostic Characters - It is found inside the red 

blood cells of the host fish and is smaller than the 

nucleus of the red blood cell. It has 1, sometimes 2, 

nuclei that appear dark in the light cytoplasm. 

Records - It occurred in Atlantic mackerel from 

Maryland, USA (RTLA 4423); and off the coast of 

England. In the English specimens, 45% of the fish 

were infected, but only 5% of the red blood cells were 

affected. 

Geographic Range - Northern Atlantic. 

15

16 


Location in Host - In the cytoplasm of red blood cells 

Length - 2.0-6.0 Φm. 

Host Specificity - It only occurs in Atlantic mackerel. 

Damage to Host - This parasite distorts the shape of the red blood cells and 

may increase their fragility and lyse them. 

Genus Kudoa Meglitsch 

This genus was named for a famous protozoologist, Dr. R. R. Kudo. It is 

important because most of the 37 described species (and all of the species 

discussed here) dissolve or liquefy fish muscles either while the host is alive or 

very soon after death. Thus they cause enormous economic loss. 

The unique feature of these multicellular parasites is the polar capsules that 

contain a coiled spring-like filament that is extruded when stimulated. They 

may be used to attach to host tissue. The number and arrangement of these 

polar capsules and the shape of the spore are diagnostic for each species. 

Although the polar capsules are the dominant feature of this parasite, the 

sporoplasm (also called the infective unit) is the cell that infects the host. 

Cysts can be seen in fillets held in front of a bright light. This process, 

called "candling", has historically been used to find defects and parasites in 

fishes. Cysts or small amounts of liquefied tissue can be mechanically removed 

from muscle, placed on a microscope slide with a drop of saline and gently 

squashed with a cover slip to release the spores. Spores or cysts can be preserved 

in 5% formalin. Muscle with cysts can be preserved in 10% formalin. 

Kudoa clupeidae (Hahn) 

The presence of this parasite in tunas makes them 

unsuitable for consumption. Liquefaction of flesh may 

occur after the fish is stored on ice. 

Name - Possibly, Kudoa sp., which occurred in the 

muscle of approximately 1% of bluefin tuna from 

Tasmania, Australia (RTLA 2128), is this species. 

Diagnostic Characters - Spores are rounded rectangular 

in apical view. They have a pointed apex when 

seen in lateral view. Trophozoites form spindle shaped 

masses that are easily seen in muscle fibers. 

Records - The muscle was uniformly infected with 3-7 

cysts per 10 cm 2 in an unidentified tuna, Thunnus sp. 

from New Jersey, USA (RTLA 2532); and bluefin tuna 

from the west coast of Africa. 

Geographic Range - Atlantic Ocean. 

Location in Host - Muscle. 

Length - Spore 4.0-5.0 X 6.3-7.5 Φm; cysts in muscle 2.0-5.0 mm. 

Host Specificity - It is almost family specific to herrings and sardines 

(Clupeidae). The few records in scombrids may be accidental infections.

MYXOZOA (MYXOZOANS) 

Significance to Sport Fishing - How often this parasite occurs has been 

employed as a biological tag to distinguish different stocks of fish. 

Kudoa crumena Iversen and Van Meter 

This protozoan heavily encysts in the muscles of 

Spanish mackerels off Florida and causes the flesh to 

rapidly deteriorate. 

Diagnostic Characters - Spores are pouch-shaped, 

with 4 polar capsules and 1 infective unit (sporoplasm). 

Elliptical white pseudocysts are obvious in muscle. 

Record - One in 9 Spanish mackerel from south 

Florida, USA, were found infected. Numerous cysts 

occurred in infected hosts, but few hosts were infected. 

Geographic Range - Unknown. 

Life History - The spores are probably inadvertently 

ingested by the fish host. In the intestine, the 4 polar 

capsules each eject a filament into the lining which 

holds the spore in place and makes an entry hole for its 

infective unit. This stage either reproduces in the 

epithelial cell or moves through the lymphatic or blood system to muscle tissue 

where it forms a pseudocyst, weakly encased by connective tissue of the host. 

Here, spores are produced which either escape when the host dies or are 

attacked and killed by the host (granulomatous inflammation). Pseudocysts are 

liberated either by digestion in the gut of a carnivorous fish or by host 

decomposition and are broadcast over the substrate. 


Length - Spores 7.5 X 9.9 Φm, filament 15.5 Φm; cysts up to 2.6 X 1.7 mm 

wide. Lom and Dykova (1992) mistakenly used Φm instead of mm for the size 

of the pseudocyst, making the macroscopic, spore-containing cyst smaller than 

the microscopic spores. 

Host Specificity - It is only known from Spanish 

mackerel. 

Kudoa histolytica (Perard) 

This parachute-shaped protozoan liquefies the 

muscles of live Atlantic mackerel. 

Name - The specific epithet,"histolytica", refers to its 

ability to liquefy the muscles of its hosts [from Greek 

"histos" = tissue, and "lytic" = dissolve]. 

Diagnostic Characters - The spore looks like a 4pointed 

star when viewed from above, or a parachute 

when viewed from the side. It has 4 polar capsules. 

Records - It occurs in Atlantic mackerel from the 

Atlantic and Mediterranean. 

Geographic Range - Northern Atlantic and Mediterranean. 

17

18 



Length - Spores 4.0-5.0 X 6.3-7.5 Φm. 

Host Specificity - It is only known from Atlantic mackerel. 

Damage to Host - Instead of being confined in host-produced connective 

tissue cysts, it liquefies the host muscle, turning it into a jelly-like mass containing 

fully formed spores. 

Detection - Liquefied areas can be seen in muscle. 

Significance to Sport Fishing - This important sport and food fish is made 

less desirable or unusable by this infection. 

Kudoa nova Naidenova 

These spindle-shape cysts infect a variety of fishes 

including bigeye tuna. It causes the flesh to rapidly 

deteriorate. 

Name - This parasite was formerly confused with K. 

clupeidae. 

Diagnostic Characters - Spindle-shaped cysts contain 

many plasmodia which form 1-8 spores each. Spores are 

pumpkin-shaped with 4 lobes and 4 polar capsules. 

Records - It occurred in bigeye tuna off the Atlantic coast of the USA. 

Geographic Range - Atlantic, Mediterranean and Black Seas. 

Ecology - It is found in both oceanic and coastal areas. 


Length - Spores 5.3-6.5 X 8.5-9.8 Φm; cysts up to 7.0 mm. 

Host Specificity - It occurs in a variety of fishes. 

Significance to Sport Fishing - This parasite causes postmortem liquefaction 

of the muscle tissues of fishes. Bigeye tuna are made less desirable to unusable 

by this infection.

FUNGI (FUNGUS) 

The fungal-like organisms that parasitize fishes were once thought to be 

simple and similar. These disease organisms are actually so diverse that they 

belong in 2, possibly 3, Kingdoms. Paecilomyces sp. (Rand, Bunkley- 

Williams and Williams 1997), the causative agent of tilapia-wasting disease 

(Bunkley-Williams and Williams 1995) is in the Kingdom Fungi; Saprolegnia 

spp., which causes problems in Puerto Rican and worldwide fresh and brackish 

water fishes, is in the Kingdom Protista; and Ichthyophonus hoferi, discussed 

below, is in an undermined Kingdom somewhere between the Animal and 

Fungi Kingdoms (Ragan et al. 1996). The name "fungus" comes from the Latin 

for "mushroom". The familiar mushrooms, lichens, yeasts and molds are in this 

kingdom. Fungi are the largest and the longest living organisms on earth. The 

forms found in fishes are not known to be dangerous to humans. Parasitic 

fungi might be useful in controlling some pest species. More than 100,000 

species are known. They range from microscopic to more than 50 acres in size. 

They come in a variety of shapes and forms. Many resemble plants, but lack 

the green pigment chlorophyll. Elaborate modes of asexual and sexual 

reproduction occur. Most are spread by spores. Some fungi, including diseasecausing 

forms, are called "imperfect" because they either lack sexual 

reproduction, or it has not been observed. Most fungi are free living, often 

feeding on decomposing materials (saprophytic). A few can alternate between 

saprophytic and parasitic existence. Others are completely parasitic (obligate 

parasites). Many organisms are infected by fungal-like organisms including 

big game fishes. A variety of marine fungi, including some that may harm 

humans, have been found in the marine waters around Puerto Rico, but none 

affect local fishes. 

Popular Reference - "Fungal Diseases of Fishes" (Neish and Hughes 1980). 

Reference - Rand (1996). 


Kingdom (undetermined) Page 

Phylum (unnamed) 

Class Ichthyophonacetes 

Order Ichthyophonales 

Family Ichthyophonaceae 

Ichthyophonus hoferi ....................................................................... 19 

Ichthyophonus hoferi Plehn and Mulsow 

This highly dangerous and destructive cosmopolitan organism kills and 

injures Atlantic mackerel, and possibly other big game fishes in the North 

Atlantic. 

Name - The condition is commonly called "ichthyophonus disease". It was 

previously confused with a microsporidian protozoan Ichthyosporidium gigan- 

19

20 


teum (Thélohan), and called ichthyosporidium disease or ichthyosporidiosis. The 

wide range of habitats and hosts for this disease suggest that a number of 

similar fungal species may be involved. 

Regan et al. (1996) erected a new group of fungal-like and extremely damaging 

parasites of aquatic organisms which includes I. hoferi, Dermocystidium 

spp. of fishes and amphibians, Psorospermium haeckelii of crayfish, and the 

"rosette agent" of salmonids. This unnamed Kingdom and/or Phylum was 

tentatively called "DRIP clade" and is phylogenetically at the divergence between 

the Animal and Fungi Kingdoms. 

Diagnostic Characters - The cysts are usually spherical to irregularly shaped 

and opaque to translucent granular with thick, multilaminate walls. It 

sometimes germinates with single or multiple, branched or unbranched thalli 

extending into host tissues. 

Records - Heavy infections occurred 

in Atlantic mackerel off the northeast 

coast of the USA and Europe. Random 

samples of Atlantic mackerel 

from sport fishing creel census conducted 

in New Jersey, USA, found 

5% of apparently healthy fishes had 

light infections. 

Geographic Range - Mass mortalities 

of marine fishes appears to be 

limited to the North Atlantic, but this 

disease is found worldwide in marine, 

brackish and fresh waters. 

Mass mortalities of cultured fishes 

have occurred in Japan and on the 

west coast of the USA. 

Life History - The resting stage 

(cyst) containing a multinucleated 

cytoplasm is usually seen in the tissues of the host. More advanced cysts 

contain a few to hundreds of distinct nuclei. Endospores germinate in the cyst, 

and pseudopodia-like thalli extend out of the cyst and invade new host tissue 

(filamentous germination). Alternatively, the entire contents (plasmodium) of 

the cyst may move into the host tissues and occupy an area of 2-3 times the 

original volume of the cyst (plasmodial germination). Both methods produce 

new resting stages until the host tissues stop further development or the host 

dies. Resting cysts may leave the host through the digestive tract, gills and/or 

skin or en masse from dead and decomposing hosts. Mother cysts produce 

amoeboblasts which await being inadvertently eaten by a new host. Amoeboid 

bodies are released and penetrate through the intestinal wall into the blood and 

become resting cysts in the tissue in which they settle. Host death can occur in 

1 month (acute) to 6 months (chronic).

FUNGI (FUNGUS) 

Ecology - This is an obligate parasite that grows best at 10ΕC but can grow 

between 3-20ΕC, but not at 30ΕC. It can continue to grow in hosts that it has 

killed. Atlantic mackerel on the east coast of the USA may be resistant to this 

disease, but can become infected and die during mass mortalities of Atlantic 

herring, Clupea harengus Linnaeus. Off Europe, mass mortalities of Atlantic 

mackerel have occurred without the stimulus of herring mortalities. 

Location in Host - During mortalities of Atlantic mackerel, the kidney, spleen 

and other visceral organs were destroyed, but no external or muscle lesions 

were present. The disease is systemic, but attacks different tissues in different 

species of hosts. 

Length - Cysts usually 10-300 Φm. 

Host Specificity - It can apparently infect almost any species of marine or 

freshwater fish and has also been reported in amphibians and copepods. 

Damage to Host - Great losses have occurred in cultured fishes and in 

repeated mass mortalities in wild fishes. Mass mortalities may limit the populations 

of herrings in the western North Atlantic. 

Detection - Multiwalled, spherical resting stage (quiescent) cysts are most 

easily seen in fish tissues. Death, emaciation and low body weight are seen in 

many cases. 

Significance to Sport Fishing - This infection reduces the numbers, condition 

and vitality of at least 1 important big game fish. 

Preparation for Study - It can be grown on Sabouraud's dextrose agar or 

glycerine-agar, both with 1% bovine serum added. Bacterial overgrowth can 

be reduced by adding 0.25 g/l each Streptomycin and Penicillin into the media 

before pouring the plates. 

Disease Prevention - Fishes or even frozen or processed fish products from 

the north Atlantic or other areas of known infections should not be cleaned or 

processed in other regions where this disease could spread. Feeding infected 

tissues spreads and intensifies the disease in new hosts. Although this disease 

is assumed to be worldwide, it behaves quite differently in different regions. 

Different regional strains, or even species may occur, and should not be spread 

to other geographic areas. 

21

22 

PLATYHELMINTHES (FLATWORMS) 

Flatworms form a phylum of soft-bodied, bilaterally symmetrical, flattened, 

worm-shaped animals. Usually each worm has a set of both female and male 

reproductive organs (hermaphroditic). They either have a primitive blind gut 

and a mouth, or absorb nutrients through their bodies. They respire through 

their skin and possess specialized cells that secrete ammonia waste products. 

There are about 20,000 species of flatworms including copepod worm, flukes, 

tissue flukes, gillworms, and tapeworms. 


Phylum Platyhelminthes - flatworms Page 

Class Udonellidea - copepod worm ............................................................ 22 

Class Trematoda - flukes and soleworms 

Subclass (Infraclass) Digenea - flukes ...................................................... 24 

Superfamily Didymozoidea - tissue flukes ...................................……. 64 

Class (Infraclass) Monogenea - gillworms ................................................ 79 

Class Cestoda (or Cestoidea) - tapeworms ................................................ 103 

UDONELLIDEA (COPEPOD WORM) 

The species below apparently represents a class of flatworms. No common 

name has been given to this animal, so we suggest copepod worm. This worm 

parasitizes fish-parasitic copepods. Some of these copepods, in turn, parasitize 

big game fishes! It has also been found on fish lice and fish-parasitic isopods. 

It superficially resembles a gillworm, but is so drastically different it must be 

separated. This distinction is based on the anatomy of the protonephridial 

excretory system. It also differs from gillworms by having a simple sac for an 

intestine and no hardened (sclerotized) attachment organs. 


Class Udonellidea - copepod worm Page 

Order Udonellida 

Family Udonellidae 

Udonella caligorum ..................................................................... 22 

Udonella caligorum Johnston 

This tiny parasite is seldom seen as an adult on parasitic copepods, but their 

eggs are occasionally found attached to the cuticle of copepods parasitic on big 

game fishes. 

Name - Several genera and a number of species have been described, but all 

appear to be synonyms.

UDONELLIDEA (COPEPOD WORM) 

Diagnostic Characters - The cylindrical body has a posterior sucker-like 

attachment organ that is not divided into sections and is not armed with anchors, 

bars or spines. 

Records - We found 1 on a male Brachiella thynni in a wahoo, 1 each on 3 

Euryphorus brachypterus in a yellowfin tuna;a nd on Caligus bonito in a snapper 

from La Parguera, Puerto Rico. It has also been reported from a little tunny off 

Chesapeake Bay, USA; and yellowfin tuna in the Pacific. 


Life History - This hyperparasite lives its whole life on para- 

sitic crustacea. The elongate, pyriform operculate eggs are 

attached in clusters to the crustacean by an adhesive disk with 

a thread attached to the egg. The young hatch out of the 

egg and are capable of attaching to the crustacean (no free 

swimming, ciliated larval stage). They possess both male and 

female organs but cross fertilize each other. This direct life 

cycle allows large numbers of them to build up and some 

copepods are "furry" with eggs. 

Ecology - This worm transfers from one copepod to another 

when these crustacean parasites are mating (a sexual disease 

of copepods). It has been found on free swimming parasitic 

copepods captured in plankton nets. 

Associations - Usually found attached on a great variety of 

caligoid copepods, but has been reported on other genera of 

copepods; a fish louse, Argulus sp.; an isopod, Livoneca 

vulgaris Stimpson; and on the gills of host fishes. 

Location in Host - Eggs are usually attached to the posterior part of a 

copepod, immature worms attach to the lateral margins of the carapace, and 

adults to the central and lateral parts of the carapace. Most worms are on 

copepods that parasitize the gills and mouth of fishes. One of our specimens 

occurred in a cavity beneath the pectoral fin of a wahoo. 

Length - 1.1-2.3 mm 

Host Specificity - This worm does not appear to prefer any particular fish 

species, but does prefer flat, round crustacean parasites, particularly copepods 

in the genus Caligus. Brachiella thynni is a new, and unusually shaped, host for 

this worm, and wahoo is a new fish host. 

Significance to Sport Fishing - This worm may feed on parasitic copepods 

instead of the fish host, and certainly covers these crustacean parasites with eggs 

that are probably detrimental. Oddly enough, this parasite may harm other 

parasites and may actually benefit the fish host. 

23

24 


DIGENEA (FLUKES) 

Flukes or digeneans (formerly called digenetic trematodes) form a class of 

flatworms. Flukes reproduce as adults and again as larvae, hence the name "digenetic" 

or 2 births. They cause serious and fatal diseases in many animals 

including humans. Bilharzia (Schistosoma mansoni Sambon) in humans is found 

in Puerto Rico. Flukes are important fish parasites with fishes serving both as 

intermediate (grubs) and final hosts (flukes). More than 9000 species have been 

described. Adults range in size from 10 cm. One of the 

largest, Hirudinella ventricosa, occurs in the stomach of local wahoo. Flukes 

usually look like typical flatworms with a mouth in the anterior region, and a 

blind gut and reproductive and other organs in the trunk region. Unlike 

generalized flatworms, many have 2 sucker-like holdfast organs. One is located 

near the mouth (oral sucker) and the other is usually in the middle of the worm 

on the same (ventral) side (ventral sucker or acetabulum). Great differences in 

shape, size and orientation of structures occur in different species. The internal 

organs illustrated in the adjacent 

drawing can be seen if worms 

are placed in saline wet 

mounts and viewed with a 

compound microscope. 

Flukes have a complex life 

cycle usually with 2-3 interme- 

diate hosts, possible transfer 

hosts and a final host. In a 

typical life cycle, eggs from the 

body of the fluke pass out the 

intestine of the final (definitive) 

host. The eggs are either eaten 

by the first intermediate host or 

hatch into a swimming ciliated 

larva (miracidium) that infects 

the first intermediate host, 

usually a snail. Once inside the 

snail the miracidium transforms 

into a sporocyst. Each sporocyst 

asexually produces many 

larval parasites (rediae) which 

in turn produce many swimming infective larvae (cercariae) that leave the snail. 

The cercariae infect the second intermediate host, encyst, and become metacercariae. 

If the appropriate final host eats this infected host, the metacercariae are 

digested out of the cyst and become adult flukes. There are many variations in 

the life cycle especially in the asexual phases. Each fluke has both female and 

male reproductive systems (hermaphroditic). Self fertilization is possible in 

some flukes but cross fertilization usually occurs.


Flukes are permanent parasites in most marine fishes, and in many fresh- 

water fishes, amphibians, reptiles, mammals and birds. Larval stages occur in 

a variety of invertebrates and vertebrates. Flukes usually occur in either the in- 

testine, stomach or mouth, or occasionally lungs and other organs. Larval forms 

occur in almost any tissue. Sometimes predator hosts temporarily support flukes 

digested from prey (false hosts), but these soon pass out of the predator. 

Didymozoids or tissue flukes belong in the digenea but their exact position 

and relationships are not clear. The classification will not be resolved until the 

life cycles and early life cycle stages are studied. We consider them separately 

after the flukes. 

Popular Reference "How to Know the Trematodes" (Schell 1970). 


Subclass (Infraclass) Digenea - flukes Page 

Order Hemiuriformes 

Family Hirudinellidae 

Hirudinella ventricosa .................................................................. 27 

Family Bathycotylidae 

Bathycotyle coryphaenae .............................................................. 29 

Family Hemiuridae 

Aponurus laguncula ..................................................................... 56 

Brachyphallus crenatus .............................................................. 56 

Brachyphallus parvus ................................................................... 30 

Dinurus scombri ........................................................................... 31 

Dinurus tornatus ........................................................................... 32 

Ectenurus lepidus .......................................................................... 33 

Hemiurus appendiculatus ............................................................. 34 

Lecithaster confusus .................................................................... 58 

Lecithochirium microstomum ....................................................... 34 

Lecithochirium monticellii ............................................................ 59 

Lecithochirium texanum ................................................................ 35 

Lecithochirium sp. .......................................................................... 35 

Lecithocladium excisum ................................................................. 36 

Myosaccium opisthonema ........................................................... 59 

Opisthadena dimidia ..................................................................... 61 

Parahemiurus merus ..................................................................... 37 

Sterrhurus musculus ....................................................................... 38 

Family Accacoeliidae 

Tetrochetus coryphaenae ................................................................ 39 

25

26 


Family Mabiaramidae 

Mabiarama prevesiculata ..................... ........................................ 40 

Order Strigeiformes 

Family Sanguinicolidae 

Paradeontacylix sanguinicoloides ................................................ 40 

Family Fellodistomatidae 

Claribulla longula ......................................................................... 57 

Tergestia laticollis ....................................... ................................. 41 

Family Bucephalidae ............................................................................ 42 

Alcicornis carangis ........................................................................ 42 

Bucephalopsis arcuata ................................................................... 43 

Bucephalopsis attenuata ................................................................. 56 

Bucephalopsis gracilescens ............................................................ 57 

Bucephalopsis longicirrus .............................................................. 44 

Bucephalopsis longovifera ............................................................. 45 

Bucephalus confusus ...................................................................... 57 

Bucephalus gorgon ......................................................................... 45 

Bucephalus varicus ......................................................................... 46 

Prosorhynchus pacificus ................................................................. 62 

Rhipidocotyle baculum ................................................................... 47 

Rhipidocotyle barracudae ............................................................... 47 

Rhipidocotyle capitata .................................................................... 48 

Rhipidocotyle longleyi .................................................................... 62 

Family Coitocaecidae 

Coitocaecum extremum ................................................................... 49 

Order Lepocreadiiformes 

Family Lepocreadiidae 

Neolepidapedon belizense ............................................................... 60 

Neolepidapedon retrusum ............................................................... 49 

Opechona orientalis ......................................................................... 60 

Pseudolepidapedon pudens .............................................................. 62 

Order Plagiorchiformes 

Family Acanthocolpidae 

Genus Stephanostomum ................................................................... 50 

Stephanostomum aulostomi ............................................................. 63 

Stephanostomum coryphaenae ........................................................ 50 

Stephanostomum dentatum ............................................................. 63 

Stephanostomum ditrematis ............................................................ 51 

Stephanostomum imparispine ......................................................... 52


Stephanostomum megacephalum .................................................... 53 

Stephanostomum sentum ................................................................ 53 

Tormopsolus filiformis ................................................................... 54 

Tormopsolus orientalis .................................................................. 54 

Family Opecoelidae 

Opecoeloides brachyteleus ............................................................ 60 

Opecoeloides vitellosus ................................................................. 60 

Pinguitrema lerneri ....................................................................... 61 

Podocotyle chloroscombri ............................................................ 61 

Podocotyle simplex ....................................................................... 61 

Pseudopecoeloides carangis ........................................................ 55 

Pseudopecoelus elongatus ............................................................ 62 

Family Gorgoderidae 

Cetiotrema carangis ..................................................................... 55 

Family Monorchiidae 

Genolopa brevicaecum ................................................................ 57 

Lasiotocus truncatus ................................................................... 58 

Miscellaneous Flukes ................................................................. 56 

Hirudinella ventricosa (Pallas) 

This giant worm is the only fluke which has frequently been 

brought to us by fishermen. A pair of these stomach worms is 

almost always found in wahoo around the world, and less 

consistently in most tunas and dolphins. 

Name - This giant fluke is so well known by sport fishermen 

that it has been given common names: "giant stomach worm", 

"giant fish stomach worm" and contracted and wrinkled flukes 

are called "walnut worms". Because of its great variability both 

in size and shape, this worm has been called by at least 39 

scientific names. The name H. marinum Garcin was first 

proposed for this fluke in 1730. It has been rejected as the name 

of this worm because it was used before Linnaeus set up the 

present system of scientific names. Hirudinella ventricosa is the 

only valid species in the genus, thus the 14 other species names 

still in use are actually synonyms. 

Diagnostic Characters - This small to very large fluke is 

unmistakable in the stomach of wahoo. It is a fleshy worm that 

varies in size and shape, with extended worms the size of a mans 

finger and contracted ones about the size of a walnut. Smaller 

worms are still massive but they are capable of contorting into 

27

28 


many shapes. It is brown to pinkish in color. The 2 suckers are easily seen and 

are close together on the anterior end of the worm. 

Records - We found 1 in an albacore, 1-4 in 23 of 40 

Atlantic blue marlin, 1 in a blackfin tuna, 1-3 in 16 of 40 

dolphin (USNPC 84887), 1-4 in 3 of 6 little tunny, 3 in a 

longbill spearfish, 1-2 in 3 of 8 skipjack tuna, 2-4 (usually 

2) in 15 wahoo (USNPC77758-60) from various localities 

around Puerto Rico; 3 in 2 of 5 dolphin, 1 in a little tunny, 

2 in each of 4 wahoo; and 4 in 1 of 2 yellowfin tuna from 

Dauphin Island, Alabama, USA. It occurred in a little 

tunny in the Gulf of Mexico off Mexico; and 19 in 1, and 

2 in 3 of 13 wahoo from Bermuda. One to 6 occurred in 

8 of 36 white marlin off Delaware and Maryland, USA 

(USNPC 80328) and 1-8 in 62 of 303 swordfish from the 

northwest Atlantic (ARC 2318); and in albacore, Atlantic 

blue marlin, Atlantic bonito, Atlantic sailfish, chub 

mackerel, little tunny, bluefin tuna, skipjack tuna, 

swordfish and yellowfin tuna from the Gulf of Mexico 

and Atlantic coasts of USA. This worm occurred once each in a pilotfish off the 

Canary Islands; in a great barracuda from the Atlantic, and a king mackerel 

from Brazil, but these may have been accidental infections. We also found it 

in wahoo from the central Pacific. It also occurs in black marlin and frigate 

tuna from the Indo-Pacific. In 14 localities across the Pacific, an average of 0.4 

(0.0-1.1) worms occurred in 878 skipjack tuna. Less than 1% of the skipjack 

tuna off Florida had this worm, but 40% of those off Brazil, 7% off West 

Africa, 21% off the Marquesas in the Pacific, and 34% off Hawaii were 

infected. 


Life History - Adults occur over a broad range of sizes. It feeds on blood, 

thus the gut of the parasite is usually filled with black, digested blood. Eggs are 

relatively small for the size of this worm. Studies have suggested that larger 

eggs occur in larger worms, but others have found egg size to be rather erratic. 

Ecology - Wahoo, particularly those shorter than 160 cm, almost always have 

a pair of these giant worms in their stomach. Two wahoo longer than 160 cm, 

off La Parguera and off Mona Island had 3 and 4 worms. Other hosts are more 

variable in the occurrence and numbers of this parasite. 

Associations - We examined 13 dolphin collected off La Parguera, Puerto 

Rico, for total parasites. Five of these fish had 2-3 of these worms, and 

thousands of other flukes in their stomachs. However, there was no obvious 

relationship between these worms and other species of flukes. 

One worm was found in a 10 mm long 2 mm wide stomach ulcer of an 

albacore from Desecheo Island. The relationship between stomach ulcers and 

stomach parasites in big game fishes is not certain. We have seen injuries to the 

stomach lining of wahoo caused by these worms, but could not be certain that 

these were not caused after the death of the host.


Location in Host - Stomach. 

Length - 8.5-100.0 mm. Live worms 

may extend to 170 mm. 

Host Specificity - Wahoo is the 

preferred host of this parasite, both by 

being almost always present, and by 

achieving a consistently large size in 

this host. It is a characteristic parasite 

of wahoo and a primary parasite of 

Atlantic blue marlin, dolphin, little 

tunny and possibly other scombrids. It 

is a secondary parasite of other 

scombrids and billfishes. Blackfin tuna, 

frigate tuna and longbill spearfish are 

new hosts for this parasite. 

Damage to Host - These very large 

worms cause wounds by penetrating the 

stomach lining to feed on blood and 

absorb a considerable amount of blood. 

Fortunately, few worms occur per host. 

Usually only 1 or 2 of the worms present in the stomach of a host are very 

large. Some mechanism of the parasites or the hosts appears to regulate both 

the numbers and sizes. 

Preparation for Study - The thick skin (tegument) of this parasite protects it 

from acid and churning food particles in the stomach, but makes the penetration 

of chemicals more difficult and preparations more likely to contract or distort. 

Specimens that have been partially damaged by long contact with stomach acid, 

after the death of the host, are particularly difficult to prepare. Worms removed 

from the stomach can be kept alive temporarily in a small amount of saline held 

in a refrigerator until preserved. 

Significance to Sport Fishing - The wounds they produce and sheer size must 

have a negative, if unstudied, impact on big game fishes. They have also been 

used as biological tags. 

Bathycotyle coryphaenae Yamaguti 

Geographically widespread, but little studied, this parasite of dolphin 

attaches in the gills, an unusual location for flukes. 

Name - Burnett-Herkes (1974) reported B. branchialis Darr in dolphin, but we 

suspect it was this fluke. 

Diagnostic Characters - This moderate-sized, elongate fluke has the ventral 

sucker located on a projection from the body. The oral sucker is smaller than 

the ventral sucker and moderately separated from it (more than 2, less than 3 

ventral sucker diameters). There is a hood-like lobe that hangs over the oral 

sucker and the ventral sucker is enclosed by a fold of the body wall. No tail is 

present. 

29

30 


Records - We found 1-10 in 12 of 40 dolphin from various 

localities around Puerto Rico; and 1 in 1 of 5 dolphin from 

Dauphin Island, Alabama, USA. From 1-14 occurred in 75 of 

125 dolphin from the Straits of Florida; and in this host from 

Louisiana, USA; and Japan. 

Geographic Range - Worldwide. Our collections are the first 

in the Caribbean and northern Gulf of Mexico. 

Life History - It does not parasitize the smallest dolphin but 

was found in fish from 12.1-138.4 mm in standard length. It 

gradually increases in numbers in larger fish and may thus 

accumulate over time. 

Associations - The numbers of this fluke were positively 

correlated with the numbers of Caligus productus and 

Euryphorus nordmanni on the gills of dolphin, which suggests 

that they may, in some way, aid each other in parasitizing this 

host. 

Location in Host - Attached to gill filaments. 

Length - 13.0-24.0 mm. 

Host Specificity - This worm only occurs on dolphin, but not 

often enough to be a characteristic parasite. 

Damage to Host - Moderate infections may injure dolphin. 

Brachyphallus parvus (Manter) 

This fluke parasitizes a variety of hosts including a few 

big game fishes. 

Name - This parasite has, at various times, been placed in 

the genera Brachyphallus, Lecithochirium and Sterrhurrus, 

but this peculiar worm does not appear to fit well in any 

described genus.The name Lecithochirium parvum was 

used for this worm in the records from Curaçao and 

Jamaica. 

Diagnostic Characters - It is a microscopic to minuscule, 

elongate fluke with the body either bulging around the 

ventral sucker or with the ventral sucker on a projection off 

the body. The oral and ventral suckers are moderately 

(approximately 1 ventral sucker diameter) separated. A 

short tail is usually withdrawn into the body. The oral 

sucker is more than 1/3 and less than 1/2 the diameter of the 

ventral sucker. The vitellaria are reduced to 2 compact, 

lobate masses just posterior of the ovary, and the testes are 

one in front of the other with the anterior one partially 

underneath the ventral sucker. Overstreet (1969) found 

considerable variation in specimens of this fluke. The tail may be extended or not, 

with the ceca extending into the tail or not, the testes may be in contact or 

separated, the sizes of the oral sucker and ventral sucker and ratios of these sizes 

varied, sizes of the eggs varied. He


found that the small size of this worm, pre-acetabular pit, weakly-developed 

cirrus sac and low number of postovarian coils of the uterus, were consistent 

characters that allowed this worm to be identified. 

Overstreet (1969) also pointed out that this species lacks the deep striations 

(cuticular plications) on the sides of the body that are characteristic of this 

genus, and placed it in Lecithochirium. Yamaguti (1970) stated it could not go 

into Lecithochirium because it lacks a prostatic vesicle characteristic of this 

genus. He did not address the striations question directly, but added "usually" 

to that character in the genus diagnosis. 

Records - It occurred in little tunny from La Parguera, Puerto Rico (USNPC 

39403); 2 in 1 of 3 crevalle jack, 2 in 1 of 18 horse-eye jack and 2 in 1 of 8 

yellow jack from Jamaica; and 1 in a greater amberjack from Curaçao. It was 

found in 1 of 3 little tunny from Dry Tortugas, Florida, USA (USNPC 37018); 

in 1 of 2 blue runner from south Florida; and in frigate tuna and yellowfin tuna 

from the western Atlantic. 

Geographic Range - Western Atlantic. 

Associations - The fluke, Lecithochirium texanum, occurred with this worm in 

a little tunny examined in the Dry Tortugas. 


Length - 0.5-1.2 mm. 

Host Specificity - This worm appears to have little host preference. 

Dinurus scombri Yamaguti 

This stomach fluke is found worldwide in little tunny. 

Name - The name, quite appropriately, refers to the family 

of the hosts. 

Diagnostic Characters - This is a tiny, elongate fluke with 

the oral and ventral suckers close together and approximately 

the same size. The tail is less than 1/3 of the total body 

length. The tubular mass of vitellaria is confined in the 

midbody and does not nearly reach the tail. 

Records - We found 2-3 in 2 of 5 Atlantic sailfish from 

various localities around Puerto Rico; 2 in a longbill spearfish 

from Aguadilla, Puerto Rico; and in a little tunny from 

Dauphin Island, Alabama, USA. One occurred in 1 of 3 little 

tunny from the Dry Tortugas off Florida, USA; and in chub 

mackerel, frigate tuna and other scombrids from Japan. 

Geographic Range - Worldwide. Our collections are the 

first in the Caribbean and northern Gulf of Mexico. 


Length - 3.6-4.8 mm. 

Host Specificity - This worm is family specific to scombrids, 

but insufficient information is available to assess its individual 

host preferences. Our records in Atlantic sailfish and longbill spearfish may 

31

32 


represent false hosts resulting from these predators eating scombrids. These 2 

billfish species are also new hosts for this parasite. 

Dinurus tornatus (Rudolphi) 

This large and impressive fluke occurs in the stomach of 

dolphin around the world. We found harmful superinfections in 

Caribbean dolphin. 

Name - Traditionally, 4 species, D. barbatus (Cohn), D. 

breviductus Looss, D. longisinus Looss and D. tornatus, are 

recognized in the stomach of dolphin.Dawes(1968) suggested 

that these worms were 1 species. We struggled to separate 

these forms, but found that the characters used to separate the 

species intergrade and are not mutually exclusive. We conclude 

that only 1, highly variable, species of this genus occurs in 

dolphin. Dinurus coryphaenae Yamaguti and probably D. 

hippurus Nadakal, Kappikarayil and Jacob are also synonyms. 

Diagnostic Characters - This is a highly variable, minuscule 

to large, elongate fluke with the oral and ventral suckers close 

together. The oral sucker is less than 1/2 to 2/3 the diameter 

of the ventral sucker. The tail is more than 1/3 to more than 

1/2 of the total body length. Deep striations (cuticular 

denticulations) occur on the sides of body. The egg filled 

uterus extends into the tail and often 1/2 way down the tail. 

The intestinal ceca extend to the end of the tail. The winding vitellaria extend 

from the posterior body into the tail. 

Records - We found 3-2805 in 40 dolphin from various localities around Puerto 

Rico (USNPC 39404-6,84882-4); 1 in 1 of 3 bar jack from Cabo Rojo, Puerto 

Rico (USNPC 85292); and 100-200 in 5 dolphin from Dauphin Island, Alabama, 

USA. It also occurred in dolphin and pompano dolphin from Cuba; more than 

200 in a dolphin from Bimini, Bahamas; 1-3 in 3 dolphin from Curaçao; in 3 

of 6, 1 of 6, and 2 of 6 dolphin from the Dry Tortugas, Florida, USA; in 

dolphin and pompano dolphin from Louisiana and North Carolina; in dolphin 

from Brazil; in Atlantic bonito and skipjack tuna in the western Atlantic; more 

than 200 in each of 2 dolphin from the Pacific coast of Panama; and in other 

scombrids in the Pacific. 


Life History - Metacercaria occur in the skin and muscle of herrings, possibly 

other fishes, and in pelagic crustacea. 

Ecology - The superinfections we found in the Caribbean have not been 

recorded elsewhere. Possibly, infections are more severe and damaging to 

dolphin in the Caribbean than in the Gulf of Mexico or Atlantic coasts of North 

and South America. Manooch, Mason and Nelson (1984) failed to find this 

fluke in 2632 dolphin collected along the Gulf and Atlantic coasts of the USA 

from Texas to North Carolina. A very heavy infection of 250 flukes was 

reported in a dolphin from India.


Associations - We found 5-2805 worms in 13 dolphin from off La Parguera. 

Hundreds of other parasites occurred with this parasite, but none showed any 

correlation with the numbers of this worm. 


Length - 2.0-22.5 mm. 

Host Specificity - This fluke is a characteristic parasite of dolphin and 

probably pompano dolphin. It appears to prefer dolphins to scombrids by 

having higher numbers and occurring in greater frequency. Some records from 

scombrids may represent false hosts from dolphin prey and/or misidentifications 

of D. euthynni Yamaguti or D. scombri. Bar jack is a new host for this 

parasite. 

Damage to Host - Superinfections of this worm must damage and limit the 

growth of this commercially and recreationally valuable fish. Encysted metacercariae 

cause "black-spot disease" in herrings from Argentina. 

Ectenurus lepidus Looss 

This parasite is found worldwide in the stomach of jacks 

and other fishes. 

Name - Ectenurus americanus (Manter), E. virgulus Linton and 

E. yamaguti Nahhas and Powell appear to be similar to this 

worm and may be synonyms. 

Diagnostic Characters - It is a minuscule to tiny, elongate 

fluke with the oral and ventral suckers close together. The oral 

sucker is less than 1/2 the diameter of the ventral sucker. The 

tail is more than 1/3, but less than 1/2 the total body length. 

Deep striations (cuticular denticulations) occur on the sides of 

body. The uterus containing eggs extends into the tail; that the 

intestinal ceca stop well short of the end of the tail; and that the 

winding vitellaria are confined in midbody and do not extent to 

the tail. 

Records - One occurred in a blue runner, 2 each in 2 of 3 

crevalle jack, 1 in 1 of 4 greater amberjack, an unknown 

number in 18 horse-eye jack and 1-2 in 2 of 8 yellow jack from 

Jamaica; 2 in 1 of 19 yellow jack from Belize (USNPC 74267); 

1-19 occurred in 52 bar jacks from Bermuda; and 3 in 1 of 2 

yellow jacks from the Dry Tortugas off Florida, USA. Despite extensive 

examinations of these same hosts in Puerto Rico, we have not seen this worm. 

It has also been reported in Atlantic bonito, rainbow runner and a variety of 

other fishes throughout the western Atlantic. 


Associations - In 52 bar jacks from Bermuda infected with this worm, 10 had 

double infections with Alcicornis carangis, 7 with Prosorhynchus pacificus, and 

10 had triple infections with all 3 flukes. 

Location in Host - Stomach or gills. Flukes in the gills have been reported by 

several authors, and probably indicate this worms' ability to leave dead fishes. 

33

34 



Host Specificity - This parasite has been reported from a wide variety of 

fishes, but it may prefer jacks. In Bermuda, it is a characteristic parasite of bar 

jack. 

Hemiurus appendiculatus (Rudolphi) 

This fluke occurs in a great variety of fishes but seldom 

in big game fish. 

Diagnostic Characters - A microscopic to tiny, elongate 

fluke with a short tail that is deeply embedded into the body. 

The oral and ventral suckers are rather close together, but 

separated by more than 1 width of the ventral sucker. The 

oral sucker is less than 1/2 as large as ventral sucker. Deep 

striations (cuticular denticulations) occur on the sides of the 

entire body. 

Records - One occurred an in Atlantic mackerel and 2 in a 

banded rudderfish, Seriola zonata (Mitchill), (USNPC 8327) 

from Massachusetts, USA. Also reported in a variety of 

fishes from the temperate Atlantic from Europe to the north 

Atlantic coast of USA and Canada, Mediterranean and Black 

Sea. 

Geographic Range - Northern Atlantic, Mediterranean and 

Black Sea. 


Length - 0.4-4.5 mm. 

Host Specificity - This fluke appears to have little host 

preference as it occurs in a great variety of inshore fishes. 

The 1 report from an Atlantic mackerel may have been an 

accidental infection or a false host. 

Lecithochirium microstomum Chandler 

This fluke occurs worldwide and has generalized host preferences. It is 

often found in big game fishes. 

Name - Manter (1947) thought that this fluke was similar, if not identical, to 

L. magnaporum Manter which occurs in a variety of Pacific big game fishes. 

We consider it a synonym. Yamaguti (1971) thought it was similar to and often 

confused with L. priacanthi Yamaguti, but did not synonymize them. 

Diagnostic Characters - It is a minuscule to tiny, bullet-shaped to oval fluke 

with the oral and ventral suckers well separated (more than 1 ventral sucker 

diameter). A moderate (approximately 1/3 of total body length) tail is usually 

not withdrawn into the body. The oral sucker is approximately 1/3 the diameter 

of the ventral sucker. The vitellaria are reduced to 2 compact, lobate masses 

just posterior of the ovary; that the testes are side-by-side diagonally, in contact 

and slightly separated from the ventral sucker; and that the uterus with eggs 

extends to the end of the intestinal ceca and both extend into the tail.


Records - Two occurred in 1 of 4 greater amberjack from 

Jamaica; and 6 in 1 of 3 pompano dolphin from Pass-a-Grille, 

Florida, USA. It was also found in skipjack tuna from the 

Pacific coast of Mexico, frigate tuna from Hawaii and in a 

variety of Pacific big game fishes. 


Life History - Pearse (1949) found a "young" fluke in a clam, 

ponderous ark, Noetia ponderosa (Say), from North Carolina 

and suggested that it was this worm. 



Host Specificity - This fluke probably prefers Atlantic 

cutlassfish, Trichiurus lepturus Linnaeus, since this fish has the 

heaviest infections (up to 250), but it occurs in a variety of 


Lecithochirium texanum (Chandler) 

This tiny stomach fluke parasitizes 

Atlantic bonito, little tunny and other fishes in the 

Gulf of Mexico. 

Name - Sterrhurus texanus C. is a synonym and Lecithochirium 

sp. of Overstreet, 1969 may be a synonym. Manter (1947) 

suggested that this worm was a synonym of S. imocavus Looss 

which was named in a tuna, Thynnus sp., from Egypt. 

Diagnostic Characters - It is a minuscule to tiny, elongate fluke 

with an ventral sucker that is relatively large and almost as wide 

as the body. The tail is relatively short and less than 1/5 of the 

body length. The ovary is widely (more than 4 diameters) 

separated from the testes. 

Records - It occurred in an Atlantic bonito from Texas, USA; 

and heavy infections in 3 little tunny from the Dry Tortugas off 

Florida, USA. 

Geographic Range - Gulf of Mexico. 

Associations - Brachyphallus parvus occurred with this worm in 

a little tunny examined from the Dry Tortugas. 


Length - 1.9-3.6 mm. 

Host Specificity - This worm is possibly family specific to scombrids. 

Lecithochirium sp. 

This new fluke occurs in low numbers in a variety of fishes in Puerto Rico 

including black jack and king mackerel. 

Name - This worm is a new species which we are in the process of describing. 

Siddiqi and Cable (1960) apparently assumed this worm was L. microcercus 

(Manter) because it occurred in the same host in the Dry Tortugas off Florida, 

35

36 


USA, and in Puerto Rico (USNPC 39402). We followed 

their lead and identified a series of specimens from Puerto 

Rico with the aid of their illustration. Only in preparing 

this book did we examine the original description and 

illustrations of L. microcercus. The Puerto Rican worms 

are similar to L.microcercus, but differ by having a genital 

pore that opens posterior to the oral sucker instead of on 

top of it [as Manter (1947) emphasized], widely separated 

(almost 1 testis diameter apart) testes instead of closely 

spaced almost touching testes, testes that are separated from 

the ventral sucker by approximately the diameter of a testis 

instead of being under the ventral sucker, a uterus that does 

not extend posteriorly passed the vitellaria instead of 

beyond the ends of the ceca and to the posterior end of the 

worm, and a uterus that is confined between the ceca 

instead of winding around and beyond the ceca. 

Diagnostic Characters - A microscopic to minuscule, bullet-shaped fluke with 

the oral and ventral suckers moderately (less than 1 ventral sucker diameter) 

separated, and a short tail that is withdrawn into the body. The oral sucker is 

about 1/2 the diameter of the ventral sucker. The vitellaria are reduced to 2 

compact, lobate masses just posterior of the ovary, the testes are side-by-side, 

widely separated (by approximately 1 diameter), widely separated from the 

ventral sucker, and the uterus with eggs is confined in the field between the 

intestinal ceca and does not extend posterior of the vitellaria. 

Records - Two occurred in a black jack from Mona Island, 1 in 1 of 6 king 

mackerel from La Parguera, Puerto Rico. 



Length - 0.6-1.0 mm. 

Host Specificity - Apparently this fluke has little host preference as 1-10 

(average 3.4) flukes occurred in 12 fish specimens of 12 fish species in 9 

families and 6 orders of fishes. 

Lecithocladium excisum (Rudolphi) 

This small, but distinctive, stomach fluke is a dominant parasite of Atlantic 

mackerel. It may be more abundant in the cooler water areas, but occurs 

throughout the world in a variety of hosts. 

Name - Ectenurus sp. of Linton, 1910, L. excisiforme Cohn, L. gulosum 

(Linton), and Paradinurus manteri Perez-Vagueras, are synonyms, and L. 

harpodontis Srivastava is similar and may also be a synonym. 

Perez-Vagueras (1958) described a new genus and new species, P. manteri, 

from a single, 11.2 mm long, worm from the stomach of a Cuban great 

barracuda. His specimen was unusual in having a body that inserted into the 

tail, instead of a tail that inserted into the body. Otherwise, this worm is 

identical to L. excisum in size, shape and arrangement of organs. The identical


and distinctive oral sucker and egg sizes (20-22 x 10-12 Φm) in 

both of these forms make a particularly compelling argument that 

they are the same. The genus Paradinurus is a synonym of 

Lecithocladium. 

Perez-Vagueras (1958) and Yamaguti (1971), who accepted 

this new genus, were apparently not aware that Linton (1910) 

described and illustrated this worm as Ectenurus sp. Linton also 

noted and illustrated that the body inserted into the broader tail in 

2 worms, but a tail that was more narrow than the body in a third 

worm. Thus the main character used to distinguish genus 

Paradinurus actually varies within the species. 

Diagnostic Characters - The distinctive, elongate oral sucker is 

shaped like a scallop shell with a wide anterior and more narrow 

posterior. The oral sucker is larger than the ventral sucker and 

they are separated by approximately 1 width of the ventral sucker. 

This elongate fluke has a tail that is more than 1/3, but less than 

1/2 of the total body length. Weak striations occur on the sides 

of the body. The pharynx is tubular. The intestinal ceca extend 

to near the end of the tail. The winding vitellaria and the eggfilled 

uterus extend into the tail. 

Records - One occurred in a great barracuda from off Havana, Cuba; in chub 

mackerel from Brazil; 3 immature worms in a frigate tuna from the Dry 

Tortugas off Florida, USA; 2 in an Atlantic mackerel (USNPC 8377), and 1-17 

(average 6.3) in 20 chub mackerel (USNPC 8375-6) from Woods Hole 

Massachusetts, USA. It is found in Atlantic mackerel and chub mackerel 

throughout the cooler waters of the north and possibly in the south Atlantic; and 

immature worms in bluefin tuna off Europe. 


Life History - Immature forms of this fluke are found in a great variety of false 

hosts. 

Ecology.- This parasite may occur more abundantly in temperate and cooler 

waters. There are records from south Florida, the northern Gulf of Mexico, 

both sides of the Atlantic, Baltic, Mediterranean, Black Sea, Pacific Ocean, 

Japan and New Zealand, but only 1 from the tropics. 


Length - 6.0-15.0 mm. These worms are about half as long when the tail is 

retracted into the body. Immature worms are 1.0-5.9 mm. 

Host Specificity - Atlantic mackerel is the dominant host of this parasite in the 

Atlantic, but it also occurs in a variety of other fishes. It may be a charac- 

teristic parasite of Atlantic mackerel, and is at least a primary parasite. 

Parahemiurus merus (Linton) 

This tiny fluke has generalized host preferences, but has only infected big 

game fishes in Jamaica. 

Name - Hemiurus sardiniae Yamaguti is a synonym. 

37

38 


Diagnostic Characters - It is an elongate fluke with a 

short, deeply invaginated tail. The oral and ventral suckers 

are close together but separated by about 1 width of the 

ventral sucker. Deep striations (cuticular denticulations) 

occur on the sides from the oral sucker to 2/3 of the way 

down the body. The uterus with eggs either does not extend 

into the tail or just barely does, and that the intestinal ceca 

extend into the tail. Vitellaria are reduced to a rounded lump 

near the ovary. 

Records - One occurred in a blue runner, 2 in 1 of 3 

crevalle jack, 1 in 1 of 4 greater amberjack, and 4 in 3 of 18 

horse-eye jack from Jamaica. Despite extensive examinations 

of these same hosts in Puerto Rico, we have not seen 

this worm in any of these hosts. It did occur in light 

infections in other fishes from Puerto Rico (USNPC 39398). 

This worm was found in Indo-Pacific sailfish from Japan. 

We found 1 Sterrhurus sp. in a great barracuda from 

Okinawa, Japan (USNPC 79985), that could possibly be this 

worm. It is also found in a variety of others hosts from the 

Dry Tortugas and throughout the western Atlantic, the Gulf 

of Mexico, Brazil, Pacific northwest coast of USA, Ecuador, 

and Japan. 



Length - 1.4-3.5 mm. 

Host Specificity - This fluke appears to have little host 

preference as it occurs in a broad range of hosts. 

Sterrhurus musculus Looss 

This tiny fluke has a worldwide distribution and a broad 

range of hosts including crevalle jack. 

Name - Lecithochirium floridense (Manter); S. floridense M.; 

Separogermiductus zeloticus Travassos, T. de Freitas and 

Buhrnheim and Sterrhurus zeloticus (T., T. and B.) appear to be 

synonyms. 

Diagnostic Characters - It is a bullet-shaped fluke with the oral 

and ventral suckers less than 1 ventral sucker width apart. The 

oral sucker is approximately 1/3 the width of the ventral sucker. 

The tail is usually extended from the body, but can be contracted 

into the body, and is more than 1/3, but less than 1/2 the total 

body length (when extended). The compact and rounded lobe of 

vitellaria is posterior to the ovary. The testes are touching or 

slightly separated,side-by-side on a slight diagonal, just posterior 

to or slightly under the ventral sucker. The egg-filled uterus


does not extend to the end of the intestinal ceca or into the tail. The intestinal 

ceca either do not extend into the tail or just barely do so. 

Records - Found in 1 of 3 crevalle jack from Biscayne Bay, Florida, USA, and 

from Brazil; and in great barracuda in the Gulf of Mexico off Mexico. It 

occurred in a variety of fishes from Puerto Rico, Curaçao and Jamaica, but not 

in big game fishes. It is known throughout the western Atlantic, eastern Atlantic 

off Europe, Mediterranean, Black Sea, and Japan in the western Pacific. 



Length - 1.0-2.1 mm. 

Host Specificity - This fluke apparently has little host preference since it 

occurs in a great variety of fishes. Its rarity in great barracuda suggests that this 

is a false host. 

Tetrochetus coryphaenae Yamaguti 

This is a common and abundant parasite in dolphin 

around the world. 

Name - This worm was placed in a new genus, Paratetrochetus 

Hanson, because it has a distinctive structure 

around the pharynx, but this character was not sufficient to 

distinguish a genus. Tetrochetus aluterae Hanson is a 

synonym, and T. macrorchis Yamaguti may be a synonym. 

Diagnostic Characters - This is a minuscule to small, 

elongate fluke with the ventral sucker attached on a 

projection from the body. The oral sucker and ventral 

sucker are close together and of similar size. No tail is 

present. 

Records - We found 1-41 worms in 8 of 13 dolphin 

examined off La Parguera (USNPC 84885), in this host and 

wahoo from Puerto Real and Mona Island, Puerto Rico; 

and 5 in 1 of 5 dolphin from Dauphin Island, Alabama, 

USA. It has also been found in dolphin, filefishes and 

puffers from Curaçao, Netherlands Antilles; Bimini, 

Bahamas (4 in a dolphin); Jamaica; Mexico; and Louisiana, 

USA, in the Atlantic; and Panama and Japan in the Pacific. 


Ecology - It occurs in both offshore pelagic and inshore fishes, and thus must 

have a broad range of tolerances and intermediate hosts. 

Associations - A variety of other worms occurred in the stomach and intestine 

of dolphin with this host, but we found no apparent interaction between this 

fluke and other parasites in the 13 dolphin we analyzed for total parasites. 

Location in Host - Intestine. 

Length - 2.3-7.8 mm. 

39

40 


Host Specificity - This worm largely occurs in dolphin and filefishes. This is 

a peculiar pattern considering that these 2 families of fishes are not closely 

related. It is a primary, but not a characteristic parasite of dolphin. The single 

records in wahoo from Puerto Rico and a porcupinefish, Diodon hystrix 

Linnaeus, from Jamaica probably represent false or accidental hosts. 

Mabiarama prevesiculata Freitas and Kohn 

This peculiar worm was described from cobia 

in Brazil. 

Name - A new family, Mabiaramidae, and new 

genus was based on this unusual worm. 

Diagnostic Characters - This small, stout worm 

has an ventral sucker that is separated from the oral 

sucker by more than 1 diameter of the ventral 

sucker, and is approximately twice as large as the 

oral sucker. The oral sucker and cecal bifurcation 

attach almost directly to the pharynx. The ceca are 

irregular in outline and extend to the posterior end 

of the worm. The vitellaria are confined to the 

lateral margins of the posterior body. 

Records - This fluke occurred in cobia from Brazil. 



Length - 6.9-11.9 mm. 

Host Specificity - Unknown. 

Paradeontacylix sanguinicoloides McIntosh 

This is the only blood fluke known from big game fishes. 

Diagnostic Characters - It is a uniformly wide, elongate worm 

with a blunt anterior end and pointed posterior end. The ceca are 

X-shaped. The middle of the body is filled with 2 long irregular 

rows of testes. 

Records - One occurred in a greater amberjack off Miami, Florida, 

USA (USNPC 34329). 


Location in Host - Blood vessels of gills. 

Length - 3.2 mm. 


Damage to Host - Two similar species from Asia cause mass 

mortalities of greater amberjack in aquaculture in Japan.


Tergestia laticollis (Rudolphi) 

This highly ornamented small fluke has distinctive 

"dreadlocks". Its variability, worldwide range, and broad 

host preferences have caused much confusion. 

Name - Yamaguti (1970) suggested that the only difference 

between this fluke and T. pectinata (Linton) was the sizes 

of the eggs, yet he listed overlapping ranges of egg size for 

these 2 species (Yamaguti 1971). We also struggled to 

separate these species, but now we recognize 1 definable 

species rather than 2 that are so similar that they confuse the 

experts. Tergestia acuta Manter, T. pauca Teixeira de 

Freitas and Kohn, T. pectinata and T. selenei Amato are 

synonyms of this highly variable fluke. 

Diagnostic Characters - This minuscule to tiny fluke has 

relatively large, distinctive, tentacle-like papillae surrounding 

the oral sucker. Muscular folds are visible in the sides of 

the neck. The ventral sucker is relatively large and 

occupies more than 1/2 of the body width. 

Records - Ten occurred in 1 of 3 bar jack from off Cabo 

Rojo (USNPC 85463), 12 in 1 of 2 bar jack (USNPC 85940) 

and in little tunny from La Parguera, Puerto Rico (USNPC 39332). One was 

found in a blue runner, 1 each in 3 crevalle jack, 9 in 5 of 18 horse-eye jack 

and 1-3 in 4 of 8 yellow jack from Jamaica; 1 in 1 of 4 bar jack, 4 in 1 of 2 

blue runner, 1-6 in 3 of 4 horse-eye jack from Bimini, Bahamas; 2 in 1 of 4 

horse-eye jack from Curaçao; 7 in 1 of 8 yellow jack from Belize (USNPC 

74189); 2 in 1 bar jack from Bermuda; and in a little tunny in the Gulf of 

Mexico off Mexico. One occurred in 1 of 3 crevalle jack from southern Brazil 

and from Florida, USA; in 1 of 2 blue runner from south Florida, and Alligator 

Harbor and Tampa Bay, Florida; in a frigate tuna, 3 little tunny, skipjack tuna 

and 2 yellow jack from the Dry Tortugas, Florida; 6 in a bullet tuna from 

Woods Hole, Massachusetts, USA (USNPC 8190); and in chub mackerel and 

skipjack tuna from the Atlantic coast of the USA. Five were found in a 

rainbow runner from the Pacific coast of Costa Rica; and in chub mackerel 

from Japan. We found it in a variety of other fishes in Puerto Rico and 

Okinawa, and it has been reported from the Black Sea, North Sea, 

Mediterranean, eastern and western Pacific. 


Life History - We found 2 immature specimens encysted in the heart of a gray 

angelfish, Pomacanthus arcuatus (Linnaeus), from La Parguera, Puerto Rico. 

These flukes differed from this species by having longer, lanceolate muscular 

lobes around the oral sucker and a smaller ventral sucker which was in the 

posterior half of the body. Possibly these structures change as the worm 

develops since we have found no other species in this genus in Puerto Rico. 


41

42 


Length - 0.9-5.4 mm. The cysts in the heart of a gray angelfish were 218 µm, 

and the encysted worms were 188 µm. 

Host Specificity - This worm has no host preference and occurs in a wide 

variety of big game fishes and other fishes. 

Damage to Host - The cysts in the heart caused little tissue damage and no 

noticeable host response. 

Family Bucephalidae 

These tiny flukes are distinct because they do not have anterior mouths 

surrounded by an oral sucker. The "oral" sucker is replaced with a more 

elaborate "rhynchus" with folds, caps and tentacles. The name means big (bu) 

head (cephalo). The mouth is on the ventral side, usually somewhere in the 

middle third of the body. The intestinal ceca are reduced to little more than a 

small oval sack which can easily be mistaken for an ovary or testis in wet 

mounts. This family seems to occur most frequently in inshore fishes. Thus 

infections in the inshore wandering jacks, barracuda, and Spanish mackerels are 

understandable, but those in little tunas and bonitos are more difficult to explain. 

These worms are rather important in big game fishes, possibly disproportionately 

so. This may be related to some aspects of the ecology or behavior of 

near-shore big game fishes. The plasticity and variability of some of these 

worms has caused much confusion in the literature which we have probably not 

been entirely successful in resolving. Some flukes are host specific to big game 

fishes, but few occur with regularity, and none qualify as characteristic 

parasites. 

The rhynchus should be examined in live flukes when 

possible. The definitive shape and structures may change 

in dead or improperly preserved samples. These flukes are 

almost too small to attract the interest of sport fishermen 

but hold a wealth of information for the eco-parasitologist. 

Alcicornis carangis MacCallum 

This fluke may damage West Indian jacks. It has 

oddly shaped tentacles on the anterior end.. 

Name - Siddiqi and Cable (1960) correctly identified this 

fluke in Puerto Rico, but Nahhas and Cable (1964) 

described it as A. siddiqii based primarily on the larger and 

more developed rhynchus in larger worms. Very likely a 

bigger fluke has a larger, more developed rhynchus. We 

do not consider these differences sufficient to distinguish a 

new species when so little is known about this fluke. The 

rhynchus was redescribed in detail by Rees (1970). 

Diagnostic Characters - It is a minuscule, elongate worm 

with a distinctive wedge-shaped, relatively large, elongate


rhynchus with 7 tentacles. Each obvious tentacle has 2 side branches of 

different sizes. The mouth opens approximately in mid-body. 

Records - Four occurred in 1 of 2 bar jack from La Parguera (USNPC 85464- 

5), 10 in a bar jack from Puerto Real (USNPC 39302), 22 in a blue runner from 

La Parguera (USNPC 85941), and more than 100 in a horse-eye jack from 

Humacao, Puerto Rico (USNPC 82997). Three were found in a bar jack from 

Curaçao (USNPC 60249); in bar jack from Cuba; 1-2 in 2 of 8 yellow jack from 

Jamaica; 1-40 in 40 of 52 bar jack from Bermuda; and "moderate numbers" in 

a bar jack from the New York Aquarium that was collected from Key West, 

Florida, USA. 


Associations - See Ectenurus lepidus. 

Location in Host - Stomach or intestine (bar jack). We found 100 flukes in 

the pyloric ceca of a horse-eye jack. 

Length - 0.9-2.5 mm. 

Host Specificity - This parasite may be genus specific (Caranx), or family 

specific to jacks. The high numbers in horse-eye jack and lower ones in bar 

jack, could suggest that horse-eye jack might be the preferred host. This 

parasite appears to be a secondary parasite of both hosts. Horse-eye jack was 

a new host (Bunkley-Williams, Dyer and Williams 1996). 

Damage to Host - The very heavy infection of more than 100 worms in the 

pyloric ceca could easily stunt or injure the host. 

Significance to Sport Fishing - If it is restricted to the West 

Indies, this fluke might have some potential as a temporary 

biological tag. 

Bucephalopsis arcuata (Linton) 

This fluke may be a characteristic parasite of Atlantic bonito 

and important in other big game fishes. 

Name - This species was briefly placed in the genera Buceph- 

aloides and Prosorhynchoides. Bucephalopsis scomberomorus 

(Corkum) appears to be a synonym. 

Diagnostic Characters - This minuscule to tiny, oval fluke has 

a relatively small, simple rhynchus. Ear-like anterior projections 

of the body occur on either side of the rhynchus. The eggs do not 

occur in the anterior part of the body. The positions of the testes 

and ovary vary considerably. This worm is very similar to 

Bucephalopsis longicirrus, with which it has often been confused, 

but it varies by having an excretory bladder than extends anterior 

of the pharynx. 

The encysted form is usually yellow, but may be silver. 

Older cysts turn brown. 

Records - Two occurred in 1 of 2, and 6 in 1 king mackerel from 

La Parguera, Puerto Rico (USNM). One was found in a king 

mackerel from Jamaica and 1 in 1 of 2 of this host from Curaçao. 

43

44 


It also occurred in 2 cero from south Florida, USA; in 3 of 11 king mackerel 

and 11 of 33 Spanish mackerel from Louisiana, USA (USNPC 70983); 1100 in 

15, 500 in 15 (USNPC 8170), 36 in 3, 22 in 5, and 18 in 4 Atlantic bonito, and 

5 encysted in 1 of 12 Atlantic mackerel, from Massachusetts, USA; in Atlantic 

bonito, king mackerel, Spanish mackerel, crevalle jack, and a few other fishes 

from North Carolina, Florida and Texas, USA. Various reports of this fluke 

from great barracuda were misidentifications of Bucephalopsis longicirrus. 

Geographic Range - Western Atlantic (not confirmed from the Atlantic coast 

of South America). 

Life History - These flukes probably enter the liver through the bile duct and 

become enclosed in host generated, connective tissue cysts. 

Location in Host - Pyloric ceca, stomach and intestine; encysted in connective 

tissue cysts in the serous coat of the pyloric ceca and liver. 

Length - 1.3-3.1 mm; cyst 3.0 mm. 

Host Specificity - This fluke may be a characteristic parasite of Atlantic 

bonito, but reports and records are too incomplete to be certain. It also occurs 

in a variety of other fishes, including many big game fishes. 

Damage to Host - The many hundreds of encysted flukes, that have been 

reported in each liver of some Atlantic bonito, could injure this host. The very 

heavy infections of gut flukes, that have frequently been reported from this 

host, must also be injurious. 

Bucephalopsis longicirrus Nagaty 

This worm is a worldwide parasite of barracudas. 

Name - This fluke has been placed in the genera 

Bucephaloides and Prosorhynchoides. 

Diagnostic Characters - It is a minuscule, elongate 

fluke with a relatively small, simple rhynchus. Ear-like 

anterior projections of the body occur on either side of 

the rhynchus. The eggs are not found in the anterior 

part of the body. This worm is very similar to B. 

arcuata, with which it has often been confused, but 

varies by having an excretory bladder than does not 

extend anterior of the pharynx. 

Records - This worm occurred in great barracuda from 

Mona Island (USNPC 39306); in 5 of 11 from 

Louisiana, USA; 1 to more than 100 in 3 from Bimini, 

Bahamas; 1 in 1 of 6 from Curaçao; 3 in 2 of 3 from 

Jamaica; Atlantic coast of Panama; Dry Tortugas and 

south Florida, USA.It has also been reported in another 

species of barracuda from the Red Sea. 


Associations - It occurred with B. longovifera and Rhipidocotyle barracudae 

from the Dry Tortugas; and with B. longovifera in 4 fish from Louisiana. 

Location in Host - Pyloric ceca and intestine.


Length - 0.7-1.3 mm. 

Host Specificity - This parasite is genus specific to barracuda (Sphyraena). 

Bucephalopsis longovifera Manter 

Great barracuda commonly have this parasite 

throughout the West Indies. 

Name - The species name is for the unusually elongate 

egg ("longovi"=long egg, "fera"=bearing). This fluke 

has been placed in the genera Bucephaloides and 

Prosorhynchoides. 

Diagnostic Characters - This microscopic to minuscule, 

oval fluke has a relatively small simple rhynchus. 

Relatively large elongate eggs are distributed throughout 

the body. 

Records - One occurred in 1 of 3 great barracuda from 

Jamaica, 2 in 1 of 6 from Curaçao, in 8 of 15 from the 

Dry Tortugas, Florida, USA; and in 5 of 11 from 

Louisiana, USA. 

Geographic Range - West Indies and Gulf of Mexico. 

Life History - Manter (1940b) noted and figured the 

odd-shaped, and highly variably eggs of this worm. 

Associations - See Bucephalopsis longicirrus. 

Location in Host - Pyloric ceca and 

intestine. 

Length - 0.6-1.0 mm. 

Host Specificity - This worm only occurs in great barracuda. 

Bucephalus gorgon (Linton) 

This fluke of amberjack has numerous distinctive tentacles 

around the anterior end. 

Name - The name "gorgon" is a name for the Medusa, a 

mythological creature with snakes for hair that turned men to 

stone. The original specimens of Gasterostomum gorgon Linton 

were lost, the original description of the species was 

incomplete, and Linton often grouped multiple species under a 

single name, thus we cannot be certain what species his name 

actually represented. His renamed Nannoenterum gorgon 

(Linton 1940) (USNPC 8185) appears to be this fluke. 

Diagnostic Characters - It is a minuscule to tiny, elongate 

worm with a rhynchus appearing like a relatively large oral 

sucker with 22 moderate-sized tentacles on the anterior end. 

The tentacles on each worm vary in size. 

Records - This worm occurred in 7 of 8 greater amberjack and 

1 of 3 banded rudderfish, from Louisiana, USA; and in a 

greater amberjack from North Carolina, USA; and 9 in 1 and 

45

46 


89 in 3 greater amberjack from Woods Hole, Massachusetts, USA (USNPC 

8185). 

Geographic Range - Atlantic and Gulf coasts of USA. 

Location in Host - Anterior intestine. 

Length - 1.6-3.3 mm. 

Host Specificity - This parasite is genus specific (Seriola). It may be a 

characteristic parasite of greater amberjack. 

Bucephalus varicus Manter 

This worldwide parasite of jacks has some 

interesting distributional gaps. 

Name - The name is appropriate as this fluke 

appears to be highly variable. Bucephalus polymorphus 

of Nagaty, 1937, and Caballero, Bravo- 

Hollis and Grocott, 1953 and B. pseudovaricus 

Velasquez are synonyms, and B. solitarius Kohn 

appears to be a synonym. This fluke needs to be 

redefined and refigured. 

Diagnostic Characters - The rhynchus is 

relatively large with 6-7 relatively large tentacles 

on the anterior end. The tentacles are often not 

protruded, resulting in knob-like structures around 

the anterior sucker. Slight pressure on live 

specimens may cause the tentacles to protrude. 

Records - One occurred in 1 of 2 bar jack, 3 in 2 

of 3 crevalle jack, 2 in 1 of 4 greater amberjack, 

1-3 in 12 of 18 and 1 in 1 of 3 horse-eye 

jack, and 1 each in 2 of 8 yellow jack from 

Jamaica; 1 in 1 of 4 bar jack, more than 300 in 1 

of 2 blue runner, 12 to more than 300 in 4 horseeye 

jack from Bimini, Bahamas; 25 in a yellow 

jack from Grand Cayman, Cayman Islands (USNPC 82471); 3-75 in 4 horse-eye 

jack (USNPC 74241) and 8-35 in 8 yellow jack (USNPC 74240, 74282) from 

Belize. It was found in 5 of 6 bar jack, 1 of 6 horse-eye jack and 1 of 2 yellow 

jack from the Dry Tortugas, Florida, USA; in 1 of 2 blue runner, and 3 crevalle 

jack from south Florida; in blue runner and crevalle jack from Alligator Harbor, 

Florida; and in 1 bar jack, 2 of 6 blue runner, 6 of 20 crevalle jack and 1 horseeye 

jack from Louisiana, USA; and blue runner from Brazil. This worm 

possibly occurs in greater amberjack and in jacks and other fishes from the 

Pacific coast of Panama; Baja California, Mexico; the Red Sea; Okinawa, Japan; 

and the Philippines. 


Ecology - This fluke is displaced by Alcicornis carangis in Puerto Rico. 

Location in Host - Pyloric ceca, occasionally in the stomach or intestine. 

Length - 0.9-2.0 mm


Host Specificity - This parasite largely occurs in jacks, particularly those in 

the genus Caranx, but not exclusively enough to be family specific. It may be 

a secondary parasite of most of the jacks listed above. 

Rhipidocotyle baculum (Linton) 

This is a rather variable tiny fluke of western Atlantic Spanish 

mackerels that has, until recently, been called by 4 names. 

Name - This fluke and R. adbaculum Manter are very similar in 

size, shape and host preferences (the name "adbaculum" means 

similar to "baculum"). They are never reported together, although 

they occur in the same hosts in the same localities. The 

uncertainty about their identity has caused them to be measured 

and partially redescribed so many times that they can now be seen 

to be synonyms. Prosorhynchus stunkardi Siddiqi and Cable, 

described from an unidentified mackerel in Puerto Rico, is also a 

synonym (Bunkley-Williams, Dyer and Williams 1996). We also 

see little reason to distinguish the very similar R. quadriculata 

Kohn, described from serra Spanish mackerel in Brazil. 

Diagnostic Characters - This minuscule to tiny, elongate fluke 

has a muscular rhynchus covered with a pentagonal, beret-shaped, 

flat hood. The pharynx is smaller than the rhynchus and is located 

approximately in mid-body. 

Records - Four occurred in a cero from La Parguera, Puerto Rico 

(USNPC 81642); 1 in 1 of 2 king mackerel from Curaçao; and 8 in 1 of 2 cero 

from Belize (USNPC 74242). It has also been reported in 1 of 3 cero, 1 

Spanish mackerel and 2 of 3 king mackerel from the Dry Tortugas, Florida, 

USA; in 2 king mackerel and 1 of 2 Spanish mackerel from south Florida; and 

1 in 1 of 33 Spanish mackerel from Louisiana, USA; it is common in Spanish 

mackerels from South Carolina, USA; and occurs in Spanish mackerel from the 

northeast Gulf of Mexico and Massachusetts, USA; and serra Spanish mackerel 

from Brazil. 


Ecology - It infects both inshore and offshore species of Spanish mackerels. 

Location in Host - Pyloric ceca and intestine. 

Length - 0.7-4.5 mm. 

Host Specificity - This parasite is genus specific to western Atlantic Spanish 

mackerels (Scomberomorus). Serra Spanish mackerel is a new host. 

Rhipidocotyle barracudae Manter 

This tiny fluke, found in great barracuda, has an oddly restricted distribution 

in south Florida, Cuba and the Bahamas. It might be useful as a biological tag. 

Diagnostic Characters - It is a minuscule, elongate fluke with a muscular 

rhynchus without papillae but covered with a pentagonal, beret-shaped, flat-hood 

with 5 lobes. The pharynx is smaller than the rhynchus and located posterior 

to mid-body 

47

48 


Records - Three occurred in 1 of 7 great barracuda from Belize 

(USNPC 74243); 1 of 2 from Eleuthera and Nassau, Bahamas; 

from Cuba and the western Gulf of Mexico off Mexico; and 1-7 

in 7 of 8 and 2 of 15 from the Dry Tortugas, Florida, USA. 

There are no records from the northern Gulf of Mexico or the 

Caribbean. We have not seen it in Puerto Rico. 

Geographic Range - Northern West Indies and southern Gulf 

of Mexico. 

Associations - See Bucephalopsis longicirrus. Manter (1940b) 

found 1 fluke infected with "very minute microorganisms filling 

the parenchyma in the pharynx region." 

Location in Host - Intestine and pyloric ceca. 

Length - 1.2-1.8 mm. 

Host Specificity - Only occurs in great barracuda. 

Significance to Sport Fishing - If the distribution of this 

parasite is as restricted as our records suggest, then it might be 

useful as a biological tag. 

Rhipidocotyle capitata (Linton) 

This fluke occurs in little tunas around the world. 

Name - Linton (1940) described Gasterostomum capitata 

before Manter (1940b) described R. nagatyi. Both names 

are synonyms. Rhipidocotyle angusticollis Chandler described 

in Atlantic bonito from Texas, USA (USNPC 36786) 

appears to be a synonym. 

Diagnostic Characters - This minuscule, elongate fluke has 

a muscular, bowl-shaped rhynchus covered with a pentagonal, 

flat-hood. The pharynx is smaller than the rhynchus and is 

located posterior to mid-body. 

Records - This worm occurred in little tunny from Puerto 

Rico (USNPC 39301). Nine were found in 3 little tunny 

from the Dry Tortugas off Florida, USA; 1 of 6 little tunny 

and 2 of 11 king mackerel from Louisiana, USA; in an 

Atlantic bonito and numerous little tunny from Texas, USA; 

17 in a bullet tuna (USNPC 8172) and little tunny (USNPC 

36707) from Massachusetts, USA; and 3 adults and a larva in 

a frigate tuna from Hawaii, USA. 


Life History - A fusiform larva was figured by Yamaguti (1970). 


Length - 1.1-2.2 mm; larvae 0.9 mm. The flukes in Pacific frigate tuna were 

smaller (1.3-1.5 mm) than those from Atlantic bullet tuna (1.1-2.2 mm). 

Host Specificity - This worm could be almost tribe specific to little tunas. 

King mackerel and Atlantic bonito may be false hosts.


Coitocaecum extremum (Travassos, Frieitas and Bührnheim) 

This fluke was found in chub mackerel from Brazil. 

Name - Nicolla extrema Travassos, Frieitas and 

Bührnheim is a synonym. Thatcher (1993) returned it to 

genus Nicolla, but did not refute the transfer to 

Coitocaecum by Yamaguti (1971). 

Diagnostic Characters - It is a minuscule, oval worm 

with a rather small ventral sucker that is larger than the 

oral sucker and located anterior to mid-body. It appears 

rather dark in color because the vitellaria extend throughout 

much of the body. The relatively large eggs are confined 

to mid-body. The testes are relatively large. The intestinal 

ceca join posteriorly forming a loop. 

Records - Reported in chub mackerel from Brazil. 

Geographic Range - Unknown 

Location in Host - intestine. 


Host Specificity - It was reported in 2 other species of 


Neolepidapedon retrusum (Linton) 

This fluke has only been found in chub mackerel 

Name - This worm has been placed in the genus Lepocreadium. 

Acanthocolpoides pauloi Travassos, Freitas and Bührnheim appears 

to be a synonym. 

Diagnostic Characters - This is a minuscule, elongate fluke with 

a relatively small ventral sucker, which is approximately the same 

size as the oral sucker, and is in the anterior 1/3 of the body. The 

vitellaria surround the intestinal ceca and extend from the posterior 

end to near the ventral sucker. The rhomboidal testes are in the 

posterior 1/3 of the body. 

Records - One to 4 occurred in 2 chub mackerel from Woods 

Hole, Massachusetts, USA (USNPC 8273-4); and in a chub mac- 

kerel from Brazil. 



Length - 1.1-3.2 mm. 

Host Specificity - This worm is only known from chub 

mackerel. 

49

50 


Genus Stephanostomum Looss 

Members of this genus possess distinctive spines in 2-3 uninterrupted rows 

encircling the mouth (circumoral spines). The variations on numbers and 

configurations of these spines would seem to provide ample diversity for easy 

species identifications, unfortunately, there is much variation in these spines 

within species. Manter (1947) discovered clusters of smaller, possibly regenerating, 

spines that appeared to be replacing missing and abnormal spines. Such 

losses and replacement with increased numbers of spines could contribute to 

some of the confusion about the number of spines in each species. The numbers 

of spines reported for the following species is the total count of all rows. 

The life cycles of a few species have been deciphered and can be used as 

possible models for the remaining species. Redia, located in the digestive gland 

of a mollusk, produce cercariae which escape and infect fishes where they 

encyst in tissues as metacercariae. We have found metacercariae of these flukes 

encysted in coral reef fishes from Puerto Rico (USNPC 82967). 

Stephanostomum coryphaenae Manter 

This small fluke is found in dolphin from the warmerwater 

portion of the western Atlantic. It might be of use as 

a biological tag. 

Diagnostic Characters - It is a small, stocky fluke with 36 

circumoral spines. The oral and ventral suckers are 

separated by more than 3 diameters of the ventral sucker and 

are approximately the same size. The vitellaria do not extend 

anteriorly to the ventral sucker. The pharynx is not near or 

in contact with the bifurcation of the intestinal ceca. 

Records - We found 2-13 worms in 4 of 13 dolphin from off 

La Parguera, Puerto Rico (USNPC 84886); and in dolphin 

from the Mona Passage (USNPC 39339). It has also been 

reported from this host from Bimini (1 in 1), Curaçao (1 in 

1 of 3), Brazil and the Dry Tortugas off Florida, USA. 

Geographic Range - West Indies and Atlantic coast of 

South America to Brazil. 

Ecology - This worm appears to be restricted to the warmerwater 

regions of the western Atlantic. What ecological 

mechanisms are involved in maintaining this distribution 

would be fascinating to explore. 

We found this fluke in a porkfish, Anisotremus virginicus (Linnaeus), from 

a shallow, inshore seagrass bed habitat adjacent to Isla Magueyes, La Parguera, 

Puerto Rico (USNPC 77726). It has previously been found only in offshore, 

pelagic habitats. 

Associations - This fluke occurred with a variety of other worms in dolphin from 

Puerto Rico, but did not appear to be interacting either negatively or 

positively with any other parasite.


Location in Host - Stomach, intestine and rectum. 

Length - 2.0-5.1 mm. 

Host Specificity - This worm only occurs in dolphin, but does not occur 

consistently enough to be a characteristic parasite of this host. The 3 flukes we 

found in a porkfish were probably an accidental infection. 

Significance to Sport Fishing - If the apparently restricted distribution of this 

fluke is correct, it might be of some use as a temporary biological tag in 

identifying dolphin that move north into the Gulf of Mexico or up the Atlantic 

coast of the USA. 

Stephanostomum ditrematis (Yamaguti) 

This elongate fluke parasitizes jacks and rarely barracuda 

around the world. It is so variable that many synonyms have been 

named. Name - It was originally described from a surf perch in 

Japan. Subsequently, it was found in jacks all over the world. This 

worm could represent a complex of 2 or more similar species. 

Stephanostomum cubanum Perez-Vigueras, S. filiforme Linton, S. 

longisomum Manter, S. manteri Perez-Vigueras, and S. microcephalum 

Perez-Vigueras are synonyms of this fluke. Stephanostomum 

ghanense of Fischthal, 1977 (USNPC 74281) appears to be 

this worm. 

This worm has been found in 3 great barracudas and described as 

a different species each time. Linton (1910) described it as Distomum 

sp. from Bermuda; Perez-Vigueras (1942) as Monorchistephanostomum 

gracile from Cuba; and Yamaguti (1970) as S. kawalea from 

Hawaii. The new genus, Monorchistephanostomum Perez-Vigueras, 

was erected because the single worm available had only 1 testis 

(monorchism). Parasitic worms that normally have 2 testes are 

sometimes found with a single testis (Williams 1976). Overstreet 

(1969) noted a specimen of the similar Stephanostomum sentum with 

a single testis. Yamaguti (1971) synonymized this genus with 

Stephanostomum. 

Diagnostic Characters - This small, elongate fluke usually has 36 

(rarely 32-50) circumoral spines. The oral and ventral suckers are 

separated by approximately 3-4 diameters of the ventral sucker, and 

the oral sucker is smaller than the ventral sucker. The vitellaria do 

not extend anterior to the ventral sucker. Eggs are relatively large 

and confined between the intestinal ceca. The testes are separated by 

approximately 1-3 of their lengths. 

Records - We found 8 in a crevalle jack from Ponce, Puerto Rico 

(USNPC 83023). It has also been reported in a bar jack and rainbow 

runner from Cuba; 1 in a blue runner, 2 in 1 of 3 crevalle jack, 4 in 3 of 18 

horse-eye jack and 3 in 2 of 8 yellow jack from Jamaica; in an unidentified jack 

from Curaçao; 2-5 in 3 of 4 horse-eye jack from Bimini; 2-3 in 2 of 4 horse-eye 

jack (USNPC 74208) and 1-6 in 8 yellow jack from Belize (USNPC 74207, 

51

52 


74281, 74283); in 1 of 6 bar jack and 1 of 6 horse-eye jack from the Dry 

Tortugas off Florida, USA; and in 1 of 2 blue runner and 2 of 3 crevalle jack 

from Biscayne Bay, Florida; 1 in 1 of 15 blue runner (USNPC 8204), 1 in a 

crevalle jack (USNPC 8203), and 5 in 1, and 28 in 3 greater amberjack 

(USNPC 8202) from Woods Hole, Massachusetts, USA. It has also been 

reported in rainbow runner and other jacks from the Pacific. 

One occurred in a great barracuda from Cuba; 3 in 1 from Bermuda; and 

2 in another species of barracuda from Hawaii, USA (USNPC 63734). 


Ecology - It appears to occur rarely in Puerto Rico. We have examined more 

than 100 specimens of jacks, but have only seen this fluke once. We examined 

45 great barracuda in Puerto Rico and 2 from Alabama, USA, but did not find 

this rather distinctive and obvious worm. 


Length - 3.2-19.0 mm (3.2-15.0 mm in jacks; 12.5-19.0 mm in barracudas). 

Host Specificity - This worm is almost family specific to jacks. It does not 

occur consistently enough to be a primary parasite of any host. The low 

numbers that rarely occur in barracudas suggest that these are false hosts that 

temporarily obtain these worms by eating jacks. The greater length of these 

worms in barracuda may only be an artifact of how seldom they have been 

measured in any fish. 

Stephanostomum imparispine (Linton) 

It occurs in cobia from the Atlantic coasts of the USA. 

Name - Distomum valdeinflatum Stossich of Linton is a synonym. 

Diagnostic Characters - This tiny to small, elongate fluke has 

33-34 circumoral spines. The oral and ventral suckers are 

separated by less than 3 diameters of the ventral sucker, and are 

approximately the same size. The vitellaria do not extend anterior 

to the ventral sucker. The pharynx is in contact with the intestinal 

ceca bifurcation. The ovary and testes are in contact. 

Metacercariae are in globular cysts often containing yellowish 

to greenish fluid. Dark pigment sometimes coats the cysts. 

Records - Five occurred in a cobia from off Madeira Beach, 

Florida; off St. John's Pass, Florida, USA; and an unstated 

number of adults in 2 of 3, and 1 metacercaria in 1 of 6 cobia 

from Beaufort, North Carolina, USA. 

Geographic Range - Atlantic and Gulf coasts of USA. 

Life History - Its metacercariae encyst in a variety of 

intermediate fish hosts. 

Location in Host - Intestine. Metacercariae encyst in 

mesenteries, liver and other internal organs. 

Length - 4.5-10.0 mm; metacercaria 0.6-2.4 mm. 

Host Specificity - The adult worm only occurs in cobia and is 

probably a secondary parasite of this host.


Stephanostomum megacephalum Manter 

This fluke occurs in jacks from warm waters 

throughout the New World. 

Name - The name "megacephala" refers to the large 

oral sucker. Stephanostomum belizense Fischthal is a 

synonym. 

Diagnostic Characters - It is a tiny, elongate fluke 

with 24-32 circumoral spines with a slight gap in the 

2 rows. The oral and ventral suckers are separated by 

more than 3 diameters of the ventral sucker. The oral 

sucker is larger than the ventral sucker, and the ventral 

sucker is separated from the body by a projection of 

the body. The vitellaria do not extend anterior to the 

ventral sucker. The pharynx is elongate and near to 

the bifurcation of the intestinal ceca. 

Records - Four occurred in 1 of 18 horse-eye jack from 

Jamaica;1-10 in 5 of 8 yellowjack from Belize (USNPC 

74282-3, 74163); 1 in 1 of 6 horse-eye jack from the 

Dry Tortugas off Florida, USA; and in 2 of 3 crevalle 

jack from Biscayne Bay, Florida. It was also found in 

other jacks from the northern Gulf of Mexico; tropical west Africa and the 

eastern Pacific. 

Geographic Range - Atlantic and eastern Pacific tropics and subtropics. 

Associations - One fluke occurred with numerous S. ditrematis in a horse-eye 

jack from the Dry Tortugas; and 1 of each species occurred in a Pacific jack 

from White Friars, Mexico and San Francisco, Ecuador. 


Length - 1.1-3.3 mm. 

Host Specificity - This worm is family specific to jacks. 

Only 1 unconfirmed record exists of a host other than a jack. 

Stephanostomum sentum (Linton) 

This fluke prefers inshore, benthic hosts, but parasitizes 

big game jacks in Puerto Rico and Jamaica. 

Diagnostic Characters - This minuscule to tiny, elongate 

fluke has 36 circumoral spines. The oral and ventral suckers 

are separated by more than 3 diameters of the ventral sucker, 

and the oral sucker is smaller than the ventral sucker. The 

vitellaria do not extend anterior to the ventral sucker. The 

pharynx is near the bifurcation of the intestinal ceca. 

Records - We found 1 in a crevalle jack (USNPC 81579) and 

12 in a horse-eye jack (USNPC 83014) from La Parguera, 

Puerto Rico. One was found in 1 of 18 horse-eye jack from 

Jamaica. 


53

54 



Length - 1.9-3.8 mm. 

Host Specificity - This fluke is found in a variety of fishes. It appears 

to occur more often in grunts, porgies and mojarras and is only a 

secondary parasite of jacks. 

Tormopsolus filiformis Sogandares-Bernal and Hutton 

This worm occurred in cobia from the Gulf coast of Florida. 

Name - The name means hair-like (filum=hair). 

Diagnostic Characters - This long slender, tiny worm has relatively 

tiny and widely spaced oral and ventral suckers. 

Records - Five occurred in a cobia from St. John's Pass, Florida, USA 

(USNPC 39003). 

Geographic Range - Unknown 

Location in Host - Rectum. 

Length - 4.8-5.7 mm. 

Host Specificity - Only reported from cobia. 

Tormopsolus orientalis Yamaguti 

This worm parasitizes amberjacks around the world. 

Name - Tormopsolus hawaiiensis Yamaguti is a synonym. 

Diagnostic Characters - This elongate, tiny to small worm 

has an oral sucker that is close (less than 1 ventral sucker 

width) from the ventral sucker and about 1/2 the size of the 

ventral sucker. The testes are elongate oval-shaped, not 

touching, and are in the posterior part of the body. The 

vitellaria do not extend to the ventral sucker. 

Records - One occurred in a greater amberjack from 

Curaçao; and 2 in 1 and 1 each in 2 of 5 lesser amberjack 

from Bermuda. Also reported from another species of 

amberjack in the eastern Pacific; an unknown amberjack, 

Seriola sp., and greater amberjack in Hawaii; and greater 

amberjack and yellowtail in Japan. Silas (1962) suggested 

that "bonito" or Atlantic bonito had erroneously been 

reported as a host of this worm from Bermuda, but this 

record refers to the common name "bonito" that Linton 

(1907) used for lesser amberjack. 


Location in Host - Pyloric ceca and intestine. 

Length - 4.3-7.5 mm. 

Host Specificity - This parasite is genus specific to 

amberjacks (Seriola).


Pseudopecoeloides carangis (Yamaguti) 

This is an oddly shaped stomach worm of jacks around 

the world. 

Diagnostic Characters - This slender fluke has an ventral 

sucker on a distinct peduncle or projection from the body. 

The peduncle is close to the anterior end of the body. The 

oral sucker is relatively large and larger than the ventral 

sucker. 

Records - One occurred in a bar jack and 1 in a blue runner 

from Curaçao; 1 in 1 of 2 bar jack from Jamaica and in 1 of 

6 bar jack and 1 of 3 yellow jack from the Dry Tortugas off 

Florida, USA. It has also been reported from various jacks 

in the Pacific. 



Length - 1.2-5.0 mm. 

Host Specificity - This parasite is family specific to jacks. 

A record from a barracuda in the Philippines could represent 

a false host. 

Cetiotrema carangis (MacCallum) 

This is an obscure urinary tract parasite of jacks. 

Name - MacCallum (1913) described Distomum 

carangis and Yamaguti (1971) changed it to Gorgoderina 

carangis, thus placing it in a genus of frog flukes. 

Manter (1947) described Phyllodistomum carangis and 

Manter (1970) changed it to Cetiotrema carangis. These 

2 forms are synonyms, but had not been compared 

previously. It is not the same as Pseudopecoeloides 

carangis (Yamaguti). 

Diagnostic Characters - This small worm has thin 

lateral margins that often fold over onto its body. The 

oral and ventral suckers are approximately the same size 

and are separated by more than 3 diameters of the 

ventral sucker. The eggs are relatively large. 

Records - One occurred in 1 of 6 bar jack from the Dry 

Tortugas off Florida; and 4 in a blue runner from 

Woods Hole, Massachusetts, USA. 

Geographic Range - Atlantic coast of the USA. 

Location in Host - Urinary bladder or ureter. 

MacCallum (1913) reported this worm from the rectum, but this location is 

adjacent to the urinary bladder and contamination could have occurred in the 

dissection. 

Length - 8.1-13.0 mm. 

55

56 


Host Specificity - Possibly genus specific to jacks (Caranx). Yamaguti (1971) 

incorrectly stated that MacCallum's host was "Carangis". 

Detection - Manter's specimen was washed from the body cavity of the host, 

but presumably came from the urinary bladder. This organ is seldom examined. 

It is often difficult to locate when damaged during the dissection process or not 

inflated. 

Miscellaneous Flukes 

Aponurus laguncula Loss - This worm was found in a chub mackerel from 

southern Brazil. It has a ventral sucker about twice as wide as the oral sucker. 

The suckers are separated by about 1 width of the ventral sucker and both are 

elevated from the body by short peduncles. The vitellaria are in 7 distinct lobes 

posterior of the ovary. This parasite also occurs in the stomach of a variety of 

marine fishes from Florida and Louisiana, USA; Brazil; and the Mediterranean. 

It is 0.5-1.3 mm long. A single record in this rather well examined host sug- 

gests that this parasite may have been an accidental infection or in a false host. 

Brachyphallus crenatus (Rudolphi) - Atlantic mackerel is a false or 

accidental host for this parasite. It is a minuscule to tiny, elongate fluke with 

a short tail that is deeply embedded in the body. The width of the tail is approximately 

1/2 of the body width. The oral and ventral suckers are approximately 

the same size and are separated by 1-2 diameters. The body is covered with 

conspicuous cuticular serrations. The 2 testes are just posterior to the ventral 

sucker, round and arranged diagonally. The oval ovary is in the posterior 1/2 

of the body. The vitellaria are formed into 2 roughly kidney-shaped masses 

indented into 2 and 3 partial lobes, and are posterior to the ovary. The uterus 

fills the body from just posterior to the vitellaria to just posterior to the testes, 

and then 1 tube passes anterior almost to the pharynx. The intestinal ceca 

extend almost to the posterior end of the tail, but the uterus does not extend into 

the hindbody or tail. One fluke occurred in an Atlantic mackerel from Woods 

Hole, Massachusetts, USA (USNPC 8348). This parasite occurs in a great 

variety of fishes worldwide, but has been noted in no other big game fishes. It 

occurs in the stomach and intestine of host fishes and is 0.9-4.0 mm long. 

Immature worms in the alimentary tract of fishes are 0.2-0.8 mm long It is 

habitat specific to the near-shore environment. 

Bucephalopsis attenuata (Siddiqi and Cable) - Great barracuda is 

a false host for this fluke. This species was placed in genus Prosorhynchoides. 

It is a minuscule, elongate fluke with a relatively large, oval rhynchus. The 

eggs are relatively large. The vitellaria are arranged in 2 lateral rows in the 

anterior part of the body. Five flukes occurred in 1 of 10 great barracuda from 

La Parguera, Puerto Rico. This parasite has only been found in Puerto Rico. 

It occurs in the intestine of the host and is 0.6-1.1 mm long. Great barracuda is 

probably a false host which obtained this worm by eating the normal host, 

Atlantic bumper, Chloroscombrus chrysurus (Linnaeus). Great barracuda was


a new host record for this fluke, although this was not noted by Dyer, Williams 

and Bunkley-Williams (1985). 

Bucephalopsis gracilescens (Rudolphi) - The identity of this worm is 

not clear as this species name may have been used for several different species 

in the western Atlantic. This fluke was placed in the genera Bucephaloides and 

Prosorhynchoides. It is a minuscule, oval worm with a relatively large, oval 

rhynchus. The relatively small and numerous eggs are distributed throughout the 

body. The vitellaria occur in 2 bunches on either side of the rhynchus. One 

immature worm occurred in 1 of 13 crevalle jack from North Carolina, USA. It 

has also been reported in frigate tuna from the Mediterranean. It is 0.5-1.7 

mm long. Crevalle jack was probably a false host. 

Bucephalus confusus Velasquez - This is a confusing and common 

parasite of many species of fishes that may only accidentally infect Spanish 

mackerel. Nannoenterum baculum Linton is a synonym, but Rhipidocotyle 

baculum (Linton) is not the same species. The rhynchus of this worm has a 

relatively large sucker with 20 relatively short tentacles on the anterior end. 

The rather large, circular vitellaria are in 2 lateral rows of 12-18 follicles each 

in the middle 1/3 of the body. The 2 testes are in the posterior 1/3 of the body. 

The uterus extends from the testes to 1/2 the distance between the anterior end 

of the vitellaria and the rhynchus. One occurred in a Spanish mackerel and 

many other fishes from Woods Hole, Massachusetts, USA, and has not been 

found elsewhere. This worm occurs in the intestine of the host, and is 1.6-4.3 

mm long. This worm may be an accidental parasite in Spanish mackerel; or 

Spanish mackerel may be a false host. The preferred or dominant host appears 

to be northern sennet, Sphyraena borealis DeKay, with many infections of up 

to 460 per host (USNPC 8180). 

Claribulla longula Overstreet - Great barracuda is a false host for this 

fluke. It is a minuscule, elongate worm with a relatively large and distinctive 

funnel-shaped or cup-shaped oral sucker. The ventral sucker is widely separated 

from the oral sucker, a little anterior of mid-body and much smaller than the 

oral sucker. Overstreet (1969) suggested that the single specimen he found in 

a great barracuda from Biscayne Bay, Florida, may represent an accidental 

infection. It is only known from south Florida. This worm is usually found in 

the pyloric ceca, but a few occur in the upper intestine. It is 0.9-2.7 mm long. 

This worm is host specific to bonefish, Albula vulpes (Linnaeus). Great 

barracuda was a false host resulting from this predator eating a bonefish. 

Genolopa brevicaecum (Manter) - This grunt fluke is probably only 

found in big game fishes that are false hosts. It has also been placed in genus 

Paraproctotrema. This elongate to oval worm has a relatively large, cup-shaped 

oral sucker. The oral sucker is approximately twice as wide as the ventral 

sucker. The genital opening, anterior to the ventral sucker, is not armed with 

57

58 


spines. The 2 compact bunches of vitellaria are lateral, in approximately the 

middle of the worm, and the single testis occurs laterally just posterior to one 

of the bunches of vitellaria. The uterus occupies most of the hind body. Two 

occurred in 1 of 8 (Nahhas and Carlson 1994) and 17 in 2 yellow jack (Nahhas 

and Cable 1964) from Jamaica; and in 1 of 3 yellow jack from the Dry Tortugas 

off Florida, USA. It is distributed throughout the New World 

tropics/subtropics. This worm occurs in the intestine of the host and is 0.6-0.9 

mm long. It is probably family specific to grunts (Haemulidae). The 2 records 

in 2 or 3 yellow jack may be false hosts due to predation on grunts. 

Lasiotocus truncatus (Linton) - This grunt fluke is probably only found 

in big game fishes that are false hosts. It has also been placed in genus 

Proctotrema. This oval worm has a relatively large, cone or funnel-shaped oral 

sucker. The oral sucker is approximately 4 times as wide as the ventral sucker. 

The genital opening anterior to the ventral sucker is armed with spines. The 2 

compact bunches of vitellaria are found laterally and approximately at the middle 

of the length of the worm, and the single testis occurs in the middle of the body 

just posterior to the vitellaria. The uterus occupies most of the hind-body. This 

fluke occurred once in a bar jack from Bimini, Bahamas. It is only known from 

Bimini and the Dry Tortugas off Florida, USA. This worm occurs in the 

pyloric ceca of the host and is 0.7-1.0 mm long. This parasite is probably 

family specific to grunts (Haemulidae). The single record in a bar jack may be 

a false host due to this predator eating grunts. 

Lecithaster confusus Odhner - Atlantic mackerel is a false host for this 

parasite. It is an elongate, tear-drop shaped, minuscule fluke with no tail and 

relatively large suckers. The oral sucker is about 1/2 the size of the ventral 

sucker. The ventral sucker is about 1/2 as wide as the greatest width of the 

body. The 2 testes are side-by-side and posterior to the ventral sucker. The 

ovary has 4 obvious lobes and is median posterior to the testes. The vitellaria 

has 7 lobes and is in the middle, posterior to the ovary. It occupies less than 

1/3 of the width of the posterior body. The uterus fills the hindbody posterior 

to the ventral sucker. Lecithaster gibbosus (Rudolphi) has also been found in 

Atlantic mackerel but off Europe in the eastern Atlantic, in bluefin tuna from the 

Mediterranean, chub mackerel in the Pacific, and in a variety of other fishes 

worldwide. It occurs on the Atlantic coast of the USA, but has not been 

reported from western Atlantic big game fishes. It differs from L. confusus by 

having a short tail, a larger asterisk-shaped, 7-lobed vitellaria that fills the entire 

hindbody and covers the ceca. The ventral sucker is smaller and is 

approximately 1/3 of the greatest body width of the worm. Eight and 10 L. 

confusus occurred in 2 Atlantic mackerel from Woods Hole, Massachusetts, 

USA (USNPC 8361). It is found in the northern Atlantic and Mediterranean. 

The first intermediate host is a marine snail and the second is a copepod. The 

metacercaria in copepods develop into adult flukes in the intestine of fishes that 

eat infected copepods. Adult flukes are 0.5-1.5 mm long, and immature worms


in the fish intestine vary from 0.2-0.6 mm long. This parasite is family specific 

to herrings (Clupeidae). The record from an Atlantic mackerel was a false host. 

Lecithochirium monticellii (Linton) - We found 1-20 of this fluke in 6 

species of fishes in 5 families and 4 orders from Puerto Rico, but none occurred 

in big game fishes. If the egg sizes in this family are as narrow and precise as 

many experts claim, then this species with eggs 18-25 X 11-14 Φm, must 

represent 2 or 3 species of flukes. Manter (1947) suggested that the original 

description of this worm probably included at least 3 species: (1) L. branchialis 

(Stunkard and Nigrelli), (2) L. texanum, and what we are now calling L. 

monticellii; and Overstreet (1969) added L. microstomum to this species-complex 

list, but suggested that L. branchialis may be the same as L. microstomum. 

Thus we cannot be certain which species of parasite this name indicated in the 

early records. It is a small, bullet-shaped fluke with the oral sucker and ventral 

sucker moderately (approximately 1 ventral sucker diameter) separated, and a 

short tail that is usually withdrawn into the body. The oral sucker is less than 

1/3 the diameter of the ventral sucker. The vitellaria are reduced to 2 compact, 

oval masses just posterior of the ovary, the testes are side-by-side, separated by 

approximately 1/2 a testis diameter and separated from the ventral sucker by 1 

diameter, the uterus with eggs does not extend to the end of the intestinal ceca 

or into the tail, and ceca either extend into the tail or do not. One fluke each 

may have occurred in 2 blue runner (USNPC 8350), 9 in a little tunny (USNPC 

8353) from Woods Hole, Massachusetts, USA; and 2-3 in 2 of 6 cobia from 

Beaufort, North Carolina, USA. It also occurs in a variety of other fishes. It 

is found in the intestine of the host. The geographic range of this parasite is the 

western Atlantic. It is 1.0-5.4 mm long. Atlantic cutlassfish is the preferred 

host of this parasite because they are consistently more heavily infected (up to 

240 flukes). 

Lecithochirium priacanthi Yamaguti - In a checklist, Yamaguti (1970) 

lists this fluke as a parasite of rainbow runner from Hawaii, but does not 

mention it in the text. 

Myosaccium opisthonema (Siddiqi and Cable) - This fluke was 

described from a herring in Puerto Rico (USNPC 39393). It was originally 

placed in Neogenolinea. It is elongate with an oral sucker about 1/2 the size of 

the ventral sucker. The 2 are separated by approximately 1 diameter of the 

ventral sucker. Striations on the body are obvious, the eggs are relatively large 

and confined posterior of the ventral sucker. Vitellaria are confined in 2 

compact lobes just posterior of mid-body. We found 7 worms in a cero from 

Humacao, Puerto Rico (USNPC 83002), but this was probably a false or 

accidental host. Parasites in this genus are host specific or at least family 

specific to herrings and sardines (Clupeidae). Similar worms have been found 

in Florida and Japan, but this species is only known from Puerto Rico. It 

59

60 


occurs in the stomach of herrings, but we found it in the intestine of the cero. 

This worm is 0.5-0.8 mm long. 

Neolepidapedon belizense Fischthal - This species was described from 

1 immature and one adult in 1 of 7 great barracuda from Belize (USNPC 

74164), but this host is probably false or accidental. It is an elongate, tiny 

worm with a relatively large oral sucker, approximately the same size as the 

ventral sucker. The 2 suckers are separated by approximately 2 of their 

diameters. The first 1/3 of the body is covered with minute spines and contains 

both suckers. The oval testes are in the posterior 1/2 of the body and the ovary 

is anterior of the testes. The vitellaria extend from the posterior end to near the 

ventral sucker. It was 3.6 mm long and occurred n the pyloric ceca of the host. 

Opechona orientalis (Layman) - This worm may indicate the migration 

of chub mackerel into the Atlantic. It was described from angelfish taken from 

the bilge tanks of a freighter - an excellent way to transport exotic fishes and 

parasites! This oval worm has a ventral sucker slightly larger than the oral 

sucker and separated from it by more than 3 of its widths. Both suckers are 

relatively small, but the eggs extending from the ventral sucker to the ovary are 

rather large. The vitellaria extend from near the ventral sucker to the posterior 

end, and the ceca also reach the posterior end. The testes are in the posterior 

1/3 of the body, in tandem, and round to irregular rhomboidal in shape. This 

parasite occurs throughout the Pacific in various fishes but prefers chub 

mackerel as a host. A collection of this parasite in chub mackerel from southern 

Brazil and its apparent absence in the rest of the Atlantic could indicate that it 

was brought into the Atlantic by chub mackerel migrating from the Pacific. 

Yamaguti (1971) illustrated the more contracted specimens of Manter (1940a) 

from the Galapagos and Mexico, which were 0.7-1.0 mm long, but cited the 

lengths of more extended specimens (1.7-3.3 mm) from other localities. 

Opecoeloides brachyteleus Manter - We found 1 worm in a bar jack 

from Aguadilla, Puerto Rico, but this appears to have been a false host. It is 

family specific to goat fishes (Mullidae) throughout the Caribbean and south 

Florida. It has a ventral sucker with 4 papillae, separated from the body on a 

peduncle. The oral sucker and pharynx are similar in size and the ventral suc- 

ker is slightly larger. The vitellaria and 2 rather large testes are confined in the 

hindbody. The uterus has rather large eggs and is confined between the testes 

and pharynx. It occurs in the intestine of its hosts, and is 1.0-2.3 mm long. 

Opecoeloides vitellosus (Linton) - There is some uncertainty if this fluke 

was correctly reported by Linton (1901) from Atlantic bonito and Atlantic 

mackerel because these early records appear to have called several species of 

flukes by this name. This parasite has never subsequently been reported from 

these hosts. This slender fluke has an ventral sucker on a distinct peduncle or 

projection from the body. The peduncle is close to the anterior end of the body.


The oral sucker is smaller than the ventral sucker. The prepharynx is short, but 

the esophagus is long. The ceca extend to near the posterior end of the worm. 

The vitellaria fill the posterior 3/4 of the body. The testes are in tandem in the 

posterior body in between the ceca. One occurred in an Atlantic bonito and 2 

in an Atlantic mackerel from Woods Hole, Massachusetts, USA. It occurs in 

fishes in Puerto Rico and Jamaica, but has not been found in big game fishes 

from the Caribbean. This worm is known from the western Atlantic. It occurs 

in the intestine and pyloric ceca of its hosts and is 1.0-3.4 mm long. It does not 

appear to have much host preference as it occurs in a great variety of fishes. 

Opisthadena dimidia Linton - One to 12 occurred in 9 of 303 swordfish 

from the northwest Atlantic (ARC 2320). This was probably a false host since 

this worm appears to be genus specific to sea chubs (Kyphosus) in the western 

Atlantic and eastern Pacific. This worm has a relatively large ventral sucker 

that almost occupies the entire width of the worm. The sides of the worm bulge 

out around the ventral sucker. The ventral sucker is more than 3 times as large 

as the oral sucker, and the 2 are less than 1 ventral sucker width apart. The 

body is elongate. The testes and ovary are in the posterior 1/2 of the body, and 

the vitellaria are confined to 2 lobes in the hind-body posterior to the testes and 

ovary. It occurs in the intestine of its hosts and is 2.8-8.4 mm long. 

Pinguitrema lerneri (Sogandares-Bernal) - We found 1 in a bar jack 

from La Parguera, Puerto Rico (USNPC 83011), but this was probably a false 

or accidental host. This parasite is apparently host specific to a mojarra 

(Gerreidae) in which it has been found from Puerto Rico and Florida, USA. 

Pachycreadium lerneri S.-B. and Pinguitrema lobatum Siddiqi and Cable are 

synonyms. It is an oval worm with a rather large, oval ventral sucker 

occupying about 1/3 to 1/2 of the body width just anterior of the middle of the 

body. Around the ventral sucker, the body covering is distorted into obvious 

oval wrinkles. The vitellaria surround the ceca from near the bifurcation to near 

the end of the worm. It occurs in the intestine and is 1.0-1.5 mm long. 

Podocotyle chloroscombri (Fischthal and Thomas, 1970) - Yellow 

jack may be a false host for this parasite of Atlantic bumper. It is similar to P. 

simplex but differs by having its ventral sucker on a distinct peduncle. Two 

mature and 3 immature worms occurred in 1 of 8 yellow jack from Drowned 

Cays, Belize (USNPC 74238). It is 2.0-3.2 mm long and occurs in the intestine 

of its hosts. 

Podocotyle simplex (Rudolphi) - This fluke may be only an accidental 

parasite of big game fishes. It is an elongate worm with the ventral sucker 

larger than the oral sucker and in the anterior half of the body. The pharynx is 

relatively large, the esophagus short, the ceca extend to near the posterior end 

of the worm, the vitellaria surround the ceca in the posterior part of the body, 

and the testes are in the posterior 1/2 of the body. This fluke was reported in 

61

62 


Atlantic mackerel from the Atlantic coast of Canada, but this may have been an 

accidental or a false host. It occurs in the intestine and pyloric ceca of its host, 

is 1.2-4.0 mm long and has been found in a variety of fishes. 

Prosorhynchus pacificus Manter - This is a parasite of western Atlantic 

and eastern Pacific groupers (Serranidae). Prosorhynchus atlanticus Manter is 

a synonym. It has a rather large, long, cone-shaped rhynchus. The vitellaria 

are confined in 2 lateral rows in the anterior 1/2 of the body, and the uterus, 

with relatively large eggs, is confined in the posterior 1/2 of the body. One to 

14 occurred in 17 of 52 bar jack from Bermuda, but this was probably a false 

or accidental host. This worm occurs in the intestine of its hosts, and is 0.7-1.7 

mm long. See Ectenurus lepidus for associations. 

Pseudolepidapedon pudens (Linton) - This fluke may have only 

accidentally occurred in a cobia. This oval fluke has an oral sucker smaller than 

the ventral sucker and separated from it by more than 2 diameters of the ventral 

sucker. A conspicuous "pit" occurs between the oral and ventral suckers. The 

prepharynx is short, the pharynx is almost as large as the oral sucker, the ceca 

are relatively thick and extend to near the posterior end of the worm. The 

vitellaria surround the cecal branches. The testes are in the posterior body, in 

tandem and occupy all the space between the ceca. The eggs are few in number 

and relatively large. A few eggs are present and the vitellaria are developed in 

immature worms in the intestine of the host. The ovary and testes in immature 

worms are approximately 1/2 the size found in adults. One fluke was found in 

1 of 6 cobia from Beaufort, North Carolina, USA. It is only known from the 

Atlantic coast of the USA. This worm occurs in the intestine of its host and is 

1.6-2.7 mm long; and immature worms are 1.2-1.4 mm. This parasite appears 

to be genus (Paralichthys), or family (Bothidae), specific to flatfishes or 

flounders. The single specimen of this fluke found in a cobia was small (1.3 

mm), in poor condition, and possibly an immature worm. Cobia could be an 

accidental or a false host for this parasite. 

Pseudopecoelus elongatus (Yamaguti) - One record in a horse-eye jack 

from Brazil could represent a false host. It has been reported in other fishes 

from Japan and Brazil. This worm is similar in general appearance to 

Pseudopecoeloides carangis, but lacks the distinct peduncle separating the 

ventral sucker from the body and the ventral sucker is larger than the oral 

sucker. It is 2.4-6.1 mm long and occurs in the intestine of its hosts. 

Rhipidocotyle longleyi Manter - The great barracuda is a false host for 

this fluke that is usually found in deeper-water fishes. This minuscule to tiny, 

elongate worm has a muscular, bowl-shaped rhynchus covered with a flat 

pentagonal hood. The sucker around the mouth is smaller than the rhynchus and 

posterior to midbody. It occurs in deep-water (256-428 m) fishes off Florida 

and Japan. This worm was reported once in great barracuda from Florida,


USA. It probably occurs worldwide, but has only been confirmed from Florida 

in the Atlantic and Japan in the Pacific. This worm occurs in the intestine of 

its hosts, and is 1.7-3.7 mm long. These worms were obtained by the great 

barracuda feeding on deep-water fishes. A great barracuda, that one of our 

students caught-and-released on the way to a dive site, followed our submersible 

down to 300 m depth in the Bahamas. Thus this predator is capable of feeding 

on fishes at 256-428 m depths. 

Stephanostomum aulostomi Nahhas and Cable - We found this fluke 

in a bar jack, but it may be a false host. This is a tiny, elongate, fluke with 36 

circumoral spines. The oral and ventral suckers are separated by more than 3 

diameters of the ventral sucker, and the ventral sucker is approximately twice 

as large as the oral sucker. The vitellaria extend to the ventral sucker. The 

pharynx is almost in contact with the intestinal ceca bifurcation. Seven occurred 

in 1 of 2 bar jack from La Parguera, Puerto Rico (USNPC 85943). This worm 

occurred singly in a peculiar attachment position, between the stomach and the 

intestine, in 4 of 9 trumpetfish, Aulostomus maculatus Valenciennes, from 

Curaçao; but in the intestine in a bar jack. This worm is 5.5-7.0 mm long. Bar 

jack is a new host, but it may be a false or accidental one. 

Stephanostomum dentatum (Linton) - This fluke was reported once from 

cobia and pompano dolphin, but they are probably false or accidental hosts. 

Cercaria dipterocerca Miller and Northrup is the cercarial stage of this fluke. 

This is a minuscule, stocky fluke with 54-58 circumoral spines. The oral and 

ventral suckers are separated by more than 3 diameters of the ventral sucker, 

and are approximately equal in size. The vitellaria extend to the ventral sucker. 

The pharynx is in contact with the intestinal ceca bifurcation. One immature 

worm occurred in 1 of 6 cobia; and 2 immature in 1 of 3 pompano dolphin from 

Beaufort, North Carolina, USA. This fluke has been noted in a great variety of 

inshore fishes from the Atlantic and Pacific USA coasts and in the Caribbean. 

Redia inside eastern mudsnail, Iyanassa obsoleta Say, produce cercariae which 

escape and penetrate the pharyngeal region of Atlantic silversides, Menidia 

menidia (Linnaeus). The resulting encysted metacercariae become adults when 

the silversides are eaten by an appropriate final host. The adult fluke is found 

in the intestine of the final host and is 1.1-2.8 mm long. This is a generalized 

and widespread parasite of flounders and groupers. The worms described from 

pompano dolphin, as lacking oral spines (Linton 1905), were probably 

misidentified, and were inadequately described. 

63

64 


DIDYMOZOIDEA (TISSUE FLUKES) 

These vivid and often spectacular worms are found encapsulated in the 

tissues of marine fishes, including big game fishes, but a few (probably of 

marine origin) infect freshwater fishes. These capsules (and the damage they 

cause) are frequently seen by fishermen but are rarely mentioned in general 

parasitology, and even many fish-parasitology, texts. Possibly more editors 

need to be taken big game fishing! No common name has been given to these 

worms so we suggest "tissue flukes". This name distinguishes tissue flukes 

encapsulated in the tissues of their hosts from the loose or free-moving flukes 

in the gut and other passages of fishes, and frees us from trying to pronounce 

"didymozoid". The scientific name is from "didymos" which means double or 

twin, and "zoë" for life. It refers to the characteristic 2 worms which usually 

occur in each capsule. The bright, usually yellow, color that makes these 

capsules so conspicuous is due to masses of eggs which occupy much of their 

bodies. Tissue flukes do not produce a cyst around themselves and are seldom 

enclosed in a heavy, host-produced capsule. Usually, they are surrounded by 

a thin connective tissue layer produced by the host, or occasionally there is no 

connective tissue. We use the term "capsule" to refer to the encapsulation of the 

worms in the tissue of the host. 

Tissue flukes are not known to harm humans. Eggs of these worms from 

flyingfishes have probably been ingested by humans. Nikolaeva (1985) suggests 

that metacercariae in raw fish could adapt to humans or migrate through human 

tissues and become dangerous. Economically, tissue flukes in muscle tissue of 

fish may cause it to deteriorate more rapidly, and makes it less desirable for 

consumers. Heavily infected Atlantic mackerel flesh has to be discarded in 

processing, and wahoo in eastern Australia is undervalued due to its reputation 

for having large tissue flukes (a third are infected). Heavy to very heavy 

infections are routinely found in big game fishes. In the Indian Ocean, 100 or 

more capsules are found in every wahoo and half the tuna. Superinfections, up 

to 1167 capsules, have rarely been reported in tuna from the southern Gulf of 

Mexico. In contrast, in the central and eastern Atlantic, the highest infection 

reported from tuna was 148 capsules. 

Approximately 200 species are known, and many probably await discovery 

due to their unusual and seldom examined locations in hosts. Ironically, a few 

of these 200 species are duplicate descriptions of the same worms (synonyms) 

due to their complexity, unavailability and difficulty in removing intact 

specimens. Encapsulated pairs of various species range from a few millimeters 

to the size of your fist. They are usually colored yellow or orange, but blue 

capsules occur in wahoo from Australia. Worms vary in length from a few 

millimeters to over 12 meters. In some species, the sizes of both capsules and 

worms vary between different tissues in the same fish. The body of the worm 

may be elongated, filamentous or ribbon-like, and intricately tangled, or divided 

into a narrow anterior and a broadly swollen posterior. 

No complete life cycle is known. Release of eggs from encapsulated worms


is accomplished annually in those from 

gills, ovaries, or other exposed locations. 

More deeply embedded or inaccessible 

worms may only release eggs after the 

death of the host and survive passage 

through the gut of the predator. In some 

cases, the capsule and surrounding tissue 

breaks down or ulcerates when the tissue 

flukes mature, releasing both eggs and 

adults. A non-ciliated miracidium with 

2 or more circles of spines around the 

oral sucker hatches from the egg. Tissue 

fluke cercariae occur in plankton, and 

metacercariae in arrow worms, bar- 

nacles, copepods, krill, squids, small 

bony fishes and sharks. Very heavy 

larval infections in squid suggest that 

they are important intermediate hosts of 

the life cycle of many tissue flukes. 

Also, actively migrating, larval stages 

have been described in fishes. Juvenile 

stages may be found in a variety of 

hosts, but only those in the proper final 

hosts develop into adults, the rest act as 

Larvae of Didymocystis acanthocybii. 

from wahoo 

transport hosts and are important in enabling parasites to reach their final hosts. 

All species are hermaphroditic (have both sexes in each worm), but often 

associate in pairs, one individual of which has female organs more developed 

and a smaller partner that has more developed male organs. Other pairs are 

fused to each other in the genital region. Presumably, the first worm to arrive 

at a site in the host becomes the "female" and the later arrival the "male". 

Usually a single male and female occur in each capsule, but 2-3 males may 

occasionally be found with a female. They encapsulate in the gills, skin, under 

scales, in connective tissue, muscle, fat, bone, teeth, eye socket (orbit), oral or 

nasal cavity, viscera, in the body cavity, vascular system or most any other 

tissue or organ. They are permanent or semi-permanent parasites in the final 

hosts. Most are assumed to survive for the life of the host or at least 

identifiable remains persist. Those in the gills or gonads may be lost each year. 

Tissue flukes are found largely in tropical and temperate pelagic fishes, but 

also occur in some more sedentary fishes, such as groupers in Puerto Rico. 

Scombrids appear to be the preferred hosts with approximately 65% of tissue 

fluke species occurring in them. Approximately 100 species of fishes, including 

other pelagics, such as flyingfishes and ocean sunfishes, Mola sp., 8 species of 

barracuda (but not our great barracuda) and jacks, are infected. Two thirds of 

the big game fish species in this book are infected. Fewer species of tissue 

flukes occur in western Atlantic big game fishes compared to other warm-water 

regions (see Checklists). More tissue fluke species are reported from the Indo- 

65

66 


Pacific and the eastern Atlantic. This disjunct distribution may simply be due 

to greater numbers of studies on these animals in Japan, Hawaii and Europe, 

but, if many of these species are truly found in the eastern but not the western 

Atlantic, they could make good biological tags to document trans-Atlantic 

migrations. 

Modern classifications place tissue flukes as superfamily or a family in the 

Subclass Digenea (flukes). Their unique morphologies and habitats in the host, 

occasionally almost separate sexes, and life cycles possibly without a mollusk 

first intermediate host, make them worthy of superfamily status. Tissue flukes 

were once separated into their own subclass largely because they were 

incorrectly thought to have a direct life cycle. Even though schistosome flukes 

have separate sexes and sanguinicolid flukes possibly equally odd life cycles, the 

combination of differences still distinguish tissue flukes into a superfamily. We 

treat them separately because fishermen can readily distinguish these relatively 

large, colorful and abundant parasites of big game fishes. 


Subclass (Infraclass) Digenea - flukes Page 

Superfamily Didymozoidea - tissue flukes 

Family Didymozoidae 

Subfamily Didymozoinae 

Didymocystis acanthocybii ............................................................... 67 

Didymocystis scomberomori ............................................................ 68 

Didymocystis thynni ......................................................................... 69 

Didymocystis wedli .......................................................................... 70 

Didymozoon auxis ........................................................................... 79 

Didymozoon longicolle .................................................................... 71 

Platocystis sp. .................................................................................. 79 

Subfamily Colocyntotrematinae 

Colocyntotrema sp. ......................................................................... 71 

Subfamily Koellikeriinae 

Koellikeria bipartita ....................................................................... 72 

Koellikeria globosa ........................................................................ 73 

Koellikeria orientalis ..................................................................... 74 

Subfamily Nematobothriinae 

Atalostropion sardae ...................................................................... 74 

Atalostropion sp. ............................................................................ 79 

Nematobothrium pelamydis ........................................................... 75 

Nematobothrium scombri .............................................................. 76


Subfamily Neodidymozoinae 

Maccallumtrema xiphiados .............................................................. 77 

Neodidymozoon macrostoma ........................................................... 78 

Miscellaneous Tissue Flukes ............................................................ 79 

Didymocystis acanthocybii Yamaguti 

This worm occurs in the tissues of the 

head of wahoo, possibly around the world. 

Name - Nigrelli (1939) described the same 

worm as D. coatesi Nigrelli. He was 

apparently not aware that it had been 

described the year before. 

Diagnostic Characters - It usually occurs 

as rather large, lumpy orange to yellow 

masses on the gill arches, eye sockets or 

other tissues of the head. In worms 

removed from the capsule, the forebody 

projects from one end of the hindbody and 

is more than 1/2 the length of the 

hindbody. The hindbody is bulbous and bean-shaped. 

Records - Two capsules occurred in 2 of 15 wahoo from Puerto Rico. One 

infected fish was from La Parguera and 1 from Arecibo, Puerto Rico (USNPC). 

Four and 38 capsules were found in 2 of 10 yellowfin tuna from the southern 

Gulf of Mexico. This parasite also occurs in wahoo on the USA Atlantic coast 

and in the Pacific. 

Geographic Range - Worldwide. Our records are the first in the Caribbean. 

Life History - Larvae (posttorticaecum) in fish are 2.4-3.0 Φm long and 0.3-0.7 

µm wide (see figures in Tissue Fluke introduction). Eggs produced by the 

larger larvae were slightly smaller than those in adults. Larval forms have been 

found in the tissues of the head of wahoos. 

Associations - A very heavy infection of gillworms, Neothoracocotyle acanthocybii, 

also occurred on the fish from Arecibo. These may have affected the 

health of the fish, but probably did not otherwise interact with the tissue flukes. 

Location in Host - It is encapsulated in the outer layers of gills, mouth, eye 

sockets and face; and in internal tissues of the head. We found capsules in the 

roof of the mouth (La Parguera), and gill filaments near the gill arch (Arecibo). 

Length - Worm 2.8-23.0 mm (forebody 1.0-11.0, hindbody 1.8-13.0 mm), 

capsules 3.0-25.0 mm. Yamaguti (1970) found capsules and worms of different 

sizes in different tissues of the heads of wahoos in Hawaii. 

Host Specificity - It only occurs in wahoo. The report of this parasite from 

yellowfin tuna (Nikolaeva 1968) requires further confirmation. 

Detection - These capsules were obvious in the gills and mouth of the fish we 

examined. They should be equally apparent on the face and other exposed 

surfaces of wahoo, but may be less noticeable in other tissue of the head. 

67

68 


Preparation for Study - Careful dissection may be necessary to find hidden 

capsules and to ascertain which tissue was infected. Larval forms have been 

rinsed from dissected heads of wahoo. 

Didymocystis scomberomori (MacCallum and MacCallum) 

This potentially harmful parasite occurs as 

a bright mass in the gills and viscera of 

inshore, but not offshore, Spanish mackerels 

throughout the western Atlantic. 

Name - It was originally placed in genus Koellikeria, 

and was refigured and redescribed by 

Overstreet (1969). 

Diagnostic Characters - Capsules usually 

occur as rather large, lumpy orange to yellow 

masses. When removed from the capsule, the 

forebody projects from one end of the bulbous 

and bean-shaped hindbody and is less than 1/2 

the length of the hindbody. 

Records - We found 2-6 capsules in 7 of 30 

cero from La Parguera (USNPC); more than 

50-100 capsules in 2 of 3 cero off Humacao, Puerto Rico (USNPC 82966); and 

a few capsules in 2 serra Spanish mackerel from Cartagena, Colombia 

(USNPC). Twenty-five capsules were found in 2 Spanish mackerel and 1 of 2 

cero from Biscayne Bay, Florida, USA (USNPC 71319); and 4 worms and 1 

capsule in 2 Spanish mackerel from Woods Hole, Massachusetts, USA. 


Life History - The capsules in the gills, intestinal wall and liver might suggest 

that larval forms penetrate the blood stream of the host and are transported to 

these sites. 

Ecology - This worm has been found in all 3 species of western Atlantic, 

nearshore Spanish mackerels, but not in the more offshore king mackerel. 

Other parasites follow this same pattern (see Discussion). 

The area in eastern Puerto Rico with the heavy infection rates is known to 

be contaminated with heavy metals and other pollutants (Tetra Tech 1992) (see 

Hysterothylacium reliquens). La Parguera in western Puerto Rico is less 

polluted. However, our data are too limited to make any environmental 

correlations. 

Associations - A female-male pair of isopods, Livoneca redmanii, often occur 

in the gill chamber with the tissue fluke capsules. However, our limited 

statistical analysis seems to show no relationship between presence of the 

isopods and the capsules. In the fish with more than 100 capsules, heavy 

infections of encysted larval roundworms, H. relinquens, and tapeworms also 

occurred in the same areas of the gut. 

Location in Host - It is encapsulated in the gills, usually in an arch, but 

occasionally in filaments; in the wall of the stomach or intestine; exceptionally,


in the liver. Capsules were found in the liver only in the heavily infected cero. 

This may represent an unusual site only utilized in heavy infections. In some 

fish, the capsules were limited to the gills or the gut, but in others all sites were 

involved. This variation has not been explained. 

Length - Worms 0.9-2.2 mm. Small capsules of a few millimeters occur in the 

intestinal wall and liver, but small to large ones up to 24.5 mm long and 12.0 

mm wide are found on the gill arches. 

Host Specificity - It is genus specific (Scomberomorus). Apparently it does 

not occur in king mackerel. This worm is a secondary parasite of Spanish 

mackerels. Serra Spanish mackerel is a new host. 

Damage to Host - Heavy to very heavy infections of this worm had not 

previously been reported. The levels we found were probably sufficiently high 

to cause injury to the host. More study is needed to determine how often and 

under what condition heavy infections occur. An infection in the gills of 

Spanish mackerel from Florida, USA, produced a fibroblastic encapsulation 

(RTLA 4501). 

Detection - The rather large, bright orange to yellow lumps on the gills or 

organs are obvious. 

Harm to Humans - We have not hesitated to eat numerous cero we collected 

that were infected with this worm (possibly not an endorsement, as you might 

become horribly transformed like us into parasitologists!). 

Significance to Sport Fishing - These large, obvious to spectacular lumps on 

the gills and guts of Spanish mackerel may offend fishermen cleaning their 

catch, but the worms are not known to harm humans. 

Didymocystis thynni (Taschenberg) 

This is a potentially 

damaging parasite of tunas and 

little tunas across the Atlantic. 

Name - Didymocystis reniformis 

Ariola is a synonym. 

Diagnostic Characters - Capsules 

appear as small, lumpy 

yellow masses. The forebody of 

the female is relatively short and 

shorter than the length of the 

hindbody. The oral sucker is 

relatively large and wider than 

1/2 the width of the anterior 

expansion. 

Records - Three to more than 70 capsules occurred in 2 little tunny from La 

Parguera, Puerto Rico. This parasite did not occur in 4 other little tunny we 

examined from Puerto Rico or 1 from the northern Gulf of Mexico off 

Alabama, USA. It also occurs in albacore, bluefin tuna and little tunny in the 

Mediterranean and bluefin tuna and skipjack tuna off Europe in the Atlantic. 

69

70 


Geographic Range - Atlantic and Mediterranean. Our records are the first in 

the western Atlantic. 

Associations - Very heavy infections of 4 species of flukes occurred in the 

intestinal tracts of 2 little tunny with tissue flukes. 

Location in Host - Most capsules occurred in the throat and 3 occurred in the 

wall of the anterior stomach of 1 little tunny; and 3 capsules occurred in the 

intestinal wall just posterior to the stomach of a second fish. This worm was 

previously reported encapsulated on the gills and inside the gill cover 

Length - Worm 5.1-5.6 mm (forebody 2.6-2.8, hindbody 2.5-3.0 mm), 

capsules 0.7-13.0 mm (in throat 0.7-2.0, gills and gill cover 2.0-4.0, stomach 

7.0-13.0 mm). 

Host Specificity - It is family specific to scombrids. 

Damage to Host - The heavy infection of this tissue fluke in combination with 

a very heavy infection of flukes in little tunny would probably be a sufficient 

parasite load to injure this host. 

Detection - These small, bright yellow capsules are easily seen in the gills, gill 

cover or throat, but may be less obvious when more deeply embedded or in the 

intestinal tract wall. 

Significance to Sport Fishing - The potential for damage to big game fishes 

makes further study of these parasites prudent. 

Didymocystis wedli Ariola 

This is a potentially dangerous 

parasite of tunas, little tunas and 

mackerels around the world. 

Diagnostic Characters - In worms 

removed from the capsule, the forebody 

projects from one end of the hindbody 

and approximately 1/2 the length of the 

hindbody.The hindbody is elongate with 

a 2 lobes on the front (bilobed anterior) 

and a twisted tail (spirally twisted). 

Records - We found 17 capsules in 1 of 

2 yellowfin tuna from La Parguera, Puerto Rico; and 26 in 1 of 2 yellowfin 

tuna off Dauphin Island, Alabama, USA. From 4-526 capsules (average 333) 

occurred in 10 yellowfin tuna in the southern Gulf of Mexico. This worm was 

also found in albacore and bluefin tuna in the Mediterranean, bluefin tuna, chub 

mackerel and skipjack tuna in the Pacific, and frigate tuna in India (BMNH 

1981.6.9.5). 

Geographic Range - Worldwide. Our collections were the first from the 

Caribbean and the northern Gulf of Mexico. 

Location in Host - Gills filaments. 

Length - Worm 4.8-5.6 mm (forebody 0.9-1.4, hindbody 4.5-5.1 mm). 

Host Specificity - This worm is family specific (Scombridae) and almost tribe 

specific to tunas, but it also infects a little tuna and a mackerel.


Damage to Host - Nikolaeva (1968) reported hundreds of capsules of this 

worm and Didymozoon longicolle in yellowfin tuna which were also parasitized 

by heavy infections of 2-3 other tissue flukes (up to 1167 capsules). Superinfections 

are sufficient to stunt or injure these hosts and are rather alarming 

because few studies have targeted these parasites in the western Atlantic. We 

cannot be certain that damaging levels are not routine. "Large numbers" have 

also been reported in the gills of bluefin and skipjack tuna in Japan. 

Didymozoon longicolle Ishii 

This parasite is potentially damaging to tunas, little tunas 

and mackerels around the world. 

Name - The Didymozoon sp. described from a chub mackerel 

by Linton (1940) was probably this worm. 

Diagnostic Characters - Capsules are as small yellow, oval 

and elongate. In worms removed from the capsule, the forebody 

is elongate and narrow and the hindbody is cylindrical 

with a conical anterior end and abroadly rounded posterior end. 

Records -Ten to 120 capsules (average 51) occurred in 6 of 10 

yellowfin tuna from the southern Gulf of Mexico; 1-3 capsules 

in 2 chub mackerel from Woods Hole, Massachusetts, USA 

(USNPC 8389). It was also found in bluefin tuna, chub 

mackerel, skipjack tuna and yellowfin tuna in the Pacific. 

Geographic Range - Worldwide. The 2 rather tenuous 

Atlantic records need to be confirmed with new collections. 

Location in Host - Gills. It was reported in the skin of 

yellowfin tuna from the Gulf of Mexico. 

Length - Worm 4.3-18.0 mm (forebody 0.8-3.8, hindbody 3.1- 

8.5 mm); capsule 3.0-5.0 mm. 

Host Specificity - This worm is family specific (Scombridae). 

Damage to Host - See Didymocystis wedli. 

Colocyntotrema sp. of Nikolaeva 

These bright orange globular capsules in 

the gills and internal organs are easily seen by 

fishermen,particularly when skinning sailfish. 

It is quite possible that this parasite damages 

billfishes. 

Name - Colocyntotrema auxis Yamaguti was 

reported in frigate tuna in the Pacific, but this 

worm appears to be a different species. 

Diagnostic Characters - Capsules appear as 

orange to yellowish, flattened oval masses, 

under the outer layer of organs. An elongated 

forebody projects from the globular 

hindbody when removed from the capsule. 

71

72 


Two identical worms in each capsule are fused at their posterior ends. 

Records - A heavy infection of this worm occurred in 1 of 5 Atlantic sailfish 

caught off Arecibo, Puerto Rico (USNPC), but not in 2 from the northern Gulf 

of Mexico off Alabama, USA. It has also been reported from Atlantic sailfish 

and Atlantic blue marlin from the southern Gulf of Mexico. The unidentified 

tissue flukes associated with longbill spearfish stomach ulcers (see Other 

Diseases and Conditions) could be this worm. 

Geographic Range - Northern Caribbean and southern Gulf of Mexico. Our 

collection is the first in the Caribbean. If its range is restricted, it might be of 

value as a biological tag. 

Life History - This parasite may be transported by the circulatory system of the 

host since it is found both in the gills and internally. 

Location in Host - More than 50 capsules occurred in the gills, 15 in the outer 

layer of the stomach and intestine, and 5 between the skin and body muscle in 

an Atlantic sailfish from Puerto Rico. This parasite was reported from the 

pyloric ceca of Atlantic sailfish and Atlantic blue marlin in the Gulf of Mexico. 

Length - The capsules varied in size from 2-3 mm in the outer layer of the gut, 

4-6 mm in the gills, and 5-8 mm under the skin. Whether these sizes were due 

to differences of nutrients available at different sites, or the duration of the 

different infections, could not be determined. 

Host Specificity - It is possibly family specific to billfishes. 

Damage to Host - The capsules were numerous enough to cause some impairment 

of the gills. The few capsules under the skin and on the gut probably 

cause little harm. 

Detection - The orange oval capsules are obvious. 

Significance to Sport Fishing - The potential of heavy infections injuring 

billfishes makes this parasite worthy of additional study. 

Koellikeria bipartita (Wedl) 

This Atlantic-wide parasite 

of tunas and greater 

amberjack, is either more 

important in Europe or 

better studied there. 

Name - It has also been 

placed in genus Wedlia. 

Diagnostic Characters - A 

larger female and smaller 

male worm pair are found 

in each capsule. In worms 

removed from the yellow 

capsule, the female has an 

elongate forebody with an expanded anterior end. The forebody is attached in 

the middle of the bean-shaped hindbody. The female worm looks like a cobra 

uncoiling from a basket. The smaller male has a similar forebody with a


relatively short neck about as long as the anterior expanded region, but a 

relatively small, globular hindbody. 

Records - One to 2 capsules occurred in 3 of 10 yellowfin tuna from the 

southern Gulf of Mexico; 4, 8 and 30 or more capsules in 3 bluefin tuna from 

Woods Hole, Massachusetts, USA (USNPC 8392); albacore, bluefin tuna and 

greater amberjack from the eastern Atlantic; and bluefin tuna and greater 

amberjack from the Mediterranean. It is either more abundant in Europe or less 

studied in the western Atlantic. 

Geographic Range - North Atlantic and Mediterranean. 

Location in Host - It is encapsulated in the wall of the intestine, pyloric ceca, 

gills, gill arches and skin of the head. 

Length - Female 1.6-9.0 mm (forebody 0.4-0.6, hindbody 1.2-2.0 mm), male 

2.9-9.0 mm (forebody 2.0-2.5, hindbody 0.9-1.4 mm), immature worm in a 

capsule 1.2 mm; capsules 3.0-9.0 mm. Capsules in the intestinal wall are 

smaller than those in the pyloric ceca. 

Host Specificity - It is almost tribe specific to tunas, but is also found in 

greater amberjack. 

Detection - Capsules in the intestine look like small yellow spots. 

Koellikeria globosa Ishii 

This marble-shaped parasite occurs 

worldwide in tunas and occasionally skipjack 

tuna and yellowtail. 

Name - The name "globosa" aptly 

describes the spherical hindbody of the 

female. This worm had also been placed 

in genus Wedlia. 

Diagnostic Characters - A larger 

female and smaller male worm pair are 

found in each capsule. The hindbody of 

the female is oval to round instead of 

bean-shaped and the oral sucker is 

relatively small. The male has a similar 

forebody, a neck longer than the 

expanded anterior, and a hindbody about 

the same size as the expanded anterior. 

Records - Two to 87 capsules (average 20) occurred in 9 of 10 yellowfin tuna 

from the southern Gulf of Mexico; several capsules in 1 of 64 yellowfin tuna 

from west Africa; and in bluefin tuna, skipjack tuna and yellowtail from Japan. 


Location in Host - It is encapsulated in the mouth, esophagus, intestine and 

pyloric ceca. 

Length - Female 9.0-19.4 mm, male 6.6-8.0 mm; capsule 2.0-5.0 mm. 

Host Specificity - It is almost tribe specific to tunas, but occasionally found 

in little tunas and jacks. 

73

74 


Koellikeria orientalis (Yamaguti) 

This is a worldwide parasite of tunas and 

occasionally skipjack tuna. 

Name - This worm has also been placed in 

genus Wedlia. 

Diagnostic Characters - A larger female 

and smaller male worm pair are found in 

each capsule. The female has an elongate 

forebody with an expanded 

anterior end. The forebody is 

attached in the middle of the 

bean-shaped hindbody. The 

female worm looks like a 

cobra uncoiling from a basket. 

The male has a similar 

forebody with a relatively long 

neck about 4 times the length of the anterior expanded region, but 

a relatively small bean-shaped hindbody. 

Records - Fifteen capsules occurred in 1 of 10 yellowfin tuna 

from the southern Gulf of Mexico; in bluefin tuna from the 

Mediterranean; albacore from the eastern Atlantic; and bluefin 

tuna, skipjack tuna, yellowfin tuna from Japan. 


Location in Host - It is encapsulated in the gills, esophagus, intestine, stomach 

and around the anus. 

Length - Female 2.5-2.8 mm (forebody 1.6- 

1.8, hindbody 0.9-1.1 mm), male 2.0-2.4 

(forebody 1.5-1.7, hindbody 0.5-0.7 mm); 

capsule 1.0-3.5 mm.Capsules in the gills and 

esophagus are larger (1.5-3.5mm) than in the 

stomach and intestine (1.0-2.0 mm). 

Host Specificity - This parasite is almost 

tribe specific to tunas, but has also been found 

in skipjack tuna. 

Atalostropion sardae MacCallum 

This is a strange and tangled worm. 

Name - Yamaguti (1971) refigured pieces of 

this worm. 

Diagnostic Characters - It occurs as a 

tangled mass of long, narrow, ribbon-shaped 

filaments in the tissues of fishes. Only the 

extreme anterior and posterior ends are 

shown. 

Records - Four to 43 capsules (average 14)


occurred in 6 of 10 yellowfin tuna from the southern Gulf of Mexico; many 

hundreds in Atlantic bonito, and in 7 of 16 Atlantic bonito from Woods Hole, 

Massachusetts, USA (USNPC 36309). 


Location in Host - In connective tissues of yellowfin tuna, and under the 

mucous membranes in the mouth of Atlantic bonito. 

Length - 140.0-190.0 mm, immature worms 50.0-70.0 mm (total adult lengths 

were estimated from pieces of worms). 

Host Specificity - This worm may only occur in Atlantic bonito. More 

collections are needed to confirm its occurrence in yellowfin tuna. 

Preparation for Study - Dissecting complete specimens from the tissues of the 

host is difficult, bordering on the impossible. 

Nematobothrium pelamydis (Taschenberg) 

These obvious, orange, often damaging gill capsules in 

Atlantic bonito are well known in the eastern Atlantic and 

Mediterranean. Their importance in the western Atlantic 

is more obscure. 

Name - Didymozoon pelamdis Taschenberg, N. sardae 

MacCallum and MacCallum, Unitubulotestis pelamdis (T.) 

and U. sardae (M. and M.) are synonyms. 

Diagnostic Characters - The orange capsules are oval and 

flattened. When removed from the capsule, worms are 

elongate with the anterior end much more narrow, but not 

distinctly set off as a forebody. The narrow anterior end 

is more than 4 times as long as wide. 

Records - One to 6 capsules occurred in 6 of 13 Atlantic 

bonito, and, in a study especially designed to determine 

prevalence and intensity of this worm, 2-3 capsules were 

found in 83 of 100 Atlantic bonito from Woods Hole, Massachusetts, 

USA. It is found in this host from the eastern 

Atlantic off Europe, the Mediterranean and Black Sea. 

Geographic Range - Northern Atlantic, Mediterranean 

and Black Sea. 

Life History - These parasites do not infect larval or small 

fishes, but Atlantic bonito 25.0-42.0 cm long as well as 

adults are infected. One larva enters a gill lobe blood 

vessel, a second larva joins the first, they grow, distort the 

artery and elicit host tissue response to form a capsule in 

the artery and enlarge, forming a sacculated aneurism. The 

first larva to arrive becomes the "female" (female gonads 

predominate) and the second becomes the "male". 

Metacercariae of this worm and N.scombri parasitize sprat, Clupea sprattus 

Linnaeus, and rarely 9 other species of Black Sea fishes. The number of these 

larval forms in sprat from the Black Sea have been monitored since 1950. From 

75

76 


1959 to 1961, 93.5% were infected with 1-116 metacercariae each. In 1975, the 

first of the final hosts of these tissue flukes, Atlantic bonito, became drastically 

reduced in numbers, and the second host, Atlantic mackerel, 

disappeared from the Black Sea. In 1977, only 1.0% of sprat 

were infected with 1 metacercaria each. 

Location in Host - Capsules usually occur along the outer layer 

of the gill filaments, rarely in the viscera. Larval forms were 

found under the lining of the gill chamber. 

Length - Adult worm 15.0-50.0 mm, immature worm encapsulated 

in the gills 18.0 mm, free juvenile 0.7-14.0 mm; capsule 

7.0-12.0 mm (3.0-3.5 mm wide). 

Host Specificity - It only occurs in Atlantic bonito. This worm 

can occur commonly in parts of the Mediterranean and Black Sea, 

but it is a secondary parasite of this host. 

Damage to Host - Very heavy infections, possibly capable of 

injuring the host, have been reported in fish during their return 

migration from the Black Sea to the Sea of Marmara. In some 

studies, 64.8% of the fishes were infected with up to 10 capsules 

per gill arch. Although the blood flow in the gill filament is 

clearly impaired by larva settling in the artery, little host reaction 

was apparent. A small host, 20 cm long, with a heavy infection 

of 10 capsules per gill arch, had pale gill filaments. 

Detection - The orange capsules are obvious on the gills. 

Significance to Sport Fishing - This rather important parasite 

of Atlantic bonito is probably the best studied of the tissue flukes, 

but it has been ignored in the western Atlantic. 

Nematobothrium scombri (Taschenberg) 

This parasite has only been found once in the western Atlantic 

in a butterfish, but occurs around the world in big game fishes. 

Name - It was previously placed in the genus Didymozoon 

(actually as the type species). 

Diagnostic Characters - The pale yellow capsules are usually 

isolated, but sometimes occur in groups. Worms are elongate 

with the anterior end much more narrow, but not distinctly set off 

as a forebody. The narrow anterior end is approximately twice as 

long as wide. 

Records - Seven worms occurred in a butterfish, Peprilus 

triacanthus (Peck), from Woods Hole, Massachusetts, USA 

(USNPC 8386); and in chub mackerel from southern Brazil. This 

parasite was also found in Atlantic mackerel from off Europe and 

the Mediterranean, chub mackerel from the Mediterranean, and 

chub mackerel and skipjack tuna from Japan. 


Life History - Capsules usually contain 1 pair of worms, but may


hold up to 16. See previous worm. 

Location in Host - This worm occurred in the inner surface of mouth, gill 

cover or gill arches of mackerels, but in the intestinal wall of butterfish. 

Length - Worms 15.0-65.0 mm; capsules 4.0-7.0 mm. 

Host Specificity - It is almost tribe specific to mackerels. The record in a 

butterfish may have been an accidental host or a data-recording error. 

Maccallumtrema xiphiados (MacCallum and MacCallum) 

Capsules of this cauliflower-shaped worm may 

damage or deteriorate the muscle of swordfish. 

Name -Yamaguti (1970) named a new genus in 

honor of the MacCallums based on their 

Koellikeria xiphiados. This worm has also been 

placed in the genera Didymocystis and Wedlia. 

The genus Makairatrema Yamaguti was also 

based on a species, Makairatrema musculicola 

Yamaguti (=Maccallumtrema musculicola) from 

the ventral abdominal muscle of a billfish. We 

fail to see any adequate distinctions between these 

2 genera and suggest that Makairatrema is a 

synonym of Maccallumtrema. This combines 2 

rather similar species from the muscles of 

billfishes into 1 genus. This action seems justified 

by the overlapping characters of the 2 genera, and 

particularly in a family with too many genera for its number of species. 

Diagnostic Characters - Capsules appear as large, ovid masses in the 

abdominal muscle. The capsule has extensions (vascular septa) that separate the 

hindbodies of the 2 enclosed worms. In worms removed from capsules, the 

forebody is attached in the middle of the hindbody, and the hindbody is 

cauliflower-shaped with lobes. 

Records -Several capsules occurred in swordfish from Woods Hole, Massachusetts, 

and Hawaii, USA. 


Location in Host - In the abdominal muscle in the upper (dorsal) flanks, the 

same muscle that is so commercially prized. 

Length - Worms 17.0-29.5 mm (forebody 3.0-6.5, hindbody 14.0-23.0 mm), 

exceptionally 40.0 mm; capsule up to 50 mm. 

Host Specificity - This worm only occurs in swordfish. 

Detection - Larger, yellow capsules near the surface of the muscle are obvious 

as soon as the fish is skinned. Smaller, less developed (white instead of 

yellow) or more deeply embedded capsules may be less obvious. 

Harm to Humans - Indirectly, the presence of these capsules could harm 

humans by causing the muscle tissue used as human food to deteriorate more 

rapidly than would be expected. Capsules should be picked out (mechanically 

removed) from swordfish steaks to prevent contamination and deterioration. In 

77

78 


cases of heavy infections, it may be wise to cook the flesh as soon as possible. 

Cooking fresh infected steaks before freezer storage may be prudent. 

Preparation for Study - Locating capsules without damaging them and 

dissecting out intact capsules and worms is a bit more of a challenge in muscle 

tissue than in dealing with capsules on the surface of other tissues. Sections of 

muscle, no more than a few centimeters thick, with capsules should be excised 

from freshly caught swordfish and placed in 10% buffered formalin (10 parts 

fluid for 1 part tissue). 

Significance to Sport Fishing - This parasite should be of considerable 

interest to fishermen, seafood inspectors and seafood processors. Our almost 

utter lack of knowledge of this important problem is astounding. 

Neodidymozoon macrostoma Yamaguti 

This worm is encapsulated on the gills of 

billfishes around the world. 

Diagnostic Characters - It occurs in a 

relatively large, ovid flat capsule with host 

tissue intruding between the 2 worms. Worms 

removed from the capsule have a cauliflowershaped 

hindbody with small slender forebody 

attached in the middle. 

Records - One capsule occurred in 1 of 40 

Atlantic blue marlin from various localities, 6 

in a longbill spearfish from Aguadilla, and 1 in 

a white marlin off La Parguera, Puerto Rico. 

This parasite was described from Hawaii in 

Indo-Pacific blue marlin and 2 other billfish 

species. 

Geographic Range - Worldwide. Our collections are the first in the Atlantic. 

Location in Host - In the outer 

layer of the gill arch in our 

specimens; and reported previously 

in the connective tissues of the gill 

arch, and in fat and muscle around 

the gill chamber. 

Length - Worm 11.0-25.5 mm 

(forebody 3.0-5.5, hindbody 8.0-20.0 

mm); capsule up to 12.0 mm. 

Host Specificity - This parasite is 

family specific to billfishes. Our 

hosts are new records for this 

parasite. 

Detection - The larger capsules are 

obvious on the arches and in the gill 

chamber.


Miscellaneous Tissue Flukes 

Atalostropion sp. of Grabda. - Grabda (1991) reported Atlantic mackerel 

from the Georges Bank in the northwest Atlantic with this worm intertwining 

between the muscle fibers and forming orange-colored centers. These parasites 

hinder commercial processing of Atlantic mackerel by ruining fillets. Fillets 

with worms were soft, watery and grayish in color. This worm may also be the 

tissue flukes found in connective tissue between the muscle bundles of Atlantic 

mackerel (RTLA 4503). This parasite is causing problems in a commercial 

fishery and should be defined and evaluated. 

Didymozoon auxis Taschenberg - The Didymozoon sp. described from 

a bullet tuna by Linton (1940) could possibly be this worm. New collections 

will be necessary to resolve this question. Worms removed from the capsule 

have a distinctive, long, narrow body that is bent at a right angle. The forebody 

is elongate and narrow. One capsule possibly occurred in a bullet tuna from 

Woods Hole, Massachusetts, USA (USNPC 8388). It has also been reported 

from bullet tuna in the Mediterranean and frigate tuna from Japan. This parasite 

probably occurs worldwide. Capsules are found on the gills of hosts. Worms 

are 3.9-12.0 mm (forebody 0.8-1.8, hindbody 3.1-8.2 mm) long; capsule 4.0 

mm. It is genus specific (Auxis). 

Didymozoon minor Yamaguti - The name of this worm, found in skipjack 

tuna from Japan, was changed to Didymozoon minus, but most confusingly, both 

names are found in the literature and in parasite checklists. 

Platocystis sp. of Nikolaeva - This worm apparently differs from 

Platocystis alalongae Yamaguti found in the skin of Pacific albacore. The flat 

capsule contains 2 worms. Worms liberated from the capsule have an elongate 

forebody attached to one end of a globular, semicircular (slice of watermelonshaped) 

hindbody. Two capsules occurred in the connective tissues of 1 of 10 

yellowfin tuna from the southern Gulf of Mexico. 

MONOGENEA (GILLWORMS) 

The name "monogenea" means born once, and refers to the simple life 

cycle. In heavy infections, they can kill captive fishes and occasionally wild 

ones. More than 1500 species have been described, but this is probably only a 

small percentage of those existing. Adults range from 30 Φm to 20 mm in 

length and are translucent, cream or pink. Gillworms have a distinct attachment 

organ on their posterior end called a haptor (or opisthaptor) with hardened 

anchors or specialized clamps to pierce the epithelium and hold on to the host. 

Sclerotized marginal hooks often surround the haptor, and bars, disks, scales or 

spines may occur on or near the haptor. The head sometimes has eye spots and 

79

80 


specialized holdfast organs. Many monogenean parasites of big game fishes 

have large suckers or numerous clamps adapted for holding on to these fast 

moving animals. Most reproduce by laying eggs that hatch ciliated larvae (oncomiracidia) 

and quickly mature and attach to the host. Since no stages on 

intermediate hosts are necessary, they can multiply rapidly. When intensive 

culture crowds fish together, most gillworm offspring survive and can quickly 

begin to kill fishes. Gillworms are permanent parasites in the gills, mouths or 

on the bodies of fishes. Some occur in the nares, pockets in the lateral line or 

rarely in the gut of fishes. Some species occur in the urinary bladder of fishes, 

frogs or turtles. They generally feed on mucus, epithelial cells or blood. Gillworms 

are common on fishes in all aquatic environments. 

Popular Reference "How to Know the Trematodes" (Schell 1970). 


Class (Infraclass) Monogenea - gillworms Page 

Order Capsalidea ....................................................................................... 81 

Family Capsalidae 

Caballerocotyla manteri .................................................................. 82 

Capsaloides cornutus ....................................................................... 83 

Capsaloides magnaspinosus ............................................................ 83 

Nasicola klawei ................................................................................ 84 

Tristoma coccineum ......................................................................... 85 

Tristoma integrum ............................................................................ 86 

Tristomella laevis ............................................................................. 86 

Tristomella lintoni ............................................................................ 103 

Tristomella onchidiocotyle .............................................................. 103 

Family Dionchidae 

Dionchus agassizi ........................................................................... 88 

Dionchus remorae .......................................................................... 102 

Order Mazocraeidea .............................................................................. 88 

Family Axinidae 

Allopseudaxine katsuwonis ............................................................ 89 

Family Heteraxinidae 

Allencotyla mcintoshi .................................................................... 89 

Cemocotyle carangis ..................................................................... 90 

Cemocotyle noveboracensis .......................................................... 91 

Cemocotylella elongata ................................................................. 91 

Helixaxine winteri ......................................................................... 92 

Family Allopyragraphoridae 

Allopyragraphorus incomparabilis .............................................. 93


Family Hexostomatidae 

Hexostoma euthynni .................................... .................................. 94 

Hexostoma lintoni ......................... ................................................ 94 

Family Protomicrocotylidae 

Protomicrocotyle mirabilis .................. ......................................... 95 

Family Chauhaneidae 

Pseudochauhanea sphyraenae ................ ..................................... 96 

Family Gastrocotylidae 

Pseudaxine mexicana ................................... ............................... 96 

Family Neothorcocotylidae 

Neothoracocotyle acanthocybii .................................................... 97 

Scomberocotyle scomberomori ..................... .............................. 97 

Thoracocotyle crocea ................................................................... 98 

Family Gotocotylidae 

Gotocotyla acanthophallus ........................................................... 99 

Family Mazocraeidae 

Grubea cochlear .......................................................................... 100 

Kuhnia scombercolias ................................................................. 100 

Kuhnia scombri ............................................ ............................... 101 

Miscellaneous Gillworms ............................................................ 102 

Capsalidea - Capsalids 

These relatively large, broad and flat gillworms attach to the host with a 

large, cup-shaped haptor on the posterior and 2 smaller suckers on the anterior 

of the worm. A microscopic pair of anchors are found on the haptor. Larger 

adult worms are easily visible on the host and small or developing worms can 

be found in wet mounts of skin scrapings or clippings of gills observed with a 

compound microscope. Absence in samples does not assure that fishes are free 

from these parasites. They attach to the skin, gills and inside the nares and 

mouth of big game fishes. Sometimes there may be so many worms on the skin 

that they appear to be the scales of the fish. They can be relaxed by freezing in 

sea water first, and fixed in 5% formalin. 

Our examinations of adult wild fishes suggest that these worms usually do 

little physical damage. They can become a serious problem in hatcheries or netpen 

culture, where fishes are crowded together, water quality is poor and water 

exchange rates are low, enhancing the accumulation of worms. In heavy infections, 

attachment causes skin or gill irritation and production of mucus. Fishes 

81

82 


may exhibit flashing behavior and scrape their bodies on the sides of the tank 

or pen. Bacteria can enter damaged areas, further weakening the fish. 

In netpen culture, plastic sheeting can be used to isolate individual pens for 

treatments using formalin. This is extremely labor intensive and time 

consuming. When the plastic is removed, fishes again become exposed to 

reinfection of parasites from wild sources. Increasing the water flow rate in 

hatchery situations may flush out early stages of gillworms in the water and slow 

their accumulation on the gills. It should be noted that no treatment will 

eliminate all parasites, some will always remain. Treatments reduce the 

numbers to tolerable loads for fishes. Local stocks should be developed for any 

marine fish enhancement programs rather than importing fishes to new areas 

where they may be more susceptible to local parasites or introduce new parasites 

to local wild stocks. 

Only large gillworms (Order Capsalidea and Order Mazocraeidea) occur on 

big game fishes. Smaller gillworms (Order Dactylogyridea and Order Gyrodactylidea) 

dominate (in number of fish species infected and number of individual 

worms per host) in most other fishes and in most other habitats. Certainly 

larger fishes might be expected to have larger parasites, but this does not explain 

the total exclusion of these otherwise omnipresent forms. Some capsalids are 

less host specific than is usually found in the majority of gillworms which normally 

occur on one species or genus of host. Capsalids often occur on many 

species of hosts in one or more families of offshore game fishes. This ability 

is of great advantage in exploiting such highly dispersed and fast swimming 

fishes. A similar situation is found in some of the copepods parasitizing these 

fishes, obvious examples of parallel evolution. These worms probably do little 

harm but in large numbers they could cause the fish to stop feeding and thus 

become less susceptible to hook and line fishing. 

Caballerocotyla manteri (Price) 

The relatively large suckers on the anterior 

end of this minuscule, and little known, worm 

look like a pair of spectacles. It occasionally 

occurs in low numbers on little tunny and wahoo. 

Name - It was originally placed in Capsala. 

Diagnostic Characters - This worm is elongate 

in body outline and the sucker-like haptor extends 

beyond the body outline. The cephalic suckers are 

large. The testes are all inside the cecal loop. 

Records - One occurred in a wahoo off La Parguera, 

Puerto Rico (USNPC 82585). Three were 

found in a little tunny from the Dry Tortugas, 

Florida, USA (USNPC 37228-9); and on this host 

from Chesapeake Bay and Woods Hole, Massachusetts, USA. 

Geographic Range - Western Atlantic (not confirmed form the Atlantic coast 

of South America). Our collection is the first from the Caribbean.


Life History - Justine, Lambert and Mattei (1985) described the shape and 

form of the spermatozoa, and Justine and Mattei (1987) spermatogenesis. 

Location in Host - Gills. It occurred on the tongue of wahoo. 

Length - 2.1-2.6 mm. 

Host Specificity - It is probably host specific to little tunny and wahoo is a 

false host. It had not previously been reported from wahoo. 

Capsaloides cornutus (Verrill) 

This little known, worm occurs in the mouth of 

longbill spearfish and white marlin. 

Diagnostic Characters - This oval worm has a 

sucker-like haptor divided into 7 large and 2 small, 

shallow depressions around the border and 1 in the 

middle. The haptor diameter is about 1/5 the length of 

the body of this parasite and only slightly extends past 

the body outline. It is pink when alive. 

Records - Four occurred in a longbill spearfish from 

Aguadilla, Puerto Rico (USNPC). Light infections of 

this parasite have also been found in white marlin from 

Woods Hole, Massachusetts (USNPC 7178) and Block 

Island, Rhode Island, USA (USNPC 35136). 

Geographic Range - Unknown. Our collection is 

the first from the Caribbean. 

Associations - This parasite and Tristomella laevis were found in the mouth 

and on the gills of the longbill spearfish and Capsaloides magnaspinosus 

occurred in the nares of the same fish. Possibly these 3 parasites also co-exist 

in white marlin, but this requires confirmation. 

Location in Host - Gills and mouth. 

Length - 5.3-8.0 mm (white marlin); 4.3-5.5 mm (longbill spearfish). 

Host Specificity - This parasite may be genus specific (Tetrapturus). Longbill 

spearfish is a new host for this worm, but this is not surprising since the fish has 

not previously been examined for parasites. 

Interestingly, this parasite was not found in 3 

white marlin we examined in Puerto Rico, or 

in 2 from Alabama, USA. 

Capsaloides magnaspinosus Price 

This wrinkled worm has rarely been seen 

because the nares of billfishes are seldom 

examined. 

Diagnostic Characters - This tear-drop or 

pear-shaped (pyriform) worm has a sucker-like 

haptor divided into 7 large and 2 small shallow 

depressions around the border and 1 in the 

middle. The diameter of the haptor is 1/3 to 

1/2 the length of the body of this parasite and 

83

84 


is entirely within the body outline. The margins of the body are scalloped or 

crenelated and the worm is much thicker than C. cornutus. It is also pink in 

life. 

Records - Three occurred in a longbill spearfish from Aguadilla, Puerto Rico 

(USNPC); and 3 in a white marlin from Woods Hole, Massachusetts, USA 

(USNPC 35648). 

Geographic Range - Unknown. Our record is the first from the Caribbean. 

Associations - See previous parasite. 

Location in Host - Nares. 

Length - 5.4-6.6 mm (white marlin); 4.3-5.5 mm (longbill spearfish). 

Host Specificity - This parasite may be genus specific (Tetrapterus). Longbill 

spearfish is a new host for this parasite. Interestingly, this parasite was not 

found in 3 white marlin we examined in Puerto Rico, or in 2 from Alabama, 

USA. 

Nasicola klawei (Stunkard) 

This round nasal worm is consistently 

found in yellowfin tunas and never in the 

similar bigeye. It may be used to distinguish 

these similar fishes. 

Name - The genus name refers to the location 

in the nares. It has also been placed in genus 

Caballerocotyla. Tristoma sp. of Rossignol and 

Repelin, 1962 is a synonym; and Capsala 

thynni (Guiart) is probably a synonym. 

Diagnostic Characters - This large, almost 

circular worm has a relatively small, sucker-like 

haptor divided into 7 relatively large 

depressions around the border and 1 in the 

middle. The haptor does not extend beyond the body outline. The anterior 

suckers are reduced and the pharynx is constricted in the middle. 

Records - We found 4 in each of 2 yellowfin tuna from off La Parguera and 

Desecheo Island; and 4 in 1 blackfin tuna from Boqueron, but not in 1 from 

Aguadilla, Puerto Rico; and 1-7 in 2 yellowfin tuna from Dauphin Island, 

Alabama, USA. This worm has previously been reported in blackfin tuna and 

yellowfin tuna from Puerto Rico; and yellowfin tuna from the Bahamas, 

Venezuela; very light to light infections in albacore and yellowfin tuna from the 

Atlantic and Gulf coasts of the USA; the eastern Atlantic and from widely 

separated localities in the Pacific (USNPC 59865). 


Location in Host - Nasal capsules. From 1-5 worms were found in each capsule, 

but usually 2 were present. It has rarely been reported from the gills or 

body where the worms were probably washed from the nares after the host died. 

Length - 7.5-11.9 mm (yellowfin tuna); 8.2-10.9 mm (blackfin tuna). Bane 

(1969) reported the average and maximum lengths of 10.4 and 11.9 mm in 11


worms from yellowfin tuna (4 from Puerto Rico), and 9.6 and 10.9 mm in 30 

blackfin tuna (from Aguadilla, Puerto Rico). He suggested that the difference 

in worm size was because blackfin tuna are smaller than yellowfin tuna and 

therefore the nasal capsules were smaller, restricting the growth of the worms 

inside. He also noted a correlation between the length of each worm and the 

length of its fish host. 

Host Specificity - This worm is a characteristic parasite of yellowfin tuna. It 

appears to occur less often in blackfin tuna. 

Significance to Sport Fishing - Bane (1969) suggested that since this worm 

always occurs in yellowfin tuna and never in bigeye tuna, the parasite could be 

used to distinguish these morphologically similar species of tuna. 

Tristoma coccineum Cuvier 

This parasite is more numerous on western 

Atlantic stocks of swordfish than the smaller T. 

integrum. The ratio of numbers of these 2 parasites 

reverses from the western to eastern Atlantic 

and may be used to identify these stocks. 

Name - Tristoma aculeatum Grube and T. 

papillosum Diesing are synonyms. 

Diagnostic Characters - This almost circular 

parasite has a sucker-like haptor divided into 7 

large shallow depressions around the border and 

1 in the middle. The haptor does not reach the 

posterior margin of the body. The numerous 

testes are confined to the intercecal area. The 

margins of the body have rows of minute spines radiating outward. If these 

rows are counted on one side, there are 43-54. This parasite is larger than the 

similar T. integrum. 

Records - One to 79 (average 12.6) occurred in 264 of 303 swordfish (ARC 

2322), and 233 (average 12.3) in 19 of 24 from the northwest Atlantic. It has 

also been found on swordfish from Woods Hole, Massachusetts (USNPC 4877, 

7168, 35124, 35645-7), the Atlantic coast of Canada to Chesapeake Bay and the 

Mediterranean. 

Geographic Range - Atlantic and Mediterranean. 

Associations - see T. integrum. 

Location in Host - Gills and gill cavity. 

Length - 10.0-16.4 mm. 

Host Specificity - This worm is only found on swordfish. It is a primary 

parasite of this host in the western Atlantic, but a secondary parasite in the 

eastern Atlantic. A record from smooth hammerhead, Sphyrna zygaena 

(Linnaeus), was probably accidental or erroneous. 

Preparation for Study - Iles (1971) suggested that "deep frozen" worms do 

not contract and match the sizes reported in the literature. Those killed in 

formalin were 40% smaller due to contraction and distortion. 

85

86 


Significance to Sport Fishing - Iles (1971) reported that in the northwest 

Atlantic the abundance of this species is much higher than the similar T. integrum, 

but the opposite ratio of these 2 worms occurs on Mediterranean 

swordfish. She suggested that this relative abundance might be used as a 

biological tag to distinguish stocks of swordfish. 

Tristoma integrum Diesing 

This parasite is more numerous on 

eastern Atlantic stocks of swordfish 

than the larger T. coccineum. The ratio 

of numbers of these 2 parasites 

reverses from the western to eastern 

Atlantic and may be used to identify 

these stocks. 

Diagnostic Characters - This worm is 

similar to T. coccineum, but is smaller. 

The margins of the body have rows of 

minute spines radiating outward. If 

these rows are counted on one side, 

there are more than 300. 

Records - One to 12 (average 3.0) 

occurred in 134 of 303 swordfish (ARC 2321), and 37 (average 3.7) in 10 of 24 

from the northwest Atlantic. It has also been found on swordfish from the east 

coast of the USA and the Mediterranean. 

Geographic Range - Atlantic and Mediterranean. 

Associations - Tristoma coccineum occurred in 19, T. integrum in 10, and both 

worms occurred together in 7 of 24 swordfish from the northwest Atlantic. 

Location in Host - Gills, between gill filaments. 

Length - 5.8-12.0 mm. 

Host Specificity - This worm is only found on swordfish and is a primary 

parasite in the eastern Atlantic, but a secondary parasite in the western Atlantic. 

Significance to Sport Fishing - See previous parasite. 

Tristomella laevis (Verrill) 

This worm attaches on any external 

surface of billfishes around the world. 

Name - This parasite has also been placed 

in the genera Capsala and Tristoma. Tristomum 

poeyi Vigueras is a synonym. It was 

redescribed by Price (1938). 

Diagnostic Characters - This parasite is 

circular in outline. The haptor is on the 

posterior portion of the body and extends 

past the body outline. The testes are 

numerous and occur outside and inside the


boundary of the ceca. The vitellaria extend nearly to the margin of the body. 

Records - Six to more than 100 occurred on 40 Atlantic blue marlin from off 

various locations around Puerto Rico (USNPC 81999-82003), 25-100 on 4 of 5 

Atlantic sailfish from Arecibo, 55-70 on 2 of 3 white marlin from La Parguera, 

and 3 on a longbill spearfish from Aguadilla, Puerto Rico. It has also been 

found on white marlin from Havana, Cuba; and Block Island, Rhode Island, 

USA (USNPC 7179); and billfishes from the eastern and southern Atlantic and 

Indo-Pacific. 

We have observed very heavy infections on freshly caught blue marlin, but 

worms gradually fell off or were knocked off in handling, and were reduced to 

moderate infections by the time the fish was examined at the dock. The 

parasites of most big game fishes may be lost or reduced in numbers during 

handling of the host, but this worm seems to be particularly vulnerable. Thus 

the numbers per host reported may actually represent a small percentage of what 

was on the live host. 


Associations - This parasite was found in the mouth and on the gills of a longbill 

spearfish along with Capsaloides cornutus. Capsaloides magnaspinosus also 

occurred in the nares of this fish. These 3 parasites are also found on the white 

marlin and may co-exist on that host. Tristomella laevis was found alone on 

Atlantic blue marlin we examined. 

Location in Host - Body, gills and inside mouth. This worm is either a real 

generalist in choosing attachment positions, or it moves or is washed into 

different locations when the host is handled. Examination of freshly caught fish 

by fishermen could explain this little mystery. 

Length - 7.75-11.5 mm. Puerto Rican specimens were smaller (5.4-10.0 mm). 

Host Specificity - This worm is a characteristic parasite of Atlantic blue 

marlin. It is at least a primary parasite of Atlantic sailfish and white marlin. 

This gillworm may be family specific to billfishes. Records from Atlantic and 

Indo-Pacific swordfish, and Brazilian dolphin, and northwest Atlantic skipjack 

tuna, appear to have been cases of accidental parasitism, erroneous 

identifications, or contamination from billfishes. Atlantic blue marlin was 

reported as a new host by Dyer, Williams and Bunkley-Williams (1992). 

Significance to Sport Fishing - This is probably one of the most familiar 

parasites for big game fishermen. It appears as soft, translucent-white, scalelike 

attachments to the outside of billfishes. They are most numerous when the 

fish is first boated but begin to fall off almost immediately. They may litter the 

deck and wiggle among the mucus and blood. Many remain attached and some 

move inside the gill covers to avoid desiccation. Probably, they do little harm 

to the fish but very heavy infections may cause skin irritation. 

87

88 


Dionchus agassizi Goto 

This worm occurs in cobia and remoras around the world. 

Its presence suggests that these fishes are related. 

Name - Dionchus rachycentris Hargis was described in August, 

1955; and D. hopkinsi Koratha in September, 1955, from cobia 

in the northern Gulf of Mexico (USNPC 54754). Both are 

synonyms of this worm. It was redescribed by Price (1938). 

Diagnostic Characters - The haptor of this elongate parasite is 

formed into a sucker and has 2 stout, centrally placed anchors. The 

anterior portion of the worm is triangular in shape and has 

numerous cephalic glands that open along the lateral margins. This 

worm differs from the similar D. remorae because the 2 anchors on 

the haptor are larger than 1/4 the diameter of the haptor; and the 

blade of these anchors is shorter than their base. 

Records - Two to 4 occurred in 3 cobia from Dauphin Island, 

Alabama; in light infections from Tampa Bay, Florida; Grande 

Isla, Louisiana; 8 worms in 4 of 9 cobia from Port Aransas, Texas, USA; and in 

this host from Australia. This worm also occurs on inshore remora, shark 

remora and spearfish remora around the world. 


Associations - This worm and D. remorae occur on the gills of inshore remora 

together. The association of this worm with remoras and cobias suggests these 

2 families are either related or the same (see Discussion). 

Location in Host - Gill filaments. 

Length - 3.0-5.1 mm (cobia); 0.8-3.7 mm (remoras). Cobia grow to a larger 

size than remoras, and larger worms on larger hosts is not surprising. The size 

of many parasites is directly related to the size of their host. 

Host Specificity - This worm is probably a primary parasite of cobia, inshore 

remora, shark remora and spearfish remora. 

Mazocraeidea 

These worms have more complex haptors than capsalids. They can usually 

be easily seen with the naked eye. These worms have intricate attachment 

organs with a series of complicated clamps or suckers often on extensions of a 

complex haptor. They generally feed on blood. They produce few eggs and 

usually occur in low numbers on the host. They are largely marine. The few 

that do occur in fresh water are usually on hosts of marine origin. They do not 

increase in numbers rapidly, but since they feed on blood, they can severely 

damage their hosts with even slight increases in numbers. Usually big game 

fishes are not held in hatchery or culture conditions where worms can 

accumulate and cause problems. They could become a problem if culture for 

restocking of marine sport fish, as is being studied in Florida, is attempted. 

These worms can be relaxed in dilute formalin solutions (1 part formalin to 

4000 parts water) and cleared and mounted in glycerine jelly. Alternatively,


they may be placed under a microscope coverslip on a slide and bathed in 5% 

formalin so that they are fixed flat. They can be stained and permanently 

mounted as described for flukes. Fishes infected with these worms can be 

treated with formalin as described for capsalids above. 

Allopseudaxine katsuwonis (Ishii) 

This distinctively triangular gillworm occurs on the 

gills of skipjack tuna around the world. 

Diagnostic Characters - This worm is narrow anteriorly 

and broadens posteriorly. It has a short but broad haptor, 

slightly constricted from the body of the worm, and 24-27 

clamps along the posterior border. It has 60-79 testes. 

In the Indo-Pacific, Allopseudaxinoides vagans (Ishii) 

shares the gills of skipjack tuna with A. katsuwonis and 

may possibly be found in the Atlantic. The name A. 

vagans is from a former name (synonym) of skipjack 

tuna, Katsuwonus vagans. It is similar in size (6.0-7.0 

mm) and general shape and organ arrangement, but 

differs from A. katsuwonis by having 13-15 clamps, and 

30-35 testes. The clamps are also more complex 

possessing accessory sclerites. 

Records - One occurred in 1 of 3 skipjack tuna from Arecibo, Puerto Rico 

(USNPC). It has also been found on this host in the north central and 

northwest Pacific. Lester (pers. comm.) found what was probably this worm 

and A. vagans on skipjack tuna from the southwest Pacific. 

Geographic Range - Worldwide. Our collection is the first record of this 

parasite from the Atlantic Ocean. 


Length - 7.0-8.0 mm. 

Host Specificity - This worm only occurs on skipjack tuna 

and is probably a characteristic parasite of this host. 

Allencotyla mcintoshi Price 

This is a little-known gillworm on the well-known 

greater amberjack. 

Name - It was once placed in genus Heteraxine. 

Diagnostic Characters - The haptor is asymmetrical with 

a longer row of larger clamps and a shorter row of smaller 

ones. The space between the lateral rows of dark vitellaria 

(containing internal organs), is dumbbell-shaped with large 

numbers of eggs above and large numbers of testes below. 

Records - Five occurred in greater amberjack off Miami, 

Florida, USA (USNPC 37730-1). 



89

90 

Length - 8.0-10.0 mm. 



Cemocotyle carangis (MacCallum) 

This elongate gillworm was thought to only occur in blue 

runner, but we have seen it on 3 other species of jacks. 

Name - Cemocotyle borinquenensis is a synonym described 

from Puerto Rico by Price (1962) from a fish he identified as 

green jack, Caranx caballus Gunther, a Pacific fish which 

does not exist in Puerto Rico. He also redescribed C. carangis. 

Diagnostic Characters - The anterior end of this parasite is 

wider than the "neck" and has 2 sucker-like organs in the 

lateral margins. The clamps on the haptor are in 2 rows, a 

short row with 11-14 clamps and a long row with 36 clamps. 

The genital opening is surrounded by spines. 

Records - We found 1 in 3 of 15 bar jack; 1 in a blue runner; 

1-15 in 5 of 9 crevalle jack (USNPC 82581, 85935); and 2 in 

1 of 17 horse-eye jack from various localities around Puerto 

Rico; and 10 in a crevalle jack from Dauphin Island, Alabama, 

USA. Two occurred in a blue runner from Woods Hole, 

Massachusetts (USNPC 35170,37735); 4-13 in 2 from the New 

York Aquarium (USNPC 37737); 68 from Alligator Harbor, 

Florida, USA; and from Mexico. 

Geographic Range - Western Atlantic. Our collections are the first in the 

insular Caribbean. 

Life History - This worm was not in 22 immature crevalle jack (3-6 cm long) 

from 4 localities near Dauphin Island, Alabama. It may only parasitize adults. 

Associations - Four species of gillworms occur together on jacks of the genus 

C aranx. The intensities of infection that we found on these fish are as follows: 


Length - 3.7-12.0 mm.


Host Specificity - This worm was thought to only occur in blue runner, but 

we have seen it on 3 additional jack species. It is genus specific (Caranx), and 

is a primary parasite of blue runner. In Puerto Rico it is also a primary parasite 

of crevalle jack and a secondary parasite of bar jack and horse-eye jack. Bar 

jack, crevalle jack and horse-eye jack are new hosts. An immature worm in a 

pompano, Trachinotus carolinus (Linnaeus), from the New York Aquarium was 

probably an accidental infection (USNPC 37742). 

Cemocotyle noveboracensis (Price) 

This parasite was thought to occur only in crevalle jack, but 

also occurs in horse-eye jack in Puerto Rico. 

Name - It was redescribed by Price (1962). 

Diagnostic Characters - This worm is shaped like an upright 

vacuum cleaner. It is similar to C. carangis but differs by having 

a larger number of clamps on each side of the haptor (43-57 and 

15-17). Slightly more clamps were found on the long side in local 

specimens than had been reported previously. 

Records - One to 6 occurred in 6 of 9 crevalle jack (USNPC 

84688), and 1-4 in 8 of 17 horse-eye jack from localities in Puerto 

Rico. One to 5 worms were on 14 of 96 adult and 1-15 on 29 of 

50 subadult crevalle jack from Venezuela; on crevalle jack from 

New York (USNPC 37738-41) and Florida, USA, and Mexico. 

Geographic Range - Western Atlantic (not confirmed from the 

Atlantic coast of South America). Our collections are the first in 

the insular Caribbean. 

Life History - This worm did not occur in 22 immature crevalle 

jack (3-6 cm long) from 4 localities around Dauphin Island, 

Alabama; or in 10 immature horse-eye jack (3-7 cm long) from 3 

localities around Puerto Rico. It may only parasitize adults. 

Associations - See C. carangis. This worm occurred with Protomicrocotyle 

mirabilis, Allopyragraphorus incomparabilis and Cemocotylella elongata in 

crevalle jack from Venezuela, but its numbers were only closely correlated with 

those of C.elongata (Bashirullah and Rodriguez 1992). 


Length - Up to 4.5 mm. 

Host Specificity - This worm was previously thought to only occur in crevalle 

jack, but we found it in this host and horse-eye jack from Puerto Rico (Bunkley- 

Williams and Williams 1995). It is genus specific (Caranx), a primary parasite 

of crevalle jack, and a secondary parasite of horse-eye jack. 

Cemocotylella elongata (Meserve) 

This worm has been found in crevalle jack from Venezuela and another jack 

from the Pacific coast of Panama. 

Diagnostic characters - This parasite is similar to the 2 species of Cemocotyle 

described above. It differs by having fewer clamps on each side of the haptor 

91

92 


(24-25 clamps on one side and 4-5 on the other) and by lacking 

spines around the genital opening. 

Records - One to 23 worms occurred on 57 of 96 adult, and 1- 

8 on 18 of 50 subadult, crevalle jacks from the east coast of 

Venezuela. It was described from a Pacific jack off Panama. 


Associations - This worm occurred with Protomicrocotyle 

mirabilis, Allopyragraphorus incomparabilis and Cemocotyle 

noveboracensis in crevalle jack from Venezuela, and its 

numbers were closely correlated with those of A. incomparabilis 

and C. noveboracensis (Bashirullah and Rodriguez 1992). 


Length - 2.1-3.7 mm. 

Host Specificity - It is a primary parasite of crevalle jack in 

the southern Caribbean. 

Helixaxine winteri Caballero and Bravo-Hollis 

This worm has both an unusual shape and 

distribution. Its distinctive haptoral clamps are situated 

on peduncles of tissue, and it has only been found from 

2 Mexican states. 

Diagnostic Characters - This small worm has a haptor 

that is curled and is nearly as long as the length of the 

worm. The clamps on the haptor are elongate in shape 

and are situated on small peduncles of tissue. 

Records - Six occurred in an unstated number of horseeye 

jack from Tuxpan, Veracruz and 13 in an unstated 

number of crevalle jack from Campeche, Campeche, 

Mexico. 


Ecology - Fewer jacks were collected from Campeche, 

but they had more worms. The infection levels could 

have been due to differences in habitat or season (June 

in Campeche, November in Tuxpan). 


Length - 1.4-2.1 mm. 

Host Specificity - Only known from crevalle jack.


Allopyragraphorus incomparabilis (MacCallum) 

This worm has a distinctive fish-tail-shaped 

haptor. It is found in the gills of jacks throughout 

the western Atlantic. 

Name - Allopyragraphorus hippos (Hargis) only 

differs from A. incomparabilis in body size and 

extent of vitellaria. The smaller worms (2.9-3.3 

mm) on which the description was based were 

probably less mature and had less developed 

vitellaria. We consider these 2 species to be the 

same. It was redescribed by Hargis (1957). 

Diagnostic Characters - It has an S-curled, 

wedge-shaped (or fish-tail-shaped) haptor with 62- 

89 pairs of attachment clamps arranged in a 

double row. The clamps are not located on stalks. 

There are 56-75 testes. 

Records - One to 5 occurred in 10 of 15 bar jacks 

from various localities around Puerto Rico 

(USNPC 85297, 85299), 1 in a bar jack from 

Desecheo Island, and 1-3 in 4 bar jack from Mona 

Island; 3 in a blue runner from La Parguera (USNPC 85942), and 1 in 1 of 2 

blue runner from Aguadilla; 1-4 in 6 of 9 crevalle jack (USNPC 82580); 1 in 

7 of 17 horse-eye jack from various localities around Puerto Rico. From 1-26 

worms were found on 66 of 96 adult and 1-11 on 23 of 50 subadult crevalle 

jacks from the east coast of Venezuela. Five occurred in a bar jack from the 

New York Aquarium; this fish had been collected from Key West, Florida 

(USNPC 36528); 13 in 7 crevalle jack from Alligator Harbor, Florida, and 10 

in 2 of 10 crevalle jack from Port Aransas, Texas, USA. 


Life History - This worm did not occur in 22 immature crevalle jack (3-6 cm 

long) from 4 localities around Dauphin Island, Alabama; in 10 immature horseeye 

jack (3-7 cm long) from 3 localities around Puerto Rico; or in 78 juvenile 

crevalle jack (fork length 4.0-11.0 cm) in Venezuela. It may only parasitize 

adults and subadults. Bashirullah and Rodriguez (1992) suggested that juvenile 

crevalle jack are found in inshore, shallow areas isolated from the adult jacks 

infected with gillworms. 

Associations - See Cemocotyle carangis and C. noveboracensis. It almost 

always occurred with Cemocotylella elongata on Venezuelan crevalle jack. 


Length - 2.9-5.0 mm. 

Host Specificity - This worm is a characteristic parasite of crevalle jack and 

horse-eye jack. Too few collections are available to determine its status on 

other jacks. This parasite is genus specific (Caranx). Blue runner is a new host 

for this parasite. Bunkley-Williams and Williams (1995) reported horse-eye jack 

as a new host. 

93

94 


Hexostoma euthynni Meserve 

This worm parasitizes the gills of 3 species of little tunas 


Name - This worm has also been placed in genus Neohexostoma. 

H. macracanthum Fujii, H. pricei Koratha, and N. 

kawakawa Yamaguti are synonyms. It was redescribed by 

Millemann (1956). 

Diagnostic Characters - This elongate worm is narrow at the 

anterior end and has a narrowed "waist" near the anterior 1/3 

of the body. It has a haptor with 8 clamps, all equal in size, 

arranged in 2 rows along the lateral borders. The clamps have 

a distinctive X-shaped sclerite in the middle. Two pair of larval 

anchors can be seen on the posterior end between the rows of 

clamps. Rohde (1980) described the ceca of this worm. 

Records - We found 1-2 in 3 of 6 little tunny from La 

Parguera, Puerto Rico (USNPC 82598); and 1 in a little tunny 

from Dauphin Island, Alabama. Two occurred in a little tunny 

from the Dry Tortugas off Florida (USNPC 36890), 3 in 1 from 

Louisiana, and from Chesapeake Bay; 4 in 2 of 10 Atlantic bonito from Port 

Aransas, Texas (USNPC 54761), USA. This worm has often been reported in 

little tunnies (Euthynnus spp.) from various localities in the Pacific including 

Hawaii, USA, (USNPC 63671, SY No. 172) and the Great Barrier Reef, 

Australia (USNPC 74151); and 4 in a yellowfin tuna from Hawaii. 

Geographic Range - Worldwide. Our collections are the first in the Caribbean. 


Length - 3.4-8.4 mm. 

Host Specificity - This worm is almost genus specific (Euthynnus), and occurs 

in all 3 species in the genus (little tunny; kawakawa, 

Euthynnus affinis (Cantor); and black skipjack, Euthynnus 

lineatus Kishinouye). The single records of light infections 

in an Atlantic bonito and yellowfin tuna may have been 

accidental. 

Hexostoma lintoni Price 

This is a mysterious worm of Atlantic bonito. 

Name - Linton (1901, 1940) called this worm Hexacotyle 

thynni Delaroche [now Hexostoma thynni (Delaroche)], and 

deposited 1 specimen (USNPC 9641). Price (1961) 

described the deposited specimen as a new species. 

Diagnostic Characters - This elongate worm has a narrow 

anterior end. The haptor has 4 pairs of clamps arranged in 

a row along the posterior border. The middle 2 clamps are 

about 1/2 as large as the other 6 clamps. The clamps have 

a distinctive X-shaped sclerite in the middle of each.


Records - One each occurred in 2 Atlantic bonito from Woods Hole, 

Massachusetts, USA (USNPC 9641). 


Location in Host - Mouth. Other species in this genus occur in the gills. This 

odd location also makes the validity of the worm questionable. Atlantic bonito 

could be a false host in which a worm was dislodged from a prey fish and 

attached in the mouth of this predator. 


Host Specificity - This worm has only been found in Atlantic bonito, but so 

rarely that it could be an accidental parasite. 

Detection - Please look in the mouth of Atlantic bonito for this tiny to small 

worm, any collections will help resolve this mystery. 

Protomicrocotyle mirabilis (MacCallum) 

This asymmetrical worm is found from Chesapeake Bay to the 

Caribbean. 

Name - This worm was redescribed by Koratha (1955b) and by 

Caballero and Bravo-Hollis (1965). 

Diagnostic Characters - It has an elongate body with 4 small 

clamps which appear to be on the side of the body anterior of the 

broad haptor. The larval anchors are widely separated and located 

at the posterior border of the haptor. 

Records - One to 3 occurred in 6 of 15 bar jack (USNPC 85298); 

1-6 in 8 of 9 crevalle jack (USNPC 82534, 82663); and 1-3 in 11 

of 17 horse-eye jack from various localities around Puerto Rico. 

One to 111 were found on 94 of 96 adult, and 1-39 in 48 of 50 

subadult, crevalle jacks from the east coast of Venezuela. It has 

also been found in crevalle jack from Alligator Harbor, Florida; 

Texas; Chesapeake Bay, USA; and the west coast of Africa; and 

in horse-eye jack from Mexico. 

Geographic Range - Atlantic. 

Life History - This worm did not occur in 22 immature crevalle 

jack (3-6 cm long) from 4 localities around Dauphin Island, 

Alabama; or in 10 immature horse-eye jack (3-7 cm long) from 3 

localities around Puerto Rico. It may only parasitize adults. 

Associations - See Cemocotyle carangis and C. noveboracensis. 


Size - Up to 5.5 mm long. 

Host Specificity - This worm is genus specific (Caranx), and a characteristic 

parasite of crevalle jack. It is also a primary parasite of horse-eye jack, and a 

secondary parasite of bar jack. 

95

96 


Pseudochauhanea sphyraenae Yamaguti 

This arrow-shaped parasite occurs in light infections on 

the gills of great barracuda around the world. 

Diagnostic Characters - The body is elongate, narrow 

anteriorly, and abruptly wider at level of vagina. The 

haptor is V-shaped with unequal rows of clamps, 28-50 on 

one side 27-37 on the other. No larval anchors are present on 

the haptor. 

Records - We found 6 in a great barracuda from Mona 

Island; 3-6 in 23 of 33 from various localities around 

Puerto Rico (USNPC 86632); 1 in 1 from Great Inagua, 

Bahamas; and 3 in 1 of 2 from Dauphin Island, Alabama, 

USA (USNPC). It has also been reported from Key 

Biscayne, Florida, and Hawaii, USA (USNPC SY No. 16). 


first records in the Caribbean and the Gulf of Mexico. 

Associations - It occurred on the gills of great barracuda 

in Hawaii with Vallisiopsis sphyraenae Yamaguti, but this 

second worm has not been found in the Atlantic. 


Length - 4.4-10.0 mm. 

Host Specificity - This worm is only found in great 

barracuda, and it is a primary parasite of this host. 

Pseudaxine mexicana Meserve 

This parasite, with a sombrero-shaped haptor, infects 

Spanish mackerels. 

Name - The broad, wide haptor looks a bit like a Mexican 

sombrero. This shape plus a distribution largely on both 

sides of Mexico makes the name, "mexicana", appropriate. 

Pseudaxine texana Koratha is a synonym. It has also been 

placed in genus Mexicotyle. 

Diagnostic Characters - It has a distinctive haptor with 

the simple anchors of the larva still present on one side and 

37-56 clamps along the posterior border. The clamps 

sometimes appeared to curl around and form a double row 

on the side opposite the simple anchors. 

Records - We found 1 each in 7, and 2-4 in 2 of 35 cero 

from La Parguera (USNPC 82660); 1-15 in 9 of 14 king 

mackerel from various localities around Puerto Rico; and 

9-20 in 2 of 4 king mackerel from Dauphin Island, 

Alabama, USA. This worm occurred in a king mackerel from Puerto de 

Veracruz, Veracruz, Mexico. It has been reported in Spanish mackerel from 

Florida, Louisiana, Texas (USNPC 54758), Chesapeake Bay, USA, and in other 

Spanish mackerels from the Pacific coast of Mexico.


Geographic Range - Western North Atlantic and the eastern Pacific. Our 

collections are the first in the Caribbean. 

Associations - We found 1-4 P. mexicana with 1-11 Thoracocotyle crocea and 

1-4 Gotocotyla acanthophallus in 3 cero; and 1 with 1 G. acanthophallus in 1 

cero from Puerto Rico. Fifteen and 6 occurred with 6 and 25 G. acanthophallus 

and 1 and 2 Scomberocotyle scomberomori in 2 king mackerel; and 1-10 with 

2-19 G. acanthophallus in 7 other king mackerel from Puerto Rico. Nine were 

associated with 18 G. acanthophallus and 1 S. scomberomori in a king mackerel 

from Alabama. On Spanish mackerels in Puerto Rico, P. mexicana and G. 

acanthophallus always occurred together when more than one worm was found. 

In 9 cero there was only a single worm. 


Length - 1.9-5.5 mm; 3.0-5.0 mm (Puerto Rico); 3.8-5.5 mm (Alabama); 3.5- 

3.9 mm (Mexico). 

Host Specificity - This worm is genus specific (Scomberomorus). It is a 

primary parasite of king mackerel and Spanish mackerel, but a secondary 

parasite of cero. 

Neothoracocotyle acanthocybii (Meserve) 

This gillworm with wrap-around clamps is found on wahoo around 

the world. The consistently heavy infections may damage this 

important sport fish. 

Name - This worm was redescribed by Yamaguti (1968). 

Diagnostic Characters - The haptor is wrapped around the elongate 

body. The 223-245 clamps occur in approximately equal rows. 

Records - We found 10-60 in 15 wahoo from various localities around 

Puerto Rico (USNPC 82582); and 50-100 in 4 wahoo from Dauphin 

Island, Alabama, USA. It also occurs in this host from Chesapeake 

Bay, USA, and the Pacific. 

Geographic Range - Worldwide. Our collections are the first in the 

Caribbean and the Gulf of Mexico. 

Associations - It occurred in a wahoo with Caballerocotyla manteri. 

Location in Host - These worms are consistently found near the distal 

ends of the gill filaments of the first 2 arches. 

Length - 6.3-11.1 mm. 

Host Specificity - It is only found on wahoo, and is a characteristic 

parasite. 

Significance to Sport Fishing - A burden of 50-100 of these 

relatively large worms must damage this important sport fish. The effect of 

these consistently heavy infections should be studied. 

Scomberocotyle scomberomori (Koratha) 

This elongate worm with a fish-tail haptor very lightly infects the gills of 


Name - Originally placed in genus Heteraxine. 

97

98 


Diagnostic Characters - This elongate worm has a broad, 

fish-tail-shaped haptor with 2 uneven rows of 85 and 50 clamps. 

In the Puerto Rican worms, the long side had 84 clamps. 

Records - We found 1-2 in 2 of 14 king mackerel from 

various localities around Puerto Rico; and 1 in 1 of 4 king 

mackerel, and 1 in each in 2 of 10 Spanish mackerel from 

Dauphin Island, Alabama, USA. It has also been reported in 

king mackerel and Spanish mackerel from Alligator Harbor 

(USNPC 37494) and Tampa Bay, Florida, and in Spanish 

mackerel from Port Aransas, Texas (USNPC 54757) and 

Chesapeake Bay, USA. 

Geographic Range - Western Atlantic (not confirmed from 

the Atlantic coast of South America). Our collections are the 

first in the Caribbean. 

Associations - One and 2 S. scomberomori were found with 

15 and 6 Pseudaxine mexicana and 6 and 25 Gotocotyla 

acanthophallus in 2 king mackerel from Puerto Rico. One 

was associated with 9 P. mexicana and 18 G. acanthophallus 

in a king mackerel from Alabama. 


Length - 5.5-6.5 mm. 

Host Specificity - This worm only occurs on Spanish mackerel and king 

mackerel. It is genus specific (Scomberomorus), but occurs so infrequently and 

in low numbers that it is a secondary parasite of these hosts. 

Thoracocotyle crocea MacCallum 

This parasite is almost all haptor. It occurs, often 

sporadically, and usually in low numbers, on the gills of Spanish 

mackerels. 

Name - It was redescribed by McMahon (1964). Thoracocotyle 

paradoxica Meserve is a synonym. 

Diagnostic Characters - The haptor dominates the worm and 

looks like the skirt of a high-kicking dancer. It is actually one 

side of the body of this worm, with the 7 large testes appearing 

to be in the area of the "haptor". It has 2 rows of 17-21 clamps. 

Records - We found 1 each in 16, and 11 in 1 of 35 cero from 

various localities around Puerto Rico (USNPC 82662). It also 

occurred in Spanish mackerel and king mackerel from Florida; 

1 in 1 of 5 king mackerel from Beaufort, North Carolina; 10 in 

1 from Grand Isle Louisiana, and 90 in Spanish mackerel from 

Chesapeake Bay; the New York Aquarium, USA; and Mexico. 

Geographic Range - Western Atlantic (not confirmed from the 

Atlantic coast of South America). Our collections are the first 

in the Caribbean. 

Associations - One to 11 were found along with 1-4 Pseudaxine mexicana and


1-4 Gotocotyla acanthophallus in 3 cero from Puerto Rico. 

Ecology - This worm seems to occur sporadically, sometimes rarely and usually 

in low numbers. How this species survives under these conditions is a mystery. 


Length - 1.0-4.5 mm. 

Host Specificity - It is genus specific (Scomberomorus), but is at most only a 

secondary parasite of Spanish mackerels. 

Gotocotyla acanthophallus (MacCallum and MacCallum) 

This elongate worm with a long, V-shaped haptor 

occurs on the gills of Spanish mackerels in the 

western Atlantic. 

Name - Microcotyle scomberomori Koratha is a 

synonym. 

Diagnostic Characters - This parasite has an extended 

"V" shaped haptor with an even number of 

clamps along each arm of the "V" and 60 testes in the 

posterior half of the body. The cirrus and genital 

atrium are armed with a series of spines of similar 

lengths in an arrangement that looks like the bristles of 

a test-tube brush and is characteristic of this worm. 

Records - We found 1 each in 7, and 4 in 1, of 35 

cero from La Parguera (USNPC 82661); 2-25 in 9 of 

14 king mackerel from various localities around 

Puerto Rico (USNPC 82594, 86623); and 18-96 in 2 

of 3 king mackerel from Dauphin Island, Alabama, 

USA. Heavy infections were reported in king mackerel 

and Spanish mackerel from Florida and 

Louisiana, and light to moderate infections from Port Aransas, Texas (USNPC 

54756), and Chesapeake Bay, USA. This worm has also been reported in 

bluefish, Pomatomus saltatrix (Linnaeus), from the Atlantic coast of the USA. 

Geographic Range - Western Atlantic (not confirmed from the Atlantic coast 

of South America). Our collections are the first in the Caribbean. 

Associations - One to 4 G. acanthophallus occurred with 1-4 Pseudaxine 

mexicana and 1-11 Thoracocotyle crocea in 3 cero from Puerto Rico. Six and 

25 G. acanthophallus were found with 15 and 6 P. mexicana and 1 and 2 

Scomberocotyle scomberomori in 2 king mackerel; and 2-19 G. acanthophallus 

with 1-10 P. mexicana in 7 other king mackerel from Puerto Rico. Eighteen 

were associated with 9 P. mexicana and 1 S. scomberomori in 1, and 96 with 

20 P. mexicana in another king mackerel from Alabama. On Spanish mackerels 

in Puerto Rico, P. mexicana and G. acanthophallus always occurred together 

when more than one worm was found. In 9 cero there was only a single worm. 


Length - 3.3-7.0 mm. Published lengths were 5.5-7.0 mm, but our measure- 

ments were 3.3-6.2 mm (Puerto Rico) and 3.8-6.9 mm (Alabama). 

99

100 


Host Specificity - This worm is genus specific (Scomberomorus). It is a 

primary and possibly a characteristic parasite of cero, king mackerel and Spanish 

mackerel. Cero is a new host One recorded from a striped bass, Morone 

saxatilis (Walbaum), from the New York Fish Market, was probably accidental 

or erroneous. 

Grubea cochlear Diesing 

This oddly asymmetrical worm is found in the gills of 

Atlantic and chub mackerel. 

Name - Linton (1940) described this worm as Pleurocotyle 

scombri (Gervis and van Beneden) and it and G. pneumatophori 

Price; Grubea sp. of Loftin, 1960; Grubea sp. of Rohde, 1984; 

and Grubea sp. of Wagner, 1975, are synonyms. Mamaev (1982) 

and Rohde (1987) redescribed this worm. 

Diagnostic Characters - This is an elongate worm with 4 

clamps on one side of the haptor and a single, smaller clamp on 

the other side. The sizes of clamps and the number of genital 

hooks vary among specimens from the same host and locality. 

Records - It occurred in a blue runner from Florida, and 1 was 

found in a chub mackerel from Woods Hole, Massachusetts, USA 

(USNPC 8160). It was also found in Atlantic mackerel and 1-2 

in 7 chub mackerel from Brazil; southwest Atlantic; Atlantic off 

Europe; 1-2 in 2 Atlantic mackerel from the Mediterranean; east, 

south and southwest coasts of Africa; and the eastern Pacific. Oddly, this worm 

does not occur in Great Britain (Dawes 1968). More than 1000 Atlantic 

mackerel of various sizes, examined at different times of the year, from 

southwest England had none of these worms. It apparently occurs sporadically 

and in very light infections. 

Geographic Range - Atlantic and adjacent Indo-Pacific. Rohde (1987) 

summarized the distribution records of this worm. 

Associations - In 100 Atlantic mackerel off Europe, only 3 were infected with 

this worm, but 40 had Kuhnia scombri. 


Length - 7.5-15.0 mm. 

Host Specificity - This worm is genus specific (Scomber) to the 2 Atlantic 

hosts, but is apparently replaced by Grubea australis Rohde in the third fish 

species of the genus, slimy mackerel, Scomber australasicus (Cuvier), in the 

Indo-Pacific. The single records in blue runner and eastern Pacific bonito, 

Sarda chiliensis (Cuvier), probably represent false or accidental hosts. 

Kuhnia scombercolias Nasir and Fuentes Zambrano 

It occurs sporadically and in low numbers in mackerels worldwide. 

Name - This worm was confused with K. sprostonae until recently. Kuhnia 

arabica Mamaev and Parukhin is a synonym.


Diagnostic Characters - This is an elongate worm with 

4 small clamps along each lateral border of the relatively 

small haptor that is not clearly separated from the body. 

It is similar to K. sprostonae, but differs by having 

anchors shorter than 50 Φm and with bases shorter than 

the rest of the anchor, occurring on the gills instead of 

the pseudobranchs, and being slightly smaller. Rohde 

(1989) and Rohde and Watson (1985) studied the effect 

of geographic variation on the anchors, shape and size of 

this worm. It shows considerable variability in the 

lengths of the anchors and genital hooks. 

Records - One to 2 occurred in 2 of 11 chub mackerel 

from the Gulf of Cariaco, Venezuela; and Brazil in the 

Atlantic, and on this host and slimy mackerel from the 

Indo-Pacific. 

Geographic Range - Worldwide. Rohde (1989) summarized 

the known distribution records. 

Associations - This worm is sometimes found with K. 

scombri on the gills. Kuhnia sprostonae also occurs on 

the pseudobranchs. There are always many more K. 

scombri than K. scombercolias on the gills. 


Length - 1.1-1.4 mm. 

Host Specificity - This worm is genus specific (Scomber), and occurs in 2 of 

the 3 species. 

Kuhnia scombri (Kuhn) 

This cosmopolitan parasite of mackerels is the best studied 

gillworm in big game fishes. 

Name - This worm was redescribed by Price (1961) and Nasir 

and Fuentes-Zambrano (1983). 

Diagnostic Characters - It is an elongate worm with 4 small 

clamps along each lateral border of the relatively small haptor that 

is distinctly separate from the body. Rohde (1991) and Rohde 

and Watson (1985) studied the effect of geographic variation on 

its shape and size, and Rohde (1987) variation in clamp sclerites. 

Records - One to 2 occurred in 2 of 11 chub mackerel from the 

Gulf of Cariaco, Venezuela; and from southern Brazil. This 

worm was also found on Atlantic mackerel and chub mackerel 

from Chesapeake Bay; Newport, Rhode Island; off Cape Hatteras, 

North Carolina; and Woods Hole, Massachusetts, USA; 

Canada; the eastern and southwestern Atlantic; Mediterranean; 

and chub mackerel from the Indo-Pacific. 

Geographic Range - Worldwide. Rohde (1989) summarized the 

known distribution records. 

101

102 


Life History - Euzet (1957) described the oncomiracidium that hatches from the 

egg, Llewellyn (1956) characterized the location on the gills and the larval 

development and Finlayson (1982) reported the reproductive processes of this 

worm. 

Associations - See Grubea cochlear and K. scombercolias. 

Location in Host - Gills at the base of the filaments. Llewellyn (1956) 

studied gill microecology, and Rohde (1991) and Rohde and Watson (1985) the 

effect of geographic variation and microhabitat. 

Length - 1.3-6.6 mm. This worm becomes sexually mature at a length of 3.5 

mm, but continues to grow. 

Host Specificity - It is genus specific (Scomber), and occurs in all 3 species. 

Damage to Host - Sproston (1945) and Llewellyn (1957) described the 

attachment of clamps on the gills of the host. 

Miscellaneous Gillworms 

Dionchus remorae (MacCallum) - This worm differs from D. agassizi 

because it has 2 anchors on the haptor that are smaller than 1/5 the diameter of 

the haptor. The blade of these anchors is longer than their base. One worm 

occurred on a crevalle jack from the New York Aquarium. Light to moderate 

infections were found on inshore remora worldwide. This parasite and D. agassizi 

occur together on the gills of inshore remora. This worm is probably a 

characteristic parasite of inshore remora. The record from crevalle jack was 

probably accidental and due to the artificial environment, although it has been 

reported from jacks in the Indian Ocean. 

Kuhnia sprostonae Price - It may occur worldwide on mackerels (Scomber 

spp.), but it has not been found in the western Atlantic (see K. scombercolias). 

Neobenedenia girellae (Hargis) - This highly damaging gillworm has 

been introduced into Hawaii, Hong Kong, Okinawa and throughout the main 

islands of Japan with introductions of greater amberjack for mariculture. It was 

first thought to be N. melleni (MacCallum), a dangerous worm from the western 

Atlantic. If N. girellae is introduced to the western Atlantic, big game fishes 

could be affected. The worm apparently lacks specificity. 

Pricea minimae Chauhan - This worm was described from the gills of 

"Thynus pelamys" from Bombay, India. The host is usually thought to have 

been a skipjack tuna, but Silas (1962) suggested that this fish is almost never 

landed in Bombay, and the host must have been another scombrid.


Tristomella lintoni (Price) - This species was reported from the gills of a 

skipjack tuna caught off Woods Hole, Massachusetts, USA. It was described 

from a single immature specimen which was not in good condition (USNPC 

4878). Despite the occurrence of this worm and "Capsala laevis" in checklists 

of western Atlantic skipjack tuna, we find no valid records of capsalids on this 

fish in our area. The specimen described by Price (1960) was probably then 

result of accidental infection and its status must remain uncertain. 

Tristomella onchidiocotyle (Setti) - MacCallum collected an immature 

capsalid in the gills of a "Thunnus thunnus-horse mackerel" (=bluefin tuna) 

from Woods Hole, Massachusetts, USA (USNPC 35644). Price (1939) des- 

cribed this worm as a new species Capsala maccallumi, suggested that it was 

probably identical to C. onchidiocotyle, but listed the host as "little tunny" with- 

out explanation. The haptor of this worm extends beyond the body outline. 

Testes are numerous and extend beyond the field between the ceca. The anchors 

have a distinct hook on the end. It occurred on bluefin tuna in the Mediterranean 

and bigeye tuna off west Africa, probably on bluefin tuna off the Atlantic coast 

of the USA and throughout the Atlantic. This parasite occurs on the gills and 

is 2.6 mm long. It is probably genus (Thunnus) and tribe specific. This worm 

needs to be redescribed and more collections are needed in the western Atlantic 

to resolve the mystery of its distribution. 

CESTODA (TAPEWORMS) 

Tapeworms or cestodes form a large class of the flatworms or 

platyhelminths. The common name comes from the long series of body 

segments which resemble a tape measure. Most adult tapeworms look like long, 

flat cooked noodles. Most larval forms look like the tiny pieces of noodle that 

stick in the colander. A giant, broad tapeworm that lives in the body cavity of 

freshwater fishes in Europe is apparently routinely eaten by humans who mistake 

it for parts of the fish! Tapeworms can reduce growth and affect reproductive 

success of fishes. Some that infect humans occur as immature or larval forms 

in fishes. A small-boat fishery for tunas on the northwest coast of Puerto Rico 

was closed by the government because of reports of "wormy" flesh in these food 

fishes. When we were contacted to examine these parasites, we found that they 

were larval tapeworms, and recommended that the fishery be immediately 

reopened. We knew that previous experimental attempts to infect mammals with 

similar larvae failed because the parasites mature only in sharks and rays. The 

fishery was quickly reopened, averting major losses to local fishermen. We 

occasionally receive similar cases of wormy filets from individuals. People do 

not like seeing larval tapeworms moving around in the flesh of their fish, but 

those in Puerto Rican fishes are relatively harmless (particularly when cooked). 

More than 5000 species of tapeworms are known. Adults range from less 

than a millimeter to more than 30 meters in length. Tapeworms usually consist 

of a chain of segments (proglottids) each with a set of female and male 

103

104 


reproductive organs. The segments are continuously budded in the anterior 

portion of the body or neck, and enlarge and mature as they slowly move 

posteriorly. The scolex or "head" on the anterior end is usually armed with 

various combinations of suckers, hooks, bothridia (outgrowths), or bothria (sucking 

grooves) for attachment in the host intestine. Eggs escape through pores or 

a whole mature egg-filled segment may break off and pass out of the intestine. 

A successful tapeworm may produce millions of eggs over its lifetime. 

Eggs or their hatched larvae (ciliated coracidia or unciliated hexacanth or 

oncosphere) are eaten by the first intermediate host (insect, crustacean or 

annelid) and become elongate procercoids. This host is subsequently eaten by 

a vertebrate (second intermediate host) and the larvae develop into partially 

differentiated plerocercoids or plerocerci. Plerocercoids can be passed from one 

host to another when an infected fish is eaten by another fish. Feeding infected 

viscera to fish therefore can greatly concentrate or increase the intensity of 

infection by these worms. We have seen this practice cause a superinfection in 

caged red hind, Epinephelus guttatus Linnaeus, in Puerto Rico, and in cage 

cultured fishes raised in the northern Gulf of Mexico. If viscera must be used 

as fish food, it should be cooked or frozen for several days to kill parasites. 

When the correct final or definitive vertebrate host eats the second intermediate 

host the adult tapeworms develop in the intestine. 

Most tapeworms have both female and male sex organs in each proglottid. 

A few have separate sexes. They occur in all kinds of vertebrates and in all 

habitats around the world. All tapeworms are permanent parasites. The 

intestine of these worms has been lost thus food from the gut of the host is 

absorbed directly through the body wall. Few species of adult tapeworms are 

found in marine bony fishes and only 2 species occur in big game fishes. In 

contrast, many species of larval tapeworms are found in the intestinal tract, often 

in large numbers, and a few are encapsulated in the tissues of marine bony 

fishes including big game fishes. Most of these larval forms are found as adults 

in sharks or rays. A necropsy is necessary to find adults or larvae in the gut 

and encapsulated larvae in internal organs. Adult tapeworms can be relaxed in 

tap water until they no longer react to touch, and preserved in 5% formalin. 

Although some fish tapeworms can mature in humans, none of those in marine 

fishes infect people. We will consider the adult tapeworms first and the larval 

forms (*) second. 

Reference - No recent guide to the species of tapeworms exists. "How to know 

the tapeworms" (Schmidt 1970) is a semi-popular guide to the genera and 

Schmidt (1986) is a more technical revision which includes lists of the known 

species. Khalil, Jones and Bray (1994) update the genera and higher 

classification of tapeworms, but do not list the species [see our review of this 

book (Williams and Bunkley-Williams 1995)].



Class Cestoda (or Cestoidea) - tapeworms Page 

Order Trypanorhyncha 

Family Tentaculariidae 

Nybelinia bisulcata* .................................................................. 110 

Nybelinia lamonteae* ............. .................................................. 111 

Nybelinia lingualis* ......................... ......................................... 111 

Nybelinia sp.* ................................. .......................................... 112 

Tentacularia coryphaenae* ....................................................... 112 

Tentacularia sp.* ....................................................................... 114 

Family Hepatoxlidae 

Hepatoxylon trichiuri* .............................................................. 114 

Family Sphyriocephalidae 

Sphyriocephalus sp.* ............................................................... 132 

Family Eutetrarhynchidae 

Eutetrarhynchus lineatus ........................................................ 116 

Family Otobothriidae 

Otobothrium crenacolle* ....................................................... 117 

Otobothrium dipsacum* ......................................................... 118 

Family Pterobothriidae 

Pterobothrium heteracanthum* .............................................. 118 

Family Grillotiidae 

Grillotia erinaceus* ................................................................ 119 

Pseudogrillotia zerbiae* ......................................................... 121 

Family Molicolidae 

Molicola horridus* .................................................................. 122 

Family Lacistorhynchidae 

Callitetrarhynchus gracilis* ................................................... 123 

Lacistorhynchus bulbifer* ....................................................... 124 

Family Dasyrhynchidae 

Dasyrhynchus giganteus* ....................................................... 125 

Family Gymnorhynchidae 

Gymnorhynchus gigas* .......................................................... 126 

105

106 


Species of uncertain status 

incompletely defined larval species* ...................................... 131 

trypanorhynchid plerocercoid* ............................................... 132 

Order Tetraphyllidea 

Family Prosobothriidae 

Prosobothrium armigerum* .................................................... 126 

Family Phyllobothriidae 

Ceratobothrium xanthocephalum* .......................................... 127 

Echeneibothrium sp.* .............................................................. 131 

Pelichnibothrium speciosum* ................................................. 127 

Phyllobothrium delphini* ....................................................... 128 

Rhinebothrium flexile* ............................................................ 128 

Species of uncertain status 

tetraphyllid plerocercoid* ........................................................ 129 

Order Pseudophyllidea 

Family Bothriocephalidae 

Bothriocephalus janikii ........................................................... 131 

Bothriocephalus manubriformis ............................................. 106 

Bothriocephalus scorpii* ........................................................ 130 

Bothriocephalus sp.* .............................................................. 130 

Family Triaenophoridae 

Fistulicola plicatus ................................................................. 107 

Pseudobothrium grimaldii ............................................. 132 

Miscellaneous tapeworms ............................................. 

131 

_______ 

*Larval forms 

Bothriocephalus manubriformis (Linton) 

This giant tapeworm infects billfishes around the world. Very 

heavy infections of hundreds of large worms damage these 

important sport fishes. 

Diagnostic Characters - This is a very large parasite. It has an 

elongate scolex with an apical disk and long bothria which are 

open at the anterior and posterior ends. The egg-filled segments 

(gravid proglottids) are wider than long and notched in the middle 

of their posterior margin. The posterior margin overlaps the 

anterior margin of the next proglottid (craspedote condition). 

Records - We found 2-4 in 8 of 40 blue marlin, 3-4 in 2 of 3 

white marlin, and more than 100 in a longbill spearfish off Puerto


Rico, and more than 50 in an Atlantic sailfish, and more than 100 in a white 

marlin off Dauphin Island, Alabama, USA. Numerous adults occurred in 3 of 

4 Atlantic sailfish off Miami, Florida; 108 in an Atlantic sailfish and 12-52 in 

4 white marlin from the southern Gulf of Mexico; 4-463 in 33 white marlin off 

Maryland, USA (USNPC 80329); more than 50 in a blue marlin (USNPC 

4711) and 50-77 in 2 sailfish from Woods Hole, Massachusetts, USA; and 1-12 

(average 4.8) in 52 of 63 black marlin off Queensland, Australia. 


Life History - Tapeworms must live in 

the intestine for several years, to grow to 

0.5-1.0 meters long. The preponderance 

of smaller worms suggests that 

competition of the adult worms or host 

immunity prevents most of these 

parasites from maturing. 

Ecology - Fishes caught near shore have 

fewer immature tapeworms than those caught offshore suggesting that the last 

intermediate host in the life cycle may occur offshore. 

Associations - Occasionally 3-5 tapeworms use the same perforation in the 

intestinal wall to penetrate and anchor their scolices in a capsule. 


Length - Adults 130.0-1000.0 mm or longer (more than 1 meter); immature 

worm as small as 25.0 mm. 

Host Specificity - It is family specific to billfishes. It may be a characteristic 

parasite of sailfishes and white marlin, but seems to occur more erratically in 

blue marlin. Many of the records in swordfish may be based on immature 

worms, or misidentifications, as some of the more careful and extensive studies 

have not found this worm in this host. 

Damage to Host - Adult tapeworms have been blamed for retarded growth, 

weight loss and emaciation of billfishes. The effect of larval and immature 

worms has not been established, but they are much more abundant than adults 

in many cases. Four worms formed a deep pit in the anterior intestine of a 

white marlin from Puerto Rico. 

Fistulicola plicatus (Rudolphi) 

This characteristic parasite of swordfish must damage and limit the growth 

of this valuable fish. 

Name - Pseudeubothrium xiphiados Yamaguti is a synonym. Bothriocephalus 

truncatus Leuckart and Fistulicola xiphiae (Gmelin) may be synonyms. 

Diagnostic Characters - This is a very large parasite. The scolex has an 

apical disk and a rounded shallow bothrium on each side. The scolex may be 

replaced by an elongate and sometimes swollen pseudoscolex in worms 

embedded in the intestinal wall. Segments are short and have leaf-like lateral 

expansions. 

107

108 


Records - Eight worms occurred in a 

swordfish off La Parguera, 1 in 1 off 

Guanica, and 6 in 1 off Humacao, Puerto 

Rico. One to 22 (average 5.5) occurred 

in 278 of 303 (ARC 2310), and 101 

(average 4.2) in 24 swordfish from the 

northwest Atlantic; and 1-9 in 14 from 

Woods Hole, Massachusetts, USA 

(USNPC 4736, 8861). 


Associations - Occasionally 3-5 

tapeworms use the same perforation in 

the intestinal wall to penetrate and anchor 

their scolices in a capsule. 

Location in Host - Intestine and rectum. 

Length - Adult 75.0 mm to more than a 

meter. 

Host Specificity - This parasite is only 

found in swordfish. It is a characteristic 

parasite of this host. 

Damage to Host - Hogans et al. (1983) 

reported a maximum infection of 22 of 

these large tapeworms in a host. This should have been a sufficient parasite 

load to cause problems in a swordfish. However, they reported an average of 

5.5 worms per host suggesting that few of these fish were this highly infected. 

Most perforations of the intestine occur in the rectum. Large thick host capsules 

are produced around the pseudoscolices of these worms. Even the tapeworms 

that do not penetrate the wall of the intestine produce irritation around their 

attachment positions. Blockage of the intestine by masses of these worms has 

been reported, but this needs to be confirmed. 

Preparation for Study - Skillful dissection is necessary to obtain intact worms 

from capsules. 

Larval Tapeworms 

Almost all our knowledge about tapeworms concerns adults. The adult and 

larval forms of a tapeworm are seldom encountered in the same host species or 

even in the same scientific study, and larvae often share few features with the 

adult tapeworms. Thus, larvae are usually not mentioned or are considered only 

incidentally, incompletely and often erroneously. Few complete life cycles of 

fish tapeworms have ever been determined. Most larval forms in fishes have 

been "identified to species" by guessing which adult tapeworm the larvae 

resemble. These difficulties have caused larval forms to be under-identified (to 

class, order or genus), misidentified, or ignored completely. Thus the state of 

knowledge, is what could only be called "one hell of a confusing mess." We


hope the following effort will be a framework to begin properly identifying these 

worms in big game fishes. Many more collections and examinations and much 

inspiration will be needed before we can understand the role larval tapeworms 

play in the health of these fishes. 

Larval forms in big game fishes are third stage or plerocercoid larvae 

belonging to 4 orders of tapeworms. They fall into 2 categories: (1) those 

encapsulated in the tissues or body cavity, and (2) those in the gastrointestinal 

tract. A few plerocercoid larvae in the tissues are elongate unstructured Bothriocephalus 

spp. in simple capsules. But almost all are trypanorhynchid 

plerocerci with 4 hook covered tentacles in complex capsules composed of 3 

layers: (1) an outer capsule produced by the host, (2) a blastocyst inside the 

capsule, and (3) a plerocercus larva inside one end of the blastocyst. A few 

capsules and/or blastocysts are so large and distinctive that they can be identified 

immediately to species. Unfortunately, most require careful measuring, 

dissection, and examination of the characters of each layer. The shape, 

thickness, color, location in the host, location of the blastocyst, and attachment 

of the capsule should be recorded and drawn. The capsule is dissected from the 

host tissue, placed in a container of saline, viewed with a dissection microscope 

and carefully opened to allow the blastocyst to escape. Some features are best 

seen while this form is alive and moving. The next step is to evert or pop out 

the scolex from the blastocyst. This requires experience, luck, or a large supply 

of capsules. Gentle pressure on the blastocyst with a blunt probe should pop out 

the scolex, but too much pressure may cause it to rupture into pieces. Either 

a "postlarva" with a tail (appendix) will separate from the blastocyst, or a 

"plerocercus" will evert on the end of the blastocyst, but remain attached. 

Larvae may be best seen in a wet mount viewed with a compound microscope. 

Some encapsulated larvae have ample characters to allow identification to 

species but many cannot be deciphered. In recently encapsulated larvae, 

characters may not be fully developed. Older capsules may contain dead larvae 

degenerated into dark, waxy masses or calcified and partially reabsorbed by the 

host. Mature postlarvae separate from capsules embedded in the outer layer of 

internal organs of fishes and are found free in the body cavity. 

Trypanorhynchid postlarvae and plerocerci have almost mature scolices similar 

to those found in the adult. 

The second category of tapeworm larvae are those found free in the stomach 

and intestine of fishes. These postlarvae, plerocerci and capsules have been 

digested from invertebrates or fishes eaten by the host. Some of the larger 

larvae have distinctive attachment structures on the scolex that can be used to 

identify them to species. Unfortunately, the vast majority of these larvae are 

tiny tetraphyllid plerocercoids (see species summary) that are too indistinct to 

be identified to family, much less species. This tetraphyllid reputation for being 

difficult to identify is often and unjustifiably applied to all larval tapeworms in 

fishes. It has been used as an excuse to ignore this varied and fascinating group 

of parasites. These loose larvae in the alimentary canal of big game fishes are 

in false hosts. We do not know if they are quickly digested, pass out of the 

intestine, or if they remain longer and use big game fishes as intermediary hosts. 

109

110 


Most plerocercoid larvae have no specificity. Some are restricted to particular 

habitats or regions and show promise as biological tags to distinguish 

stocks of big game fishes and their migrations. If we knew more about their 

life cycles, they could potentially tell us what, when and where the fish ate, and 

possibly reveal a variety of other biological secrets. 

We have attempted to distinguish larval tapeworms with obvious characters 

that can be seen with a hand lens or a dissection microscope. In some species, 

the hooks on the tentacles were used. This requires placing the larvae in wet 

mounts on microscope slides and viewing them with a compound microscope. 

Larval tapeworms can be relaxed and preserved in steaming, 5% formalin; 

or relaxed in tap water and fixed in 5% formalin. Neither method assures that 

tentacles, important in the identification of trypanorhynchids, will always be 

fully everted so that their hooks can be seen. Lidocaine is used to evert the 

tentacles. The best insurance for success is to study many specimens. 

Nybelinia bisulcata (Linton) 

This larval tapeworm is occasionally encapsulated 

in the stomach wall or is found as postlarvae in the 

intestine of big game fishes. 

Diagnostic Characters - Most capsules are relatively 

small and oval and contain oval blastocysts. This 

worm has the characters of Nybelinia sp. listed below. 

The tiny tail is as long as wide and less than 1/4 the 

width of the body. Bothridia are broad, bean-shaped 

and longer than 1/2 the length of the body. Spine-like 

structures (microtriches) are visible on the margins of 

the bothridia. Tentacles are relatively short. 

Records - One to 2 postlarvae occurred in 9 and 1-2 

blastocysts in 3 of 40 Atlantic blue marlin and 1-3 

postlarvae in 17 of 40 dolphin examined from various 

sites around Puerto Rico, and 14 and 27 capsules in 2 

swordfish from off Guanica Bay, Puerto Rico. 

Capsules were very numerous in 3 of 5 dolphin off 

Miami, Florida; a few capsules and postlarvae in a 

cero, a few capsules in 1 of 8 cobia, 1 capsule in 1 of 

15 crevalle jack, and 1 postlarva in a Spanish mackerel 

from Beaufort, North Carolina; 1-2 capsules occurred 

in 2 Atlantic mackerel and several capsules and a few postlarvae in 2 swordfish 

from Woods Hole, Massachusetts, USA. These larvae are more common in 

non-big game bony fishes, sharks and rays; and occasionally occur in squids. 

Geographic Range - Possibly worldwide, but most records are from the U.S. 

Atlantic. Our collections are the first in the Caribbean. 

Life History - The adult tapeworm occurs in sharks. 

Location in Host - It is usually encapsulated in the stomach wall or on the 

sides of the stomach, and postlarvae are in the body cavity or in the intestine.


Length - Plerocercoid usually 0.4-1.2 mm, exceptionally to 20.0 mm, blastocyst 

0.1-3.5 mm, capsule 0.2-4.8 mm. 

Host Specificity - Atlantic blue marlin is a new host for this parasite. 

Nybelinia lamonteae Nigrelli 

A common larva in swordfish from the northwest Atlantic. 

Our records from Puerto Rico suggest that it may be more 

widespread and may occur in other hosts. 

Diagnostic Characters - It has the characters of Nybelinia sp. 

below, but with fine spine-like structures (microtriches) visible 

along the margins of the bothridia. Tentacle sheaths are longer 

than the tentacles or the bulbs. Tentacle bulbs are more than 

3 times as long as wide and are only slightly overlapped by 

bothridia. 

Our specimens differ from the description of Nigrelli 

(1938) by having a bothridial area of the scolex that is 

rectangular, tentacle sheaths that curl near the bulbs, and a tail 

that is partially invaginated. 

Records - One blastocyst occurred in a dolphin; and 1-2 

postlarvae in 2 of 6 king mackerel from La Parguera, Puerto Rico (USNPC). 

One capsule was found in a swordfish from Nova Scotia, Canada; and 1-21 

(average 2.8) postlarvae occurred in 154 of 303 swordfish from the northwest 

Atlantic (ARC 2314). 

Geographic Range - Unknown. Our collections are the first in the Caribbean. 

Life History - Nigrelli (1938) suggested that the adult of this worm could be 

N. robusta (Linton) which parasitizes rays and is encapsulated in shark remora. 

Location in Host - These worms were found encapsulated in the mesenteries 

or as free postlarvae in the stomach. 

Length - Plerocercoid 1.3-7.0 mm. 

Host Specificity - Dolphin and king mackerel are 

new hosts. 

Nybelinia lingualis (Cuvier) 

This larval tapeworm is occasionally encapsulated 

in the viscera or is found as postlarvae in the intestine 

of big game fishes. 

Diagnostic Characters - This worm has all the 

characters of Nybelinia sp. listed below. The bothridia 

are bean-shaped and lack spines along the margins. 

Tentacle sheaths are approximately as long as the 

tentacles. Tentacle bulbs are about twice as long as 

wide and are largely covered by the bothridia. 

Records - One blastocyst occurred in 1 of 2 skipjack 

tuna from off Arecibo, Puerto Rico (USNPC); and in 

111

112 


swordfish from Woods Hole, Massachusetts, USA. The postlarvae occur 

commonly in bony fishes, sharks and rays. 

Geographic Range - Possibly worldwide, but almost all records are European. 

Our collections are the first in the Caribbean. 

Location in Host - Viscera (blastocysts) and intestinal tract (postlarvae). 

Length - Postlarva 1.3-4.0 mm. 

Host Specificity - Skipjack tuna is a new host. 

Nybelinia sp. 

Name - Several larval species are included under this name. In most cases, 

characteristics were not recorded and specimens were not deposited so they 

cannot be identified to species. Some of the larvae reported may have been the 

3 species described above, but other species exist. 

Diagnostic Characters - This postlarva has a short scolex and a short, usually 

invaginated tail. Four elongate bothridia are about 1/2 the length of the scolex. 

Tentacle bulbs extend posterior to the end of the bothridia. The posterior 

margins of the bothridia are free. The hooks are almost identical throughout the 

length of the tentacle. Extreme variations in shape typically occur. 

Records - Six postlarvae occurred in 1 of 2 Atlantic blue marlin, 6-15 in 2 of 

4 white marlin; 1-5 in 4 of 10 yellowfin tuna from the southern Gulf of Mexico; 

and 3-11 in 5 of 9 swordfish from the northwest Indian Ocean. 

Geographic Range - Worldwide 

Life History - The postlarvae are common in bony fishes and elasmobranchs. 



Tentacularia coryphaenae Bosc 

Larvae of this worm are encapsulated in a variety of big 

game and other bony fishes around the world. 

Name - Tentacularia bicolor (Bartels) is a synonym. 

Diagnostic Characters - This worm has all the characters of 

Tentacularia sp., noted below. In addition, the combined length 

of the tentacle sheaths and bulbs are about 1/3 to 1/2 the length 

of the bothridia. The bothridia vary from a uniform width 

throughout to tapering to a point on the posterior end. 

Records - We found postlarvae in an albacore from Desecheo 

Island; 1-22 postlarvae in 25 and 1-3 blastocysts in 7 of 40 

Atlantic blue marlin; 3 blastocysts in a blackfin tuna from 

Boqueron; 1-14 (average 6.1) postlarvae in 7 of 13 dolphin 

from La Parguera (USNPC); 1-100 blastocysts in 14 and 1 to 

many postlarvae in 22 of 40 dolphin from various localities 

around Puerto Rico; numerous blastocysts in 2 skipjack tuna 

from off Arecibo; 1 each in 1 of 3 wahoo from Boqueron, and 

3 in a wahoo from Desecheo Island, Puerto Rico and 2-10 postlarvae in 3 of 5 

dolphin from Dauphin Island, Alabama, USA. Numerous postlarvae were found


in 2 of 5 dolphin and 1 of 2 king mackerel from Miami, Florida, USA; 1 

capsule in 1 of 9 little tunny from Bermuda; 1 and few blastocysts in 2 of 6 

Atlantic bonito; numerous capsules in a dolphin (USNPC 5483); and many in 

each of 4, 10 and 1 in 2, 3 in 1 (USNPC 4820), and 4 in 1 (USNPC 4829) 

swordfish from Woods Hole, Massachusetts, USA; swordfish off Nova Scotia, 

Canada; 1-7 postlarvae in 18 of 303 swordfish from the northwest Atlantic 

(ARC 2309); pompano dolphin from an unknown Atlantic locality; bluefin tuna, 

wahoo and yellowfin tuna off west Africa; chub mackerel, dolphin and skipjack 

tuna from the Pacific; and in a variety of other fishes and occasionally squids. 

Only 1 dead, encapsulated worm was found in 400 albacore from the southwest 

and south central Pacific. 

Capsules of this worm are visible through the peritoneum in the wall of the 

body cavity and were counted in 1529 skipjack tuna from the southwestern 

Pacific. In 1017 of these fish collected from tropical areas, the numbers of 

capsules increased (0-18 worms) with increasing fish length (33-75 cm). In 512 

fish collected from off New Zealand, the number of capsules in 45 cm fish was 

the same (N=7) as 45 cm fish collected from the tropics, but the number of 

capsules did not increase with increasing fish length (45-65 cm) (Lester, Barnes 

and Habib 1985). These data suggest that the fish off New Zealand originally 

came from the tropics when approximately 45 cm in length and remained there. 

It also suggests that the parasite must use a tropical intermediate host since no 

further infections were accumulated in the New Zealand fish. 


Life History - Adult tapeworms parasitize blue shark, Prionace glauca 

(Linnaeus), another offshore, pelagic fish, and dusky shark, Carcharhinus 

obscurus (Lesueur). 

Associations - Three postlarvae used their tentacles to penetrate the tissues of 

another postlarval tapeworm, Hepatoxylon trichiuri, that occurred with them in 

the stomach of a swordfish. More records of these attacks would be useful in 

determining if these are routine aggressive acts or mere accidents. Host 

immunity and chemical effects of other parasites have been thought to control 

the numbers of worms in the intestine of at least mammalian hosts. The 

physical combat of hooked-tentacle duels for survival among these highly 

energetic worms might also be considered. 

Location in Host - Blastocysts occur in the muscles, sides of the stomach and 

other organs. Postlarvae free from the blastocyst are found in the body cavity, 

stomach and intestine. 

Length - Postlarva 1.3-9.5 mm, capsule up to 60.0 mm. 

Damage to Host - Hard, calcified capsules covered the entire intestine and 

pyloric ceca of an Atlantic blue marlin from Puerto Rico. Three fresh capsules 

40-60 mm long were also found in this fish. This larvae has been found 

associated with inflamed areas and ulcers in the stomach of swordfish. 

Host Specificity - Plerocercoid larvae occur in a variety of fishes. The adult 

occurs only in sharks. Atlantic blue marlin and blackfin tuna are new hosts. 

113

114 


Significance to Sport Fishing - The capsule is easy to see through the 

peritoneum in the body wall of skipjack tuna, it lives for a number of years in 

the host, and is an excellent biological tag for discrimination of stocks and to 

indicate migrations. 

Tentacularia sp. 

Name - This name includes several larval species. In most cases, 

characteristics were not recorded and specimens were not deposited so they 

cannot be identified to species. Some of the larvae reported may have been T. 

coryphaenae, but other species exist. 

Diagnostic Characters - This small to moderate-sized postlarva has a long 

scolex with a short tail. The scolex has 4 shallow, elongate, separate 

attachment grooves (bothridia) without free edges; and 4 short, retractile hookbearing 

tentacles. The tentacle sheaths extend from bases of the tentacles and 

are relatively short, almost straight (not twisted), and do not extend beyond the 

posterior border of the bothridia. The hooks on the tentacles are almost 

identical (except that those at the bases of the tentacles are smaller) and closely 

spaced (homeoacanthous armature). Tentacle bulbs are banana-shaped and 

placed in the anterior 1/2 of the bothridial region. 

Records - One larva occurred in a great barracuda from Jamaica; 2 in 1 of 2 

Atlantic blue marlin, 7 in an Atlantic sailfish, 1 in 1 of 4 white marlin and 1 and 

8 in 2 of 10 yellowfin tuna from the southern Gulf of Mexico, capsules in 29 of 

45 yellowfin tuna from the northeastern Gulf of Mexico (USNPC 56909) and 

5-8 larvae in 2 of 10 Indo-Pacific sailfish from the northwest Indian Ocean. 


Life History - Adult tapeworms occur in sharks. 

Ecology - The postlarvae are found in bony fishes (teleosts) in shallow, midwater 

and continental slope habitats. 

Location in Host - Blastocysts occur in the mesenteries, muscle and other 

viscera. Postlarvae occur in the body cavity, stomach and intestine. 


Host Specificity - The larvae appear to have no specificity. Adults of most 

species are specific to 1 or a few shark species. 

Detection - Capsules in muscle or internal organs are difficult to find. Soft 

organs can be examined by squashing. In muscle cut into thin strips and held 

up to a light, capsules appear as opaque areas in the translucent flesh. 

Hepatoxylon trichiuri (Holten) 

Encapsulated larvae of this parasite occur commonly in swordfish and 

other fishes. 

Name - Modern authorities apparently disagree if these worms represent 1 or 

several species (or if H. squali (Martin) or H. trichiuri is the type of the genus). 

We consider Dibothriorhynchus attenuatus (Rudolphi), D. lepidopteri Blainville, 

D. xiphiae MacCallum, H. attenuatus (Rudolphi), H. grossus Rudolphi, H. 

stenocephala (Guiart), H. squali, and Hepatoxylon sp. of Ward, 1962 synonyms.


These larval forms are in need of additional study and 

may be shown to represent different species. Yamaguti 

(1959) suggested that H. stenocephala, represented by a 

postlarva from an unidentified dolphin in the Atlantic, 

was a distinct species. Hogans et al. (1983) considered 

this form from swordfish to be H. attenuatus. Postlarvae 

have been divided taxonomically into long and short 

forms, but these shapes are apparently due to 

developmental changes instead of species differences. 

Diagnostic Characters - Postlarvae are wrinkled all 

over except for the 4 bothridia. The tentacles are short, 

globular to conical, and vary from approximately as long 

as wide to twice as long as wide. Tentacular sheaths are 

about as long as the bulbs. Tentacular bulbs are more 

than 4 times longer than wide, and covered by the 

bothridia. The region containing the bothridia is less than 1/2 the length of the 

body. The postlarva varies greatly in shape and size. 

Records - We found 10-21 postlarvae in 15 

of 40 dolphin from various localities around 

Puerto Rico, 5 and 8 capsules in 2 swordfish 

off Guanica Bay, Puerto Rico; and 1 

postlarvae in 1 of 5 dolphin, and 3 in 1 of 4 

wahoo off Alabama, USA. Capsules were 

present in 14 of 45 yellowfin tuna from the 

northeastern Gulf of Mexico (USNPC 56907); 

1-24 blastocysts (average 3.4) in 178 of 303 

swordfish from the northwest Atlantic (ARC 

2313); 1 blastocyst in a large mass on the 

outer layer of the intestine, and 3 encapsulated 

on the outer layer of the gonads in 1 

swordfish each; many blastocysts and 

postlarvae in 8, 40 in 1, 10 each in 2, 8 in 1, 

and 2 in 1 swordfish from Woods Hole, 

Massachusetts, USA (USNPC 7732); 23 

postlarvae in a swordfish from Nova Scotia, 

Canada; postlarva in dolphin from Brazil and 

the Azores; capsules and postlarvae in albacore from the Mediterranean and the 

Pacific; and postlarvae in swordfish from the Baltic, eastern Atlantic and 

Mediterranean. The larvae of this parasite is also found in a variety of bony 

fishes, sharks and rays. 


Life History - Adults of this tapeworm are found in sharks. 

Ecology - Unlike most larval forms in big game fishes, it is common offshore 

and inshore in the Atlantic. In a study of 400 albacore in the Pacific, it was 

abundant near Australia (38%) and western New Zealand (32-75%), but declined 

115

116 


in the open Pacific (11-14%) and in the Coral Sea to the north (20%). This 

suggested that the intermediate host occurred in nearshore habitats (Jones 1991). 

Associations - One of 13 swordfish from Woods Hole was heavily infected 

with a tissue-embedding copepod (either Pennella filosa or P. instructa). The 

same fish had the heaviest infection of this larval tapeworm. 

Location in Host - Blastocysts encapsulated on the outside of the stomach and 

viscera, and postlarvae were free in the body cavity or in the intestinal tract. 

Length - Postlarva 7.5-70.0 mm; adults up to 400.0 mm. Linton (1924) found 

a 150.0 mm larva. 

Host Specificity - Wahoo is a new host. 

Significance to Sport Fishing - This parasite has been used as a biological 

tag to demonstrate how stocks of albacore migrate in the Pacific. 

Eutetrarhynchus lineatus (Linton) 

We solve a 75 year-old mystery by 

matching the larva from a big game fish 

with its adult. 

Name - The larval species Rhynchobothrius 

carangis MacCallum, a mystery 

(species in question) for more than 75 

years, is the larva of E. lineatus. Rhynchobothrius 

sp. of Linton (1908, Fig. 70) 

also appears to be a less mature specimen 

with the tentacle bulbs still bulging out 

the sides of the worm as they do in the 

larvae. 

Diagnostic Characters – Blastocyst 

white with an orange yellow posterior. 

This moderate-sized postlarva has a long 

scolex and a tail. The scolex has 2 round 

bothridia, with free posterior borders, that project from the anterior end. The 

4 tentacles are relatively long, tentacle sheaths are long and wavy or rippling, 

and tentacle bulbs are long and narrow. The body bulges out around the 

tentacle bulbs 

Records - One capsule occurred in a crevalle jack at the New York Aquarium 

that had been transported from Key West, Florida, USA; 1 blastocyst in a green 

moray, Gymnothorax funebris Ranzani; and 1-7 adult tapeworms in 4 nurse 

sharks, Ginglymostoma cirratum (Bonnaterre), from the Dry Tortugas, Florida. 


Life History - Linton's (1908) and MacCallum's (1921) records are the only 

times the larvae of this worm has been collected and described. The rarity in 

fishes, single specimens in fishes, and the almost fishless diet of the final host, 

suggest that the larvae is usually encapsulated in an invertebrate that might be 

consumed by a nurse shark.


Ecology - The final host occurs in inshore, shallow habitats. Only big game 

fish that venture inshore would be exposed to its larval forms. 

Location in Host - Both larvae occurred in the wall of the rectum. 

Length - Postlarva 4.3-11.5 mm; adult 15.0-30.0 mm. 

Host Specificity - Larvae are not specific since the 2 known specimens were 

found in different orders of fishes. The adult is host specific to nurse shark. 

Ttggggggggfdedfcvgt 

Otobothrium crenacolle Linton 

The larvae of this tapeworm occurs in many big 

game fishes. Superinfections may damage the internal 

organs of these fishes. 

Name - This species is sometimes spelled O. 

crenacollis. Diagnostic Characters - The blastocyst is 

an amber colored, oval capsule without a caudal 

extension. Postlarvae have 2 short bothridia, a short tail, 

and a longer striated body. Tentacle sheaths are spiral, 

and tentacle bulbs are short, less than twice as long as 

wide, and reach the posterior margin of the body. The 

bothridia usually show a posterior notch, and have comashaped 

sensory pits on their margins. Usually 1 larva 

occurs in a capsule, but up to 3 have been found. 

Records - Heavy infections 

of capsules occurred in dolphin and king mackerel 

and postlarvae in little tunny from Bermuda; a 

superinfection of capsules in a cero, and a very 

heavy infection of capsules in a dolphin from 

Beaufort, North Carolina, USA; a light infection 

of capsules in an Atlantic bonito, 1 postlarvae and a 

light infection of capsules in 2 blue runner, light 

infections of capsules in 4 cero (USNPC 5494), 

numerous capsules in a king mackerel, a few 

capsules in a swordfish and a light infection of 

degenerated capsules possibly of this worm in a 

white marlin (USNPC 5501) from Woods Hole, 

Massachusetts, USA. These capsules also occur 

in a variety of other bony fishes and sharks. 

Geographic Range - Atlantic including the Gulf of Mexico. 

Life History - Adults of this tapeworm occur in sharks. 

Location in Host - Blastocysts are usually found in the wall of the stomach, 

less often on pyloric ceca and other viscera, occasionally in muscle. Postlarvae 

occur in the intestinal tract. 

Length - Postlarva 0.3-2.4 mm, blastocyst 0.5-2.8 mm, capsule 1.5-7.0 mm. 

Damage to Host - Superinfections of more than 100 capsules per 13 cm 2 of 

visceral surface occur. 

117

118 


Otobothrium dipsacum Linton 

The larvae of this tapeworm is rarely encapsulated in 

big game fishes. 

Diagnostic Characters - The rather large (up to 24 mm) 

capsule is ovoid, often darkly colored. The blastocyst is 

oval and has no tail. The postlarvae have 2 wide, short 

bothridia, a short tail, and a longer striated body. The 

tentacle sheaths are spiral. The tentacle bulbs are long, 

approximately 1/2 as long as the body, more than 4 times 

as long as wide, and reach the posterior margin of the 

body. The bothridia have a posterior notch, and oval 

sensory pits on the posterolateral margins. 

Records - Three to 4 postlarvae occurred in 2 of 5 Atlantic 

sailfish, and 2-4 in 5 of 40 dolphin from various localities 

around Puerto Rico. Several postlarvae were found in 1 of 

19 great barracuda off Miami, Florida; and 1 capsule in 1 

of 2 swordfish from Woods Hole, Massachusetts, USA. 

This worm has also been found in Indo-Pacific sailfish. 

Blastocysts and postlarvae occur in other bony fishes, 

sharks and rays. 


Life History - The adult of this tapeworm occurs in sharks. 

Location in Host - Blastocysts occur on the outer layers of the viscera. Postlarvae 

are free in the body cavity or intestine. 


Host Specificity - This parasite does not appear to be a very common parasite 

of swordfish. It did not occur in 303 swordfish examined in the northwest 

Atlantic. Atlantic sailfish and dolphin are new hosts. 

Pterobothrium heteracanthum Diesing 

These elongate, brown capsules are found in the 

viscera of Spanish mackerels and inshore fishes. 

Name - Pterobothrium filicolle (Linton) is a synonym 

and P. fragile (Diesing) and Synbothrium sp. of Linton, 

1900 may be. Pterobothrium acanthotruncatum Escalante 

and Carvajal, encapsulated in dolphin from Peru, 

appears to be a different species. 

Diagnostic Characters - The elongate host capsules 

are brown. The "neck" of the blastocyst has a chara- 

cteristic constriction just before the long, rather wide tail 

joins the bulbous head. When the plerocercus everts 

from the blastocyst, it has an elongate, slender scolex 

with 4 round bothridia mounted on characteristic 

projections from the scolex. Tentacles are long and are 

along side each bothridium at the end of each projection


from the scolex. The tentacle sheaths are longer than 

the tentacles and the tentacle bulbs, relatively straight 

in the anterior, but irregularly curled near the tentacle 

bulbs. The tentacle bulbs are elongate and fill much of 

the width of the scolex and slightly bulge its width. 

Tentacles have 5 hooks in the principal rows and 

numerous smaller hooks. 

Records - Numerous capsules occurred in a cero and 

several in 1 of 2 Spanish mackerel from Beaufort, 

North Carolina, USA; 1 capsule in a greater amber- 

jack, numerous in 3 cero (USNPC 5486 and 5491), 3 

to numerous in 2 king mackerel (USNPC 4792), and 

numerous in 2 Spanish mackerel (USNPC 5487) from 

Woods Hole, Massachusetts, USA. 


Life History - Adults are not known. The plerocercus 

scolex remains attached to the blastocyst. It does not 

separate and form a postlarva. 

Ecology - Most of the known hosts occur in inshore 

habitats. Spanish mackerels may become infected when 

they feed inshore. 

Location in Host - Encapsulated in the wall of the 

stomach or intestine and in the outer layer of the liver, 

ovary and other internal organs. Occasionally capsules are 

deeply embedded in the liver. 

Length - Plerocercus 4.4-6.4 mm, blastocyst 25.0-94.0 

mm, capsule 3.0-10.5 mm. The length of the blastocyst 

varied considerably, but the plerocercus is approximately 

the same size in the longest and shortest blastocysts. 

Host Specificity – The 

occurrence of the plerocercus of this parasite in 3 of 

the 4 species of western Atlantic Spanish mackerels 

and no other big game fishes is apparently only a 

coincidence. 

Grillotia erinaceus (Van Beneden) 

This parasite is encapsuled in inshore fishes and 

occasionally big game fishes. Superinfections have 

been reported and might injure Atlantic mackerel. 

Name - Rhynchobothrium imparispine Linton is a 

synonym. Grillotia sp. Nikolaeva, 1962 may be a 

synonym. 

Diagnostic Characters - The capsule is oval or 

teardrop shaped and may be attached on a stalk. 

Capsules sometimes occur in clusters. Younger 

119

120 


capsules are white to yellow, older ones brown to black. The blastocyst is oval 

to oblong, but does not have a bulbous end and a tail. The plerocerus attached 

to the blastocyst has a long narrow, scolex, with 2 short, round bothridia. The 

bothridia have thickened, raised rims, and the lateral and posterior margins are 

free. Tentacles, tentacle sheath and bulbs are all relatively 

long. Bulbs are elongate, more than 6 times longer than 

wide, and do not reach the posterior margin of the scolex. 

Sheaths are irregularly tangled. Tentacles have 4 hooks in the 

principle rows. 

Records - One and a few cysts occurred in 2 of 31 Atlantic 

bonito; a superinfection in 1, numerous in each of 13, few to 

numerous in each of 23, a few in each of 6 and 1 in 1 of 32 

Atlantic mackerel (USNPC 4743); and few in 5 swordfish 

from Woods Hole, Massachusetts, USA. Four, 6 and 170 

plerocercoids of Grillotia sp. (possibly G. erinaceus) were 

found in 3 of 10 yellowfin tuna from the southern Gulf of 

Mexico. This parasite encapsulates in a great variety of bony 

fishes, sharks and rays. Eels and salmon carry this parasite 

from saltwater into freshwater habitats. 

Geographic Range - Possibly worldwide, but 

most records are from the northern Atlantic. 

Life History - The first intermediate host is 

probably a copepod. The second intermediate 

host is a fish. No free postlarvae occur in the life 

cycle. The adult is more common in skates and 

rays than in sharks. Parts of the life cycle were 

described by Ruszkowski (1932). A similar 

species, G. angeli Dollfus, survives for at least 

several years, if not the entire life of the host, in 

Atlantic mackerel in the Mediterranean and the 

Atlantic off Europe. 

Ecology - The capsule is more common in 

inshore than offshore fishes. 

Location in Host - Encapsulated in the outer 

layer of viscera, in the intestinal wall, or occasionally in muscle. Capsules are 

sometimes found in the intestinal tract after being digested out of prey fishes. 

Length - Plerocercoid 4.2-7.0 mm, capsule 2.5-13.0 mm, exceptionally 22.0 

mm. 

Host Specificity - This parasite occurs in a great variety of fishes, but in few 

big game fishes. It only occurred in high numbers in Atlantic mackerel. The 

report from Atlantic bonito was of free capsules in the intestine digested out of 

other fishes. It does not appear to be a very common parasite of swordfish. It 

did not occur in 303 swordfish examined in the northwest Atlantic. 

Damage to Host - Superinfections are probably injurious to the host. 

Infections of G. branchi Shaharom and Lester plerocercoids in the tissues of the


gill arches of Spanish mackerels in Australia and Malaysia caused inflammation, 

production of black pigment (melanization) and erosion of bone. 

Comments - This is a common parasite in north Atlantic fishes and has been 

the subject of a large number of articles and a book (monograph) (Johnstone 

1912). 

Pseudogrillotia zerbiae Palm 

Large spaghetti worms frequently wiggle out of highly 

prized greater amberjack fillets to the disgust of fishermen 

and cooks. 

Name - The common name "spaghetti worm" is often used 

to describe its appearance in the flesh of fishes. 

Pseudogrillotia sp. of Deardorf, Raybourne and Mattis, 

1984 is a synonym. 

Diagnostic Characters - The blastocyst in the flesh of a 

fish is a long white worm with a bulbous "head" and a 

thick, string-like body. The plerocercoid larva has a scolex 

with 2 round bothridia. The tentacle sheaths are winding 

but not spiral. 

Records - We found numerous worms in greater 

amberjack from Arecibo and Aguadilla, Puerto Rico, and 

in 10 off Panama City, Florida, USA. It has also been 

found in greater amberjack off Mississippi (BMNH 

1995.3.20.1-2) and Hawaii; and in black marlin and jacks 

in Hawaii 


Caribbean. 

Location in Host - This worm is embedded in muscle. More are found along 

the spine and in the muscle towards the tail. 

Length - Plerocercoid 23.0-25.0 mm, blastocyst 20.0-55.0 mm. 

Host Specificity - It is possibly a characteristic parasite of greater amberjack. 

This parasite also occurs in 

rudderfish and could be 

almost genus specific 

(Seriola), or almost family 

specific to jacks. Rudderfish 

is a new host. 

Detection - Actively moving 

worms can easily be seen 

when fillets are removed 

from freshly caught fish. 

Harm to Humans - Large, 

live spaghetti worms wiggling 

in the flesh of greater 

amberjack have caused many 

121

122 


fillets to be needlessly discarded. This was not much of a problem 30 years ago 

when this big game fish was considered a "trash" fish, but now it has become 

a "highly prized" food fish. These worms do not harm humans. They have 

experimentally been fed to mammals and did not develop or cause harm. They 

are sufficiently large to be picked out of the flesh. If you are too squeamish to 

pick out or eat these worms, then avoid cutting fillets too close to the spinal 

column or too close to the tail. This technique will waste some fish muscle, but 

will avoid most of the worms. You might also refrain from showing these 

parasites to non-fishing spouses, if you want to be able to eat your catch! 

Preparation for Study - These long worms are easily plucked from muscle 

and preserved. They make impressive classroom demonstrations of seafood 


Significance to Sport Fishing - The problem of wormy flesh in greater 

amberjack has been impressing fishermen for the last 25,000 years but this 

worm was only described scientifically in 1995. 

Molicola horridus (Goodsir) 

This parasite rarely is encapsulated in muscle of swordfish. 

Name - The genus was named for ocean sunfish (genus 

Mola). Molicola uncinatus (Linton) is a synonym. 

Diagnostic Characters - The blastocyst is elongate, but 

does not have a bulbous anterior or a long tail. The 

plerocercoid larvae consists of a long scolex and a tail. 

The scolex is narrow and cylindrical except for the 4 short, 

broad bothridia which are wider than the neck. The 

tentacles are long, and the tentacle sheaths long and spiral. 

The tentacle bulbs are short, oblong, about 3 times as long 

as wide, and almost fill the width of the scolex. The 

tentacles have a ring of hooks at the base which contains 

the largest hooks. 

Records - One blastocyst occurred 

in a swordfish from 

Woods Hole, Massachusetts, 

USA; and it commonly oc- 

curs in ocean sunfish, Mola mola (Linnaeus), a 

pelagic offshore fish. 


Life History - The adult of this parasite has been 

listed in tapeworm guide books from the mys- 

terious "Vulpecula marina" for more than 70 years 

although Linton (1924) calls it a "thrasher" 

[=thresher shark, Alopias vulpinus (Bonnaterre)]. 

Adults also occur in the pelagic and offshore blue 

shark.


Ecology - Members of this parasite genus occur in offshore, pelagic habitats. 

Location in Host - Blastocysts usually occur in the liver, but were found in the 

muscle near the spinal column of swordfish. 

Length - Plerocercus 2.0 mm, blastocyst 89.0 mm. 

Callitetrarhynchus gracilis (Rudolphi) 

This parasite occasionally is encapsulated in a variety of big game fishes. 

Name - This species was originally named from a plerocercoid 

larva in a bullet tuna or frigate tuna. Tentacularia lepida 

Chandler is a synonym. 

Diagnostic Characters - The host capsule is bladder-like to 

elongate and usually white, but older capsules turn brown to blueblack 

and slightly iridescent. The postlarva has an elongate 

scolex, long tail, and 2 short, heartshaped 

bothridia. The tentacle 

sheaths are tightly coiled. The 

tentacle bulbs reach the end of the 

scolex, do not occupy the entire 

width of the scolex, and are about 3 

times longer than wide. The base 

of the tentacles does not have a ring 

of larger hooks. 

Records - One to 2 postlarvae occurred in 7 of 40 

dolphin from various sites around Puerto Rico; 

capsules were common in Spanish mackerel, 1-4 

in 11 of 52 bar jack, 4 in 1 of 4 blue runner and 

postlarvae in bar jack from Bermuda; several 

capsules in 17 of 32 little tunny and/or skipjack 

tuna off Miami, Florida, USA. Two blastocysts were 

found in a dolphin and 1 in a pompano dolphin from 

Beaufort, North Carolina, USA; 1-20 capsules in 2 of 8 

Atlantic mackerel, 1 capsule in a bluefin tuna, and 1 capsule 

in a blue runner, from Woods Hole, Massa- 

chusetts, USA; in bluefin tuna from the Mediterranean; 

yellowfin tuna from west Africa; bluefin tuna, chub 

mackerel, frigate tuna and Indo-Pacific sailfish from the 

Pacific; and in other fishes, particularly groupers and 

snappers. 



Life History - The adult occurs in sharks. 

Location in Host - Blastocysts occurred in the wall of the 

stomach and intestine, or in the outer layers of the viscera. 


123

124 


Lacistorhynchus bulbifer (Linton) 

This parasite is occasionally encapsulated in a variety of big 

game fishes. It is more commonly found in inshore fishes. 

Name - Lacistorhynchus tenuis (van Beneden) was considered one 

worldwide species, but it has been shown to be a species complex 

with L. tenuis in Europe, L. dollfusi Beveridge and Sakanari in the 

Pacific, and possibly L. bulbifer in the western Atlantic. Rhynchobothrium 

bulbifer Linton is a synonym. 

Diagnostic Characters - The host capsule is swollen on one end 

and has a tail on the other. The blastocyst has a tail that varies in 

length. The tentacles of the postlarvae emerge from the sides of the 

anterior end of each bothridium, instead of on top of the scolex. 

The postlarvae has a long scolex and a short tail. The 2 bothridia 

are elongate with raised rims. The tentacle sheaths are spiral, the 

bulbs 

elongate, and more than 4 times longer than wide. The scolex 

bulges around the tentacle bulbs. 

Records - One and few capsules occurred in 2 Atlantic 

bonito; 12 in 1, few in 12 of 56, and 1 in 1 Atlantic 

mackerel; 1 in a bluefin tuna, 1 in a blue runner, several in 

a little tunny and 2 in a Spanish mackerel from Woods 

Hole, Massachusetts, USA; and in a variety of other fishes. 


Life History - Coracidia of L. dollfusi were infective to 

copepods for 2 weeks, procercoids in copepods were 

infective to bony fishes for 4 weeks, eversible tentacles 

developed in the plerocercoids in 3 months, and worms in 

the final host began to form segments in 4 months at room 

temperature. Development and survival times are longer at 

lower temperatures. Other details of the life cycle have 

been reported by Young (1954) and Sakanari and Moser 

(1985a,b). 

Ecology - Many of the final hosts for this tapeworm are smaller, inshore 

sharks. Thus the larval forms are more common in inshore fishes and are only 

occasionally found in big game fishes. 

Associations - One capsule was associated with a capsule of Koellikeria sp., 

probably K. bipartita, in a bluefin tuna. 

Location in Host - The larvae are encapsulated in the intestinal or stomach 

wall, in the outer layers of viscera, or occasionally free in the intestine, digested 

out of food items. They are also found encapsulated in the dorsal muscles of 

Atlantic mackerel. 

Length - Postlarva 1.2-3.5 mm, capsule 1.5-4.5 mm. 

Damage to Host - Five or more plerocercoid larvae of L. dollfusi developing 

into immature adults apparently killed young sharks. Infections kill 20-50% of 

the copepod intermediate hosts.


Dasyrhynchus giganteus (Diesing) 

This spectacular capsule in the common and often caught 

crevalle jack deserves more attention. 

Name - The genus is appropriately named for sting rays (genus 

Dasyatis). The Dasyrhynchus sp. of Overstreet, 1978 is the same 

as this worm (Overstreet pers. comm.). Rhynchobothrium insigne 

Linton is a synonym. 

Diagnostic Characters - This large blastocyst has an oblong 

anterior and a long tail. The tiny postlarva has a long, narrow 

scolex which gradually expands in width from the front to the base, 

and has 2 short, inverted-heart-shaped bothridia. Tentacle sheaths 

are long and tightly coiled. Tentacle bulbs are long and narrow. 

The tentacles have 9 dissimilar hooks (5 larger and 4 distinctly 

smaller) in each principle row, rows of smaller hooks occur between 

principle rows, and a double file or chainette of closely spaced 

hooks is found on one side of each tentacle. The postlarvae of D. 

variouncinnatus (Pintner) is identical, but it only occurs in the 

Pacific. 

Records - Blastocysts commonly infect crevalle jack in the northern 

Gulf of Mexico; 6 capsules in a greater amberjack off Miami, 

Florida; 1 postlarva each in 8, 2 in 1 and 3 in 1 of 303 swordfish from the 

northwest Atlantic (ARC 2414); 1 in a leatherjacket, Oliogloplites saurus 

(Schneider), from Brazil, and other fishes from Florida, USA. 


Life History - The adult occurs in several species of sharks. 

Location in Host - Usually found 

encapsulated under the skin and in the muscle 

on the top of the head of crevalle jack, 

occasionally in other muscles. It is also 

found under the skin of leatherjacket and in 

the stomach of swordfish. 

Length - Postlarva 5.0-18.0 mm, blastocyst 

anterior bulb up to 25 mm long. 

Host Specificity - The larva may be family 

specific to jacks and the adult to requiem 

sharks (Carcharhinidae). The rarity and low 

numbers, and location in the stomach suggest 

that swordfish may be a false host. 

Damage to Host - Six to 8 blastocysts on one side of the head occupy much 

of the muscle in this area, and may injure the host. 

Detection - Most of the blastocysts occur in the head where they are seldom 

seen by fishermen as this part of the fish is seldom cleaned or opened. If the 

head is cut apart, these large white capsules are prominent. 

125

126 


Gymnorhynchus gigas (Cuvier) 

Larvae of this parasite are encapsulated in inshore and offshore 


Diagnostic Characters - The plerocercus is attached to 

a large blastocyst with a long, thin tail. The 4 bothridia 

are elongate oval. The tentacles are relatively short and 

have long hooks with even longer hooks at the base. 

The tentacle sheaths are spiral. Tentacle bulbs are 

relatively short. The entacles have 9 hooks in each 

spiral row. 

Records - One plerocercus occurred in 1 of 4 white 

marlin and 1-9 (average 4) in 8 of 10 yellowfin tuna 

from the southern Gulf of Mexico; and rarely in 

swordfish from Europe. 

Geographic Range - North Atlantic. 

Life History - The adult is not known. 

Ecology - This parasite is encapsuled in ocean sunfish, 

rarely in swordfish, and commonly in yellowfin tuna. 

Many more records exist of this larva from inshore than 

offshore fishes. 

Location in Host - Muscle (swordfish) and blood vessels (yellowfin tuna). 

Length - Plerocercus 4.5-5.5 mm, blastocyst 24.0-125.0 mm. 

Prosobothrium armigerum Cohn 

This worm may only be an accidental parasite of big game fishes. 

Little is known about its postlarva. 

Name - Ichthyotaenia adhaerens (Linton) and P. japonicum Yamaguti 

are synonyms. 

Diagnostic Characters - The distinctive scolex has 4 relatively 

large, sucker-shaped bothridia which face forward (the circular flat 

faces are in the anteriormost plane). The suckers are cup-shaped 

from the sides (laterally) and are topped by circular disk-like pads. 

The spine-like structures (microtriches) that densely cover the neck of 

the adult are not apparent in the postlarva. The body of the postlarva 

is composed of a long narrow scolex and a short tail. 

Records - One postlarva occurred in 3 of 40 Atlantic blue marlin 

from various localities around Puerto Rico. 


Caribbean. 

Life History - The eggs are clustered in groups of 24 within a 

cocoon. 

Ecology - The adult of this tapeworm occurs in an offshore host, blue shark. 

Location in Host - Intestine (postlarva). 

Length - Postlarva 9.2 mm. 

Host Specificity - Atlantic blue marlin is a new host.


Ceratobothrium xanthocephalum Monticelli 

This little known plerocercoid commonly 

occurs in swordfish. 

Diagnostic Characters - The scolex is distinctive 

with 4 oval bothridia and relatively large, flat 

suckers on their anterior ends. The 2 

posteriorlateral corners of the suckers are armed 

with large muscular protrusions. The bothridial 

area makes up more than 1/2 to the entire length 

of the postlarva. 

Records - One to 99 postlarva (average 7.8) oc- 

curred in 112 of 303 swordfish from the northwest 

Atlantic (ARC 2315). It is also found in squids. 


Life History - The adult of this tapeworm occurs 

in sharks. 

Location in Host - Postlarva were found free in the intestine and rectum. 


Pelichnibothrium speciosum Monticelli 

This distinctive and widespread larva is 

encapsulated in tunas and other offshore pelagics. 

Name - The status of this species is uncertain. 

Phyllobothrium caudatum Zschokke and Heitz and 

P. salmonis Fujita are synonyms. 

Diagnostic Characters - The distinctive postlarva 

has a long body divided into a scolex with 4 

bothridia and 5 suckers; a neck with obvious 

segments, and an unsegmented tail region. Four 

of the suckers are on the anterior ends of the 

bothridia, and 1 sucker is on the anterior end of 

the scolex (apical sucker). 

Records - One to 2 postlarvae occurred in 5 

Atlantic blue marlin, and 1-4 in 6 of 40 dolphin from various localities around 

Puerto Rico. It was also encapsulated in 27 of 45 yellowfin tuna from northeast 

Gulf of Mexico (USNPC 56913); in bluefin tuna from Europe and the Pacific 

and in blue shark, another offshore pelagic fish, other fishes in the Pacific, and 

squids. 


Life History - This parasite apparently uses squids and fishes as intermediate 

hosts for its plerocercoids. This is a useful strategy in the offshore realm where 

hosts may be isolated and sparse. 

Ecology - Most of the known hosts of this parasite are offshore and pelagic 

which suggests it may be limited to this habitat. 

127

128 


Location in Host - Encapsulated in the wall of the stomach, outer layer of the 

intestine or viscera. 


Host Specificity - Atlantic blue marlin and dolphin are new hosts. 

Phyllobothrium delphini (Bosc) 

This larva is found in the stomachs of swordfish, but is better 

known for forming large capsules in marine mammals. 

Name - Hydatis delphini Bosc [=hydatid cysts of dolphins (mammal)]. 

It does not actually produce the dangerous hydatid cysts 

found in grazing land mammals. The species name has been 

treated as a larval-group name instead of a described species by 

some authors (Yamaguti 1959). Phyllobothrium loliginis (Leidy) 

from the stomachs of squids and swordfish may be a synonym. 

Diagnostic Characters - When the plerocercus is everted from 

the bladder, the blastocyst is shaped like a cherry with a stem. 

The scolex and neck are about as long as the attached bladder. 

The scolex has 4 membranous bothridia with thickened margins 

that are puckered and crumpled. The bothridia are triangular and 

each has a prominent, cup-shaped sucker on its anterior apex. The anterior end 

of the scolex is capped with a broadly rounded raised rostellum containing an 

apical sucker, which may disappear before the larva becomes an adult. 

Records - One to 102 capsules (average 7.0) occurred in 150 of 303 swordfish 

from the northwest Atlantic (ARC 2315). 


Life History - The adult form of this larva could be Phyllobothrium tumidum 

Linton a parasite of great white sharks, Carcharodon carcharias (Linnaeus), 

(which eat marine mammals around the world) and a variety of other sharks. 

Location in Host - Plerocercoids were found free in the stomach of swordfish 

and squids, but were encapsulated in the muscle of marine mammals. 

Length - Plerocercus 26.0-34.0 mm, encapsulated bladder 12.0-18.0 mm. 

Rhinebothrium flexile Linton 

Blastocysts containing this distinctive, chandeliershaped 

larva occasionally occur in big game fishes, but 

are found more often and in damaging numbers in 

inshore fishes. 

Name - It has been placed in genus Echeneibothrium. 

Diagnostic Characters - The blastocyst has a 

bulbous end containing the larva, a long string-like 

body and a smaller swelling on the posterior end. 

The postlarva removed from the capsule has a scolex 

with 4 elongate, oval bothridia on projections. The 

surface of each bothridium is divided in half by a long 

septum running down the middle, each half is then


further subdivided into elongate segments 

(loculi) by smaller, muscular cross-septa. 

Records - One to 14 postlarvae occurred 

in 12 of 40 dolphin from various 

localities around Puerto Rico. Numerous 

blastocysts occurred in 1 of 8 cobia from 

Beaufort, North Carolina, USA; and 1 

capsule in a chub mackerel from Woods 

Hole, Massachusetts, USA. The 

Rhinebothrium sp. of Arandas-Rego and 

Santos, 1983 in chub mackerel from 

Brazil may be the same species. 

Geographic Range - Worldwide. Our 

collections are the first in the Caribbean. 

Ecology - The plerocercoid occurs more commonly and in much higher 

numbers in inshore bottom fishes than in offshore, big game fishes. 

Location in Host - Encapsulated in intestine, liver and viscera. 

Length - Postlarva 1.0-1.2 mm, blastocyst 8.6-21.2 mm. 

Host Specificity - Encapsulated larvae occur in a variety of fish hosts, and the 

adults parasitize several species of rays. Dolphin is a new host. 

Damage to Host - Very heavy infections injure the viscera of inshore fishes 

and could damage big game fishes. 

129 

Tetraphyllid plerocercoid 

These tiny, white, nondescript larvae occur in the 

intestine of all marine fishes. Superinfections may 

harm swordfish. 

Name - Most plerocercoid larvae are undeveloped and 

cannot be identified beyond order. "Scolex pleuronectis" 

Mueller is a group name for these unidenti- 

fiable larvae. Scolex 

polymorphus Rudolphi 

and S. delphini Stossich 

are synonyms of this 

group name. 

Diagnostic Characters - It is a simple, white 

conical shaped plerocercoid with a bulbous scolex 

and a tapering tail. The 4 bothridia have pleated 

margins on the scolex which are difficult to see. 

The suckers on each bothridium are obvious in 

wet mounts viewed with a compound microscope. 

A terminal sucker usually occurs on the apex of 

the scolex. Live worms are highly active,

130 


contracting and relaxing, and may invaginate the scolex into the body. 

Records - These worms are found in all big game fishes and practically any fishes. 

We could list thousands of records, but this would be meaningless. For example, 

1-1900 occurred in 277 of 303 swordfish in the northwest Atlantic (ARC 2412). 


Location in Host - Intestine, stomach, pyloric ceca, bile duct and gall bladder. 

Length - Plerocercoid 0.4-2.0 mm. 

Damage to Host - Very heavy infections may block the bile duct or gall bladder. 

Large numbers in the stomach and intestine of hosts appear to have little effect. 

Superinfections have been reported in swordfish and in other fishes. 

Bothriocephalus scorpii (Mueller) 

This widespread parasite occurs in a variety of hosts, but 

rarely in big game fishes. 

Name - Leeuwenhoek named this worm in 1722 before the 

present system of scientific names was established. 

Diagnostic Characters - The immature worm has an elongate 

scolex with the lateral bothria becoming more shallow towards the 

posterior ends. The immature segments are as long as wide or 

longer. The plerocercoid is a flat, elongate worm that can only 

be identified by the presence of accompanying immature worms. 

More work is required before plerocercoid larvae of this genus 

can be easily identified to species. 

Records - One immature and 1 plerocercoid occurred in 1 of 3 

Atlantic mackerel from Woods Hole, Massachusetts, USA. 


Location in Host - intestine. 

Length - Immature 5.0 mm, plerocercoid 2.0 mm. 

Host Specificity - Plerocercoids and immature worms occur in a wide variety 

of hosts. 

Bothriocephalus sp. of Linton 

The encapsulated larva occasionally occurs in big game fishes. 

Name - The records below probably represent more than 1 species. Little 

effort has been made to distinguish these rather formless larvae. The records 

may include larvae of B. manubriformis and B. scorpii. 

Diagnostic Characters - These plerocercoid larvae are flat, vase-shaped, flaskshaped 

or slender worms with calcarious bodies in their tissue. The scolex is 

simple, has 2 lateral grooves and is covered with minute spine-like projections 

(microtriches). 

Records - One and a few encapsulated in 2 of 38 Atlantic bonito (USNPC 

4785,6528), 1 to a few capsules in 8 of 192 Atlantic mackerel (USNPC 8890), 

1 in a greater amberjack (USNPC 8891), and 5-12 capsules in 12 chub mackerel 

from Woods Hole, Massachusetts, USA.


Location in Host - These larvae were encapsulated in the mesenteries or outer 

layers of the viscera, in the wall of the stomach or intestine, or occasionally 

loose in the intestine where they were digested out of prey. 

Length - plerocercoid 0.4-13.0, capsule 1.0-8.0 mm. 

Miscellaneous Tapeworms 

Bothriocephalus janikii Markowski - This worm was described from 

specimens collected from the stomach of a dolphin during the Discovery Cruise 

apparently in the south central Atlantic (2 latitudes and no longitude given?); and 

it was reported in the intestine of dolphin from the Bay of Bengal off India. 

Schmidt (1986) did not list this tapeworm. 

Echeneibothrium sp. of Nikolaeva - The primary characteristic of this 

genus and its plerocercoid is the presence of 4, elongate, oval bothridia that are 

distinctly divided by ridges into longitudinal rows of 1 or 2 rectangular 

compartments. The whole bothridia resembles the sucker disks of remoras 

(genus Echeneis). Three capsules with plerocercoids occurred in 1 of 10 

yellowfin tuna from the southern Gulf of Mexico and plerocercoids of 

Echeneibothrium sp. of Rego, Carvalho-V., Mendonca and Afonso-R., 1985, 

occurred in less than 10% of 80 Atlantic mackerel from Portugal. Adults 

usually occur in skates. This parasite has rarely been found in big game or any 

fishes. The normal intermediate hosts for the plerocercoids may be 

invertebrates, and fishes may be accidental hosts. More collections are needed 

to identify this form to species and to determine their importance in big game 


Incompletely defined trypanorhynchid larval species – The species 

listed below have been named from big game fishes, but so few details were 

presented that they cannot be identified to genus or species. These names cannot 

and should not be used. They are only presented to prevent confusion, if they 

are seen in the literature. (1) Bothriocephalus claviger Leuckart - This worm has 

also been called "Tetrarhynchus" claviger (Leuckart), and is found in swordfish. 

(2) Dasyrhynchidae of Ward (1962) - Capsules were found in 21 of 45 

yellowfin tuna from the northeast Gulf of Mexico (USNPC 56911). (3) 

Dibothriorhynchus speciosum MacCallum - Up to 20 capsules occurred in 

Spanish mackerel and many other fishes from Bermuda. Plerocercoids were 

20.0-22.0 mm long. (4) Rhynchobothrium ambiguum Diesing - This worm was 

found in swordfish. (5) Rhynchobothrium longispine Linton - Several and 1 

capsule in 2 of 11 Atlantic mackerel and 3 in 1 of 3 Spanish mackerel from 

Woods Hole, Massachusetts, USA; and possibly in cobia from Beaufort, North 

Carolina, USA. It is 0.8-3.8 mm long. (6) Tetrabothriorhynchus scombri 

Diesing - in Atlantic mackerel. (7) Tetrarhynchus papillosus Rudolphi - in 

dolphin. (8) Tetrarhynchus scomber-pelamys Wagner - in Atlantic bonito. (9) 

Tetrarhynchus scomber-rocheri Wagner - in bullet tuna or frigate tuna. (10) 

131

132 


Tetrarhynchus scomber-thynnus Wagner - in bluefin tuna. (11) Tetrarhynchus 

thynni Wagner - in tuna. 

Pseudobothrium grimaldii Guiart - This adult (?) tapeworm was found 

out of the geographic range of this book in an albacore from the eastern Atlantic 

off the Azores. Both the genus and species are in question. We include it 

because: (1) the existence of this worm is not mentioned in modern guides to 

tapeworms and it is thus being forgotten; (2) few adult tapeworm species have 

ever been found in big game fishes and this worm could be very interesting; and 

(3) because we would like someone to find adult tapeworms in albacore. The 

description of this worm was inadequate to distinguish the genus or species. 

The body was badly contracted. It apparently had no attachment organs on the 

scolex. The posterior margin of the segment (proglottid) overlaps the anterior 

margin of the next proglottid (craspedote condition), and the posterior border has 

10-12 pleats. It was about 55.0 mm long. 

Sphyriocephalus sp. - This larval parasite could possibly be the same as S. 

dollfusi Bussierae and Aldrin which has been found in bigeye tuna in the eastern 

Atlantic. Postlarvae have a scolex that is thicker than wide and characterized 

by a deep bothridial cavity. The 2 ventral and dorsal bothridia are fused 

together around the cavity. The postlarva looks like a rather thick ping pong 

paddle. The tentacles emerge through the bothridia cavities and look something 

like cacti in a planter. The tentacles, tentacle bulbs and sheaths are all short. 

Two and 6 blastocysts were encysted in the viscera of 2 of 10 yellowfin tuna 

from the southern Gulf of Mexico and capsules occurred in 14 of 45 yellowfin 

tuna from the northeast Gulf of Mexico (USNPC 56908). 

trypanorhynchid plerocercoid - Most of these plerocercoid larval 

forms in fishes can be identified, at least to genus, because the scolices are well 

formed and they have 4, distinctive, hook-covered tentacles. Some forms may 

lack characters for identification beyond order, particularly those that are less 

developed, badly contracted during preservation, or degenerated in capsules. 

The best way to resolve these identification problems is to obtain a larger series 

of specimens to find more mature plerocercoids in better condition.

NEMATODA (ROUNDWORMS) 

133 

Roundworms, threadworms or nematodes are a phylum. They, along with 

flatworms and spiny-headed worms, are sometimes called "helminths". 

Roundworms cause serious diseases and even death in humans. The recent 

popularity of Japanese raw-fish dishes has caused an increase in the number of 

fish-roundworm-related illnesses around the world. Modern refrigeration of fish 

catches has also allowed dangerous roundworms, that would have been discarded 

by quick cleaning, to migrate from the gut and mesenteries into the edible flesh. 

The unwise game of "live fish 

swallowing" has produced severe 

gastric distress in humans when 

roundworms from fish burrowed 

through the intestinal wall into the 

body cavity of humans. Treatment 

requires surgical removal. One 

case is known of a roundworm from 

the flesh of a Hawaiian jack 

penetrating a wound in the hand of 

a man cleaning the fish. The entry 

was painful and the worm could 

only be removed with surgery. 

More than 12,000 species are 

described of the more than 500,000 

to 2 million that probably exist. 

They are one of the most abundant 

multicellular organisms on earth, 

both in total numbers of individuals 

and in number of species. Roundworms 

occur in such high numbers 

in almost all vertebrates and 

invertebrates that it has been suggested 

that the shapes of all living 

animals could be seen from space 

by merely seeing the mass of 

worms in each animal. 

Most free-living forms are 

small to microscopic, but parasitic forms are large, up to 8 meters long. Roundworms, 

as the name implies, are circular in cross-section. The body is non-segmented, 

elongate and slender, often tapered near the ends, and covered with 

cuticle. Three to 6 lips surround the mouth. The digestive tract is complete. 

Musculature has longitudinal fibers. A pseudocoel (false body cavity) is present. 

A nerve ring is usually visible in the anterior end. Sexes are separate. The

134 


male has a cloaca, a pair of chitinized copulatory structures 

(usually spicules), often with a variety of papillae, alae (long, 

thin flaps of cuticle), suckers in or on the posterior end. All of 

these male structures are important in identifying species. Eggs 

are released into the intestine of fishes, or through holes in the 

host skin in tissue-dwelling roundworms, into the water. Eggs 

may contain developed roundworms, while others are expelled 

while less developed. Some larvae are eaten by fishes and 

develop directly into an adult. Usually, larvae must go through 

2-5 molts in 1 or more crustacean and/or fish intermediate 

hosts. They are found in all marine, freshwater and terrestrial 

habitats. Flying insects, birds and bats take them into the skies. 

None of the roundworms found in Puerto Rican fishes are 

known to normally infect humans. Nevertheless, consuming 

live roundworms is ill advised as these parasites may cause 

considerable gastric distress or even penetrate the wall of the 

stomach or intestine. Humans in the continental USA have been 

infected with roundworms (anisakiasis) from raw skipjack tuna 

and yellowfin tuna. Any of the anisakids could potentially harm 

humans. Those found in the muscle of edible sport fishes are 

of particular interest as they are likely to be consumed by 

humans. Thorough cooking or freezing for several days kills all 

roundworms in muscle. If you must eat raw-fish dishes, make 

certain they are prepared by a professional chef who has been 

trained to avoid parasites. Most cases of anisakiasis in the USA 

have been the result of do-it-yourself preparations. The 

Japanese are thought to be the leading experts and demand the 

best quality fish, but they freeze most raw fish long enough to 

kill roundworms. 

A necropsy is needed to find the larvae in the organs and 

mesenteries and the adults in the stomach and intestine. Roundworms 

can be relaxed in acetic acid and stored in a mixture of 

70% ethanol with 5% glycerine. Adult roundworms from 

tissues of fishes are exceedingly delicate and tend to explode if 

placed in fresh water or preservatives. These small worms can 

be placed in a steaming 0.8% saline and 5% formalin solution. 

Once fixed (15 minutes for small worms up to 24 hours for 

large ones), worms can be rinsed in fresh water, and slowly 

transferred into gradually increasing concentrations of ethanol, 

until stored in a mixture of 70% ethanol and 5% glycerine.


Roundworms are usually examined in wet mounts. Semipermanent mounts may 

be prepared using glycerine jelly. In most larval species, identification to 

species is very difficult, but most genera can be determined. Study of the entire 

life cycle of a roundworm may be necessary to determine the species of larvae. 

No treatment is possible for roundworms in the body cavity or tissues of 

fishes and is seldom necessary for intestinal roundworms. Worms that perforate 

the intestine of humans must be surgically removed. 

Most roundworms in fishes are rather similar in shape. Thus the outlines 

of their entire bodies are not of much use in telling them apart. These worms 

are distinguished by relatively small structures on their anterior ends and on the 

posterior end of males. We illustrate a whole female Hysterothylacium aduncum 

here to show the general shape, but only illustrate enlargements of the anterior 

end and the posterior end of the male for each species described. 

Reference - Anderson, Chabaud and Willmott (1974-83) is an excellent 

taxonomic guide to genera and higher classification of roundworms, and Bruce, 

Adlard and Cannon (1994) a taxonomic guide to species of fish-parasitic 

ascaridoids (anisakids and ascarids in big game fishes), but no general guide to 

all the fish roundworms has been crafted since Yamaguti (1961). 


Phylum Nematoda - roundworms Page 

Class Enoplea (Adenophorea) 

Order Enoplida 

Family Cystoopsidae 

Cystoopsis scomber ....................................................................... 154 

Class Rhabditea (Secernentea) 

Order Strongylida 

Family Ichthyostrongylidae 

Ichthyostrongylus thunni ................................................................ 154 

Order Ascaridida 

Family Ascarididae 

Genus Ascaris ....................................................................................... 153 

Porrocaecum paivai ...................................................................... 155 

Family Anisakidae 

Anisakis simplex* ........................................................................... 136 

Genus Contracaecum* ......................................................................... 153 

Goezia pelagia ................................................................................ 138 

Genus Hysterothylacium ...................................................................... 139 

Hysterothylacium aduncum ........................................................... 141 

Hysterothylacium cornutum .......................................................... 142 

Hysterothylacium corrugatum ...................................................... 143 

135

136 


Hysterothylacium fortalezae ........................................................ 144 

Hysterothylacium pelagicum ....................................................... 145 

Hysterothylacium reliquens ......................................................... 146 

Iheringascaris inquies .................................................................. 147 

Maricostula histiophori ............................................................... 148 

Maricostula incurva .................................................................... 149 

Maricostula sp. ............................................................................ 150 

Raphidascaris anchoviellae ......................................................... 155 

Family Cucullanidae 

Cucullanus carangis ..................................................................... 153 

Cucullanus pulcherrimus .............................................................. 153 

Order Spirurida 

spiruroids ....................................................................................... 155 

Family Camallanidae 

Oncophora melanocephala ........................................................... 151 

Family Philometridae 

Philometra sp. ………………........................................................ 154 

Philometroides sp. .......................................................................... 154 

Family Cystidicolidae 

Ctenascarophis lesteri .................................................................... 151 

Parascarophis galeata ................................................................... 154 

Prospinitectus exiguus .................................................................... 152 

Miscellaneous Roundworms ........................................................... 153 

_______ 

*Larval forms 

Anisakis simplex Rudolphi 

These tiny, inconspicuous worms 

cause severe pain, nausea, vomiting and 

can rarely kill humans. But this 

dangerous parasite can be easily avoided 

even in raw-fish dishes. 

Name - This name has variously been 

used for a presumably single, widespread 

species in the northern Atlantic, similar 

species or any Anisakis sp. larva. We 

assume the "Anisapis sp." of Chen and 

Yang, 1973 in tunas off Taiwan was a 

typographical error for Anisakis sp.,


otherwise it would be a rather amusing roundworm. The disease this worm 

causes in humans is called "anisakiasis" or "sushi disease". 

Diagnostic Characters - A cream-colored cyst encompasses the larvae which 

is coiled in a tight spiral (as are most other larval roundworms encysted in fish). 

When freed from the cyst, the anterior end of the worm has 3 relatively small, 

inconspicuous lips with an obvious, center off-set, forward projecting tooth 

(anteroventral projecting boring tooth). The relatively long esophagus is 

followed by a glandular region. There is no appendix or caecum. It has a 

bluntly rounded tail. 

Records - One to 9 larvae occurred in 63 of 303 swordfish examined in the 

northwest Atlantic (ARC 2323). It has also been found in Atlantic bonito, 

Atlantic mackerel, skipjack tuna and yellowfin tuna off the northeast coast of the 

USA. This worm probably encysts in all big game fishes in this region. The 

larval Anisakis sp. reported in chub mackerel from southern Brazil may be this 

worm. 

Geographic Range - Worldwide. It is well known in the North Atlantic from 

North America to Europe, and similar forms occur the north Pacific and in the 

cooler waters of the southern hemisphere. 

Life History - Adults parasitize the gastrointestinal tract of dolphins (mammal), 

seals and whales. Eggs pass out of the intestine of the host. Second stage 

larvae escape from the egg in a few days. This free-swimming stage may be 

eaten by a crustacean transport host in which no development will occur, until 

this host is eaten by an appropriate intermediate crustacean or fish host, or it 

may be eaten directly by an intermediate host. The larvae, a few millimeters 

long, migrates to the hemocoel (crustacean) or mesenteries (fish), is soon 

encapsulated and develops into a third-stage larvae. When the appropriate 

marine mammal final host eat the intermediate host, the worm is digested out 

and develops into an adult. Third-stage larvae can infect humans. Larvae from 

Atlantic herring have been raised to adults in laboratory culture media without 

hosts. 

Ecology - In the Atlantic it is limited to the colder regions (not in the Gulf of 

Mexico or the West Indies). However, in the Pacific it occurs from cold waters 

to tropical Hawaii and Australia. 

Location in Host - These worms are usually found in the mesenteries around 

the gut and internal organs. Holding dead, fish, that have not been cleaned 

(eviscerated), on ice or under refrigeration allows these larvae to migrate into 

the flesh of the host. 

A European herring fishery was converted from small boats with immediate 

cleaning practices to large vessels with refrigerated storage and shore 

processing. This gave the worms an opportunity to invade the flesh and 

widespread outbreaks of anisakiasis occurred in people consuming raw-herring 

dishes. Adequate freezing of the herrings before sale solved the problem. 

Length - 11.2-34.5 mm. 

Host Specificity - It is most common in inshore fishes and sometimes infects 

almost all fishes. 

137

138 


Damage to Host - Heavy infections of larvae in fishes can cause severe 

inflammation and death. 

Detection - Once a fish is killed and held on ice, these larvae borrow out of 

their cysts and migrate into the muscle. These worms are relatively small and 

cream colored which makes them difficult to find in muscle. Studies have 

estimated that industrial techniques for detecting these and other larger worms 

in fish fillets can only find about half the worms. 

Harm to Humans - Juveniles of this worm survive in refrigerated, iced, 

inadequately frozen, salted and pickled fish flesh. They penetrate the stomach 

of humans 1-12 hours after infected seafood is eaten, or may penetrate the 

intestine up to 14 days later. The penetrations usually cause sudden and severe 

pain in the upper stomach or abdomen. Usually nausea and vomiting are 

associated with the severe pain. Anisakiasis in the stomach can be diagnosed 

with an endoscope and the worms can be removed with biopsy forceps, if the 

surgeon is sufficiently skillful to catch these fast-moving and lively parasites. 

Worms that pass further down the intestinal tract cannot be so easily diagnosed 

and removed. In some cases penetration may occur without symptoms or with 

mild pain. Once the worms penetrate into the digestive tract wall or into other 

organs in the body cavity, diagnosis is more difficult as the symptoms causes by 

these parasites are similar to other, more common diseases. Immunological tests 

are available. Surgery may be necessary in some cases. 

Experimental infections in monkeys are successful if the larvae enter an 

empty stomach, but not when the stomach contains food. This may explain the 

higher Japanese vs. Chinese infection rates, as raw fish is traditionally consumed 

in the beginning of the meal in Japan, but at the end in China (Williams and 

Jones 1976). 

These worms can live for 51 days in vinegar, 6 days in 10% formalin (!), 

but only 2 hours in a home freezer (-20ΕC). Fish flesh should be frozen for 24 

hours prior to being used in raw-fish preparations (Pinkus, Coolidge and Little 

1975, Hilderbrand 1984). Larger and thicker fish may take longer to freeze and 

should be held 4-5 days in inefficient or over-filled freezers. This temporary 

freezing has very little effect on the flavor or texture of fresh fish, but kills 

dangerous roundworms. Thorough cooking kills all parasites. 

Significance to Sportfishing - The larvae of this parasite may occur in 

sufficient numbers in Atlantic mackerel to injure, stunt or kill these fish. They 

are not known to occur in high numbers in other sport fishes. 

This larvae has been evaluated as a potential biological tag for Atlantic 

mackerel and other fishes. Its abundance was used to show movement of 

albacore from New Zealand to the central south Pacific (Jones 1991). 

Goezia pelagia Deardorff and Overstreet 

This spiny burr of a parasite is more impressive in life than our drawings 

suggest. It is highly irritating and damaging to host fishes, but may be useful 

as a biological tag. 

Name - The name "pelagia" means of the sea.


Diagnostic Characters - This 

worm has spines covering most of its 

body. It has over-hung lips on the 

anterior end, and a conical tail with 

a finger-like projection. Male 

spicules are approximately equal in 

length. Males have 12-19 pairs of 

papillae anterior of the anus; 2 pairs 

at the level of the anus, and 

posterior to the anus there are 4 

pairs and the third pair is doubled (2 

papillae jammed together in the 

row). The vulva of the female 

opens 29-55% of the total length of 

the worm from the anterior end. 

Records - Worms occurred in cobia (USNPC 75680-2) and other fishes off 

Alabama, Mississippi and Louisiana, USA. 

Geographic Range - Northern Gulf of Mexico. 

Ecology - These worms feed on both the blood of the host and partially digested 

food in the stomach of the host. 

Associations - Several Iheringascaris inquies occurred in 1 capsule with this worm. 

Location in Host - Free but firmly attached in the stomach or embedded and 

encapsulated in stomach wall. 

Length - Female 3.6-14.5 mm, male 3.4-12.0 mm. 

Host Specificity - This worm probably has little host preference, since the 3 

host species are found in 3 families and 2 orders of fishes. 

Damage to Host - Various amount of tissue penetration and host reaction 

occurred with encapsulated worms. Some worms in a capsule were 

degenerated. Extensive bacterial and fungal infections occurred inside the 

capsules. Inflammation and host reaction was limited to the area immediately 

around the capsules, and the capsules seemed to confine microbial infections and 

further worm penetration. Other worms in this genus have caused mass 

mortalities in fishes stocked in freshwater lakes. Some species elicit extensive 

host reaction. Deardorff and Overstreet (1980) suggested that these worms can 

detrimentally affect sport fishes. 

Significance to Sport Fishing - This worm may be a good biological tag as 

it is easy to recognize, has a limited geographic range and survives in capsules 

for several years. 

Hysterothylacium Ward and Magath 

Many species of roundworms that have incorrectly been placed in the genera 

Contracaecum and Thynnascaris Dollfus (thynnos=tuna, ascaris=worm) belong 

in the genus Hysterothylacium (hysteros=after, thylaco=bag, and refers to the 

139

140 


cecum). Thynnascaris was a marvelous name for parasites of big game fishes, 

especially tunas, but unfortunately it is not taxonomically correct for these 

worms. Deardorff and Overstreet (1981) resurrected and redescribed this genus, 

moved 55 species into it and detailed those species found in the Gulf of Mexico. 

Bruce and Cannon (1989) re-redescribed it and split off the new genus 

Maricostula. Bruce, Adlard and Cannon (1994) listed 65 species in Genus 

Hysterothylacium. The exact number cannot be determined as only a few of 

these species have been adequately described. 

Larval forms of this genus encyst in fish and are similar in appearance to 

Contracaecum spp. larvae. They differ by having the excretory pore at the level 

of the nerve ring in Hysterothylacium spp. and near the lips in Contracaecum 

spp. Members of both genera have the offset boring tooth similar to Anisakis 

spp. larvae, but differ by having a distinct caecum. A complete life cycle is 

only known for Hysterothylacium aduncum, but the other species in big game 

fishes probably have similar cycles. Eggs are laid in the intestinal tract of the 

host and are shed into the environment. Second stage larvae 0.3-0.5 mm long 

hatch from the eggs. They can stay alive without a host from 2 weeks to several 

months. Once ingested by a copepod, the larvae grow to 0.4-0.7 mm long. A 

small fish may be able to serve as a host for this stage instead of a copepod. 

When the copepod is eaten by a fish, the second stage larvae is digested out of 

the copepod, penetrates the intestinal wall of the fish, and encapsulates and 

grows to 1.0-4.0 mm long. The process may be repeated several times in 

ascending sizes of fishes (transport host), if other fishes eat the infected fish 

before the larvae has sufficiently developed, or if the host eating the infected fish 

is not the appropriate final host. Eventually, the third stage larvae grows to 22 mm 

long and its intermediate host is eaten by the appropriate final host (or the 

larvae dies of "old age" or is finally overcome by the defenses of the host). In 

the gut of the final host, the larvae is digested out of its intermediate host, 

attaches in the stomach and develops into an adult. 

Heavy infections of larvae in fishes can cause severe inflammation and 

death. Overstreet and Meyer (1981) found that a larval form of this genus, 

common in the northern Gulf of Mexico, was capable of penetrating the stomach 

and causing hemorrhage in monkeys that ingested these worms. The flesh of 

dolphins, Spanish mackerels, swordfish and tunas may become contaminated 

with this worm if they are not cleaned quickly. This is particularly true for 

uncleaned fishes held for hours on ice or under refrigeration. Most of these 

worms in the gut of the final host are either young, immature or larval forms. 

When the host dies these forms can easily penetrate the gut wall and pass into 

the flesh. These fish should be adequately frozen before being used in raw-fish 

dishes. These larvae occur in a great variety of invertebrates including 

commercial shrimp. Twelve hours in a home freezer (-20ΕC) kills these 

roundworms (Norris and Overstreet 1976), but more time may be necessary to 

kill these larvae in fish tissues. Thorough cooking kills all parasites.


Hysterothylacium aduncum (Rudolphi) 

A general and widespread 

parasite of inshore fishes. 

Apparently a few adults and 

encysted larvae are transferred to 

big game fishes through feeding on 

inshore ones. Larval forms can 

infect humans. 

Name - This species could be a 

complex of several similar species. 

Hysterothylacium longispiculum 

(Fujita) and Thynnascaris adunca 

(Rudolphi) are synonyms. 

Diagnostic Characters - The alae begin just posterior to the lips. The lips are 

wider than long, lack interlabial grooves, are not indented in their sides, and 

taper toward the anterior margin. There is a deep constricted at the base of the 

lips. The tail gradually tapers and ends with a patch of small caudal spines 

(called "cactus-tail" by some authors). The cervical alae are narrow. Males 

have 2 pre-anal pairs of papillae, 2 pairs of papillae at the same level as the 

anus, and 5 pairs posterior to the anus. The spicules are approximately equal 

in length. The female has a vulva opening in the first 30-50% of the body 

length. The encysted, third stage larvae may be identified by the cactus tail. 

Records - One to 21 worms occurred in 26 of 303 swordfish from the 

northwest Atlantic (ARC 2410); in Atlantic mackerel and white trevally off the 

Atlantic coast of the USA, in northern bluefin tuna off Europe, and in other 

jacks around the world. Encysted larvae were found in northern bluefin tuna 

and swordfish off Europe and in albacore in the southwest Pacific. Jones (1991) 

found larvae of this worm in 0-33% of over 400 albacore sampled from 13 

locations in the tropical to temperate southwestern Pacific. High prevalence (27- 

33%) occurred in both tropical and temperate areas. Intensity increased with 

increases in host size. Adults and larvae have been reported from a great 

variety of fishes. 

Geographic Range - Worldwide in temperate and cooler waters. 

Life History - See description in the genus. Eggs hatch in 4-12 days. Larvae 

have experimentally developed in benthic polychaetes, planktonic copepods and 

fish fry, but are only commonly found in polychaetes in the wild. Sometimes 

as many as 13% of numerous polychaete species sampled are infected with 1, 

or rarely 2, larvae (once 150). Jones (1991) suggested that planktonic krill 

(euphausiids), plankton-associated hyperiid amphipods and squids were 

intermediate hosts in the southwest Pacific. More than 100 species of 

invertebrates in 7 phyla have been reported as intermediate hosts. This wide 

range of hosts may help to explain the great abundance and broad distribution 

of this parasite. 

Ecology - It is usually found in inshore, benthic hosts. This parasite probably 

only occurs in offshore fishes that acquire them from eating inshore fishes. In 

141

142 


Europe it is the most common larval roundworm encysted in inshore fishes, 

occurring in almost every fish in some areas. 

In contrast, Jones (1991) reported that this larval worm was more common 

(27-33%) in open oceanic situations east of New Zealand, than near island 

groups and Australia, in both the temperate and tropical regions of the southwest 

Pacific. The worms he studied may be a similar, but different species as it 

occurs strictly in temperate and colder waters in the Atlantic, and does not occur 

in the West Indies or the Gulf of Mexico. 

Associations - This worm occasionally occurred in low numbers with 

Hysterothylacium corrugatum and Maricostula incurva in swordfish in the 

northwest Atlantic, and with Hysterothylacium cornutum in bluefin tuna off 

Europe. It encysted in fish with Anisakis simplex in the southwest Atlantic and 

probably does so in the north Atlantic. 


Length - Female 16.0-94.0 mm, male 14.0-37.0 mm, third stage larva 5.9-21.6 

mm (the cyst is smaller because the worms are coiled). 

Host Specificity - It apparent has little host preference. This parasite has been 

reported from more than 100 fish species, although some of these records were 

probably in error. It is possible that big game fishes are only false hosts. 

Harm to Humans - Williams and Jones (1976) document cases of human 

infection (eosinophilic granulomata) by larvae of this worm. 

Significance to Sport Fishing - This worm apparently occurs only in cold- 

water areas. It might be useful as a biological tag of big game fishes moving 

into warmer regions. The adults may only survive a few months or a year, but 

encysted larvae should be available for a longer period of time. Jones (1991) 

was able to easily identify the encysted larvae in albacore in the southwest 

Pacific, but this biological tag was not diagnostic of movements in his study. 

Hysterothylacium cornutum (Stossich) 

Light infections probably do little 

damage to tunas around the world. 

They do pose a threat to humans who 

eat these popular and pricey fish raw. 

Name - It was redescribed by Bruce 

and Cannon (1989). Thynnascaris 

legendrei Dollfus is a synonym. 

Diagnostic Characters - The alae 

begin at the base of the lips. The lips 

are slightly shorter than wide. The tail 

is bluntly rounded with a prominent 

acute process. Males have 23-32 preanal 

papillae, 2 papillae at the level of 

the anus, and 9-13 (usually 9) papillae


posterior to the anus. There are obvious striations (modified ventral annuli) across 

the body in the area of the papillae. The spicules are approximately equal in 

length. The female vulva opens in the anterior 13.3-29.0% of the body length. 

Records - One female and 2 males occurred in an albacore and 3 females and 

a male in blackfin tuna off Puerto Rico (USNPC). It has also been found in 

albacore, bluefin tuna and yellowfin tuna in the Atlantic; and these fishes and 

additional species of tunas in the Indo-Pacific. Deardorff and Overstreet (1982) 

found this worm in yellowfin tuna off Hawaii, but did not find it in 19 yellowfin 

tuna from the northern Gulf of Mexico. Ward (1962) also failed to locate it in 

45 yellowfin tuna from this region. The Contracaecum sp. reported in the 

stomach and intestine of yellowfin tuna in the southern Gulf of Mexico by 

Nikolaeva (1968) could have been this worm. 

Geographic Range - Worldwide. Our collections are the first in both the 

western Atlantic and the Caribbean. 

Ecology - Pelagic, open ocean. 



Host Specificity - It is genus specific (Thunnus). Records from fishes other 

than tunas were other species of roundworms. Blackfin tuna is a new host. 

Harm to Humans - The relatively new and expensive flying of fresh tunas into 

Japan from around the world, should provide an ample opportunity for these 

worms to contaminate the flesh. 

Significance to Sport Fishing-If this worm occurs in the Caribbean, but not 

in the Gulf of Mexico; and in the northeast but not the northwest Atlantic, then 

it might be useful as a biological tag. 

Hysterothylacium corrugatum Deardorff and Overstreet 

This large parasite of swordfish occurs sparsely 

in the western Atlantic and eastern Pacific. Both the 

mystery of its unusual distribution and how it 

maintains such low numbers are worthy of investigation. 

Name - The name "corrugatum" means ridged and 

refers to the modified annuli on the posterior, 

ventral surface of the male. This is a useful, but 

not exactly a unique character among these worms. 

Diagnostic Characters - The alae begin just pos- 

terior to the base of the lips. The lips are distinctly 

longer than wide, are constricted in the middle, lack 

deep interlabial grooves and are not on peduncles. 

The tail gradually tapers. Males have 24-26 pairs of 

papillae anterior to the anus, 1 pair at the same level 

of the anus, and 4 pairs posterior to the anus. 

There are obvious striations (modified ventral 

annuli) across the body in the area of the papillae. 

143

144 


The spicules are approximately equal in length. The female vulva opens in the 

anterior 33.0-39.0% of the body length. 

Records - A light infection occurred in 1 of 2 swordfish caught off La 

Parguera, Puerto Rico (USNPC); 6 females and 5 males in a swordfish from 

Miami, Florida (USNPC 75844-46); from Panama City, Florida, USA; up to 99 

worms, but usually many fewer (mean 12.5), in 208 of 303 swordfish from the 

northwest Atlantic (ARC 2311); and in the eastern Pacific. 

Geographic Range - Tropical to temperate western Atlantic and tropical 

eastern Pacific. Our collection is the first in the Caribbean. 


Associations - It occurred with Maricostula incurva in swordfish off Puerto 

Rico, south Florida, and in the northern Gulf of Mexico; and with M. incurva, 

Hysterothylacium aduncum and H. reliquens from the northwest Atlantic. 



Host Specificity - It only occurs in swordfish and is a secondary parasite. 

Damage to Host - This large worm may stunt or injure swordfish particularly 

when it occurs in heavy infections of almost 100 worms. It may also contribute 

to severe injury to swordfish when they are superinfected with thousands of 

Maricostula incurva. 

Significance to Sport Fishing - Parasites extract a heavy toll on this highly 

important sport and commercial fish. Their role in its ecology must be better 

understood. If the range of this worm is as restricted as our current knowledge 

suggests, then it might be of use as a biological tag. 

Hysterothylacium fortalezae (Klein) 

This rather small worm occurs in 

low numbers in tropical and subtropical 

western Atlantic Spanish mackerels. 

Diagnostic Characters - The alae 

extend the entire length of body and are 

prominently expanded into wings the 

anterior and posteriorly ends of the 

worm. The lips are as long as wide and 

have no interlabial grooves. A tuft of 

12-14 projections occurs on the end of 

the tail. Males have 13-25 pairs papil- 

lae anterior to the anus, no papillae at 

the level of the anus, and 8 post-anal 

pairs. The female vulva opens in the 

anterior 30.0-38.0% of the body length.


Records - We found 1-2 in 3 of 35 cero and 1-5 in 8 of 9 king mackerel from 

La Parguera, Puerto Rico (USNPC); and 8-11 in 2 of 4 king mackerel and 2-24 

in 3 of 9 Spanish mackerel from Dauphin Island, Alabama, USA (USNPC). 

This worm also occurred in Spanish mackerel from Florida and Mississippi, and 

in other inshore fishes from Mississippi, USA; and in king mackerel and serra 

Spanish mackerel from Brazil. 

Geographic Range - Atlantic and Mediterranean. Our collections are the first 


Location in Host - Stomach and intestine. 

Length - Female 10.0-15.4 mm, male 12.0-23.4 mm, female fourth-stage larva 

encysted in fish 5.0-13.5 mm, third-stage larva 1.7-3.5 mm. 

Host Specificity - Spanish mackerels are the preferred hosts, but it is not 

genus specific (Scomberomorus). Cero is a new host. 

Hysterothylacium pelagicum Deardorff and Overstreet 

A common, yet little known, stomach and 

intestinal parasite of dolphin around the world. 

More fish are parasitized by this worm in the 

tropics than in the temperate areas. 

Name - The name "pelagicum" means of the sea. 

This species was confused with Hysterothylacium 

cornutum until it was separated by Deardorff and 

Overstreet (1982). It was redescribed by Bruce 

and Cannon (1989). Ascaris increscens of Linton 

and Hysterothylacium sp. of Deardorff and 

Overstreet are synonyms. Contracaecum [sp.] 

reported as extremely abundant in numerous 

dolphin stomachs caught off North Carolina by 

Rose (1966) was probably this worm. 

Diagnostic Characters - The alae begin just 

posterior to the lips. The lips are shorter than 

wide, and have deep interlabial grooves. There is a tuft of 12-14 projections on 

end of the tail. Males have 36 pairs of papillae anterior of the anus, 3 pairs at 

the level of the anus, and 11-12 pairs posterior to the anus. It does not have 

body striations (modified ventral annuli) in the area of the papillae. There is a 

tiny round ball at the tip of the tail which looks like a nipple or a spinous 

process. The female vulva opens in the anterior 30-40% of the body length. 

Records - We found 1-31 (average 5.8) in 23 of 40 dolphin from various 

locations around Puerto Rico (USNPC); and 1-20 in 2 of 5 dolphin from 

Dauphin Island, Alabama, USA. It also occurs in this host off the Gulf and 

Atlantic US coasts from Texas to North Carolina, including the Florida Keys; 

and 1-35 (mean 10) were found in 19 of 33 dolphin from Hawaii. It has also 

been found in black marlin from the Pacific, but this could represent 

roundworms from dolphin consumed by this host. 

145

146 


We identified all the gastrointestinal parasites in 13 dolphin collected off La 

Parguera, Puerto Rico. Twelve (92.3%) were infected with 1-12 males (average 

1.6) and 1-44 females (average 11.9) of this roundworm. Manooch, Mason and 

Nelson (1984) examined the stomachs of 2632 dolphins in 10 sample areas from 

Texas to North Carolina, USA. They found 132 (5%) of these fish had from 1-100 

of this worm, but most fish had few worms (mean 11). They also found more 

stomach parasites in larger fishes, but this was not apparent in our 13 fish. They 

only looked at stomachs, while we examined the entire tract. They also found few 

flukes, while we found more than 2800 in a single stomach. This suggests that 

Caribbean dolphin are much more heavily parasitized than those off the Gulf and 

Atlantic coasts of the USA. 

Geographic Range - Worldwide. Our collections are the first in the Caribbean. It 

has been listed from the Caribbean in a summary article, but this was mistakenly 

based on the Gulf of Panama record (eastern Pacific). 


Associations - This roundworm occurred together with and in similar numbers to 

a fluke, Dinurus tornatus, in 13 dolphin we examined (The abundance of this 

roundworm was strongly positively correlated at a 99% confidence level with the 

presence of the fluke). Seven males and 4 females of Maricostula makairi Bruce 

and Cannon occurred with a male and 3 females of H. pelagicum in the stomach of 

a black marlin off Australia. 

Location in Host - Stomach, pyloric ceca and intestine. 


Host Specificity - It is host specific and is a primary parasite of dolphin. Other 

big game fishes that eat dolphin are false hosts. 

Damage to Host - The very heavy infections of more than 100 of these rather 

large worms could harm or at least stunt the growth of dolphin. They also contribute 

to the injury of this host when combined with superinfections of flukes. 

Hysterothylacium reliquens (Norris and Overstreet) 

The adult of this worm is an 

incidental, uncommon and possibly 

temporary parasite obtained by big game 

fishes through eating inshore fishes, but 

the larvae is occasionally found in 


Name - The name "reliquens" refers to 

abandonment or the vacating of the host 

by these worms which pour out of the 

mouth, gills and anus, after the host is 

landed. It was placed in genus Thynnascaris. 

Diagnostic Characters - The minute 

alae begin just posterior to the lips. The 

lips are longer than wide, constricted at


their midpart, and lack interlabial grooves. The tip of the tail is covered with 

numerous minute spines. Males have 23-29 pairs of papillae anterior to the 

anus, no pairs of papillae at the level of the anus, and 4-5 pairs posterior to the 

anus with the third pair from the end doubled (2 papillae jammed together in the 

row). It does not have striations (modified ventral annuli) at the area of the 

papillae. Female vulva opens in the anterior 16-45% of the body length. 

Records - We found heavy infections of larvae encysted in 2 cero from 

Humacao, but not in 33 other cero examined from various locations around 

Puerto Rico; and 7 in 1 of 4 king mackerel off Dauphin Island, Alabama. 

Larvae also encysted in Spanish mackerel from the northern Gulf of Mexico. 

One to 40 adult worms occurred in 14 of 303 swordfish from the northwest 

Atlantic (ARC 2411). 

Geographic Range - Western Atlantic and eastern Pacific. Our collections are 

the first in the Caribbean. 

Ecology - Fishes are infected in areas with salinities above 20 ppt but not above 

35 ppt or hypersaline. The area in eastern Puerto Rico with the heavy infection 

rates is contaminated (see Didymocystis scomberomori). 

Location in Host - Stomach and intestine. The third stage larvae are encysted 

in mesenteries and the outer layer of the stomach and intestine. 

Length - Female 21.0-130.0 mm, male 25.0-40.0 mm, female fourth-stage 

larvae (encysted in fish) 8.7-22.5 mm, male fourth-stage larvae 14.0-22.0 mm, 

early fourth-stage larvae 4.3-9.2 mm. 

Host Specificity - This adult occurs in a variety of inshore bottom fishes, but 

big game fishes may be false hosts. Encysted larvae occur in a variety of fishes 

including Spanish mackerels. Cero and king mackerel are new hosts. 

Damage to Host - Heavy infections of encysted larvae may injure cero. 

Iheringascaris inquies (Linton) 

Heavy infections of this worm occur 

around the world in cobia. 

Name - Deardorff and Overstreet (1980) 

resurrected and redescribed this genus. 

Hysterothylacium shyamasundarii (Lakshmi 

and Rao) and Neogoezia elacateiae 

Khan and Begum are synonyms of this 

worm. 

Diagnostic Characters - The body is 

relatively narrow. The lips are as long 

as wide to wider than long, and do not 

have deep interlabial grooves. The tail is 

conical. Males have spicules that are 

approximately equal in diameter and 

length (20-25% of body length). There are lateral rows of 6 papillae in addition 

to the normal pre- and post-anal papillae and a medioventral preanal organ is 

present. Males have 25-29 pairs of papillae anterior to the anus, and 6 median 

147

148 


pairs posterior to the anus with third pair from the end doubled (2 papillae 

jammed together in the row). It does not have striations (modified ventral 

annuli) at the area of the papillae. The female vulva opens in the anterior 31- 

38% of the body length. 

Records - Heavy infection of both sexes occurred in cobia from off La 

Parguera, Puerto Rico; and around the world. Overstreet in Shaffer and 

Nakamura (1989) noted that whenever a cobia is dissected, the stomach was 

found to be heavily infected with this roundworm. 


Location in Host - Stomach and pyloric ceca. 


Host Specificity - It only occurs in cobia, and is a characteristic parasite of 

this host. 

Damage to Host - Constant and heavy infections of this parasite must take a 

toll on the host. Overstreet (1978) reported that hundreds of worms occur in 

large cobia, and they occasionally cause ulcerated lesions at sites where several 

worms attach clustered together. 

Maricostula histiophori (Yamaguti) 

This is a cosmopolitan parasite of Atlantic and 

Indo-Pacific sailfish 

Name - It was redescribed by Olsen (1952) and 

Bruce and Cannon (1989). 

Diagnostic Characters - The alae begin at the lips. 

The lips are wider than long, indented 1/2 way up 

their sides, and have no interlabial grooves. There 

is a deep constriction at the base of the lips. The 

tail gradually tapers. Males have 14-17 pairs of 

papillae anterior to the anus, 1 doubled (2 papillae 

jammed together in the row) pair at the same level 

as the anus and 4 pairs posterior to the anus. There 

are crests (modified ventral annuli) in the area of the 

papillae. The female vulva opens in the anterior 

26.7-56.8% of the body length. 

Records - Two to 12 occurred in 4 of 6 Atlantic 

sailfish off Puerto Rico (USNM). A male and 1 female, and a female were 

found in 2 Atlantic sailfish off Panama; and 40 in an Atlantic sailfish off the Dry 

Tortugas, Florida, USA. It also occurs in Indo-Pacific sailfish. 

Geographic Range - Worldwide. Our collections are the first in the insular 

Caribbean. 


Length - Females 57.8-84.5, male 42.5-48.0, immature male 24.0-31.0 mm. 

Host Specificity - Host specific to sailfish or genus specific to Atlantic and 

Indo-Pacific sailfish; depending on which host taxonomy you follow. The


presence of this roundworm could be part of an argument for calling sailfish 1 

species. It may be a characteristic parasite of sailfishes. 

Damage to host - Iversen and Kelly (1974) found an intense granulomatous 

inflammatory reaction (or fibrous tissue in older infections) surrounding this 

worm in gastric ulcers. They suggested that the spines of prey fishes caused 

most ulcers but that nematodes may also be involved. 

Maricostula incurva (Rudolphi) 

It is a confusing and probably wide 

ranging and important parasite of 

swordfish. 

Name - The identity of this worm has 

been confused and its biology obscured 

because redescriptions have either been 

based on Maricostula histiophori or on a 

mixture of different species. 

Diagnostic Characters - The alae begin 

just posterior to the lips. The lips are 

wider than long, not indented in their 

sides, their anterior margin not narrowed 

and straight, and they have deep 

interlabial grooves. There is a deep 

constriction at the base of the lips. The tail gradually tapers, and the cervical 

alae are moderately wide. Males have 16 pairs of papillae anterior to the anus, 

1 doubled (2 papillae jammed together in the row) pair at the level of the anus, 

and 4 pairs posterior to the anus. Crests (modified ventral annuli) occur in the 

area of the papillae, have a high profile, and overlap each other. The spicules 

are about 160% of the length of the ejaculatory duct. The female vulva opens 

in the anterior 25-40% of the body length. 

Records - More than 100 occurred in each of 2 swordfish off La Parguera, 

Puerto Rico; off St. Croix, U.S. Virgin Islands; from Cuba; off Miami, Florida, 

USA, up to 3038 (average 96.4) in 285 of 303 swordfish from the northwest 

Atlantic (ARC 2312); in the Adriatic, Baltic, Mediterranean, North Sea, and in 

the Pacific. 


Life History - Hogans and Brattey (1982) suggested that larger worms, up to 

16 cm, live for several years. 

Associations - This worm occurred in combination with Hysterothylacium 

corrugatum in swordfish off Puerto Rico, Florida, in the northern Gulf of 

Mexico; and with Hysterothylacium aduncum, H. corrugatum and H. reliquens 

in the northwest Atlantic. 


Length - Female 25.0-160.0 mm, male 17.0-34.0 mm. Linton (1897) reported 

worms up to 267.0 mm. 

149

150 


Damage to Host - This large worm may stunt or injure swordfish particularly 

when it occurs in heavy to superinfections of thousands of worms. They may 

also contribute to severe injury to swordfish when they are combined with very 

heavy infections of Hysterothylacium corrugatum. 

Host Specificity - This worm is only found in swordfish and is probably a 

characteristic parasite. The few records of this roundworm from other fishes 

appear to have been other species. 

Maricostula sp. of Bruce and Cannon 

A characteristic parasite of white marlin that occurs less often in blue marlin 

throughout the western Atlantic. Superinfections damage billfishes and disgust 

fishermen. 

Name - Bruce and Cannon (1989) distinguished this worm a new species, but 

did not name it. We are examining our samples from Puerto Rico to determine 

if this worm warrants description as a new species. The Hysterothylacium 

incurvum (or Thynnascaris incurva) reported by various authors from billfishes 

appear to be this worm. 

Diagnostic Characters - The alae begin just posterior to the lips. The lips are 

wider than long, not indented in their sides, and have deep interlabial grooves. 

There is a deep constriction at the base of the lips. The tail gradually tapers, 

and the cervical alae are narrow. Males have 27-38 pairs of papillae anterior 

to the anus, 1 doubled (2 papillae jammed together in the row) pair at the level 

of the anus, and 4 pairs posterior to the anus. Crests (modified ventral annuli) 

occur in the area of the papillae, and are low profile and do not overlap each 

other. The spicules are about 60% of the length of the ejaculatory duct. The 

female vulva opens in the anterior 25-40% of the body length. 

Records - We found 11 in a white marlin off Mayaguez, Puerto Rico (USNPC), 

and 100 each in 2 white marlin from Dauphin Island Alabama, USA; and 1-7 

in 8 of 40 Atlantic blue marlin from various localities off Puerto Rico (USNPC). 

Up to 1787 occurred in each of 4 white marlin from the southern Gulf of 

Mexico; 300 worms in 22 white marlin off Destin, Florida to Orange Beach, 

Alabama, USA (USNPC 75843); and 6-741 in 36 white marlin from South 

Carolina, USA. 

Geographic Range - Western Atlantic. Our collections are the first in the 

Caribbean. 



Host Specificity - This worm only occurs in billfishes and may be family 

specific. It is a characteristic parasite of white marlin, and a secondary parasite 

of Atlantic blue marlin. Atlantic blue marlin is a new host for this parasite. 

Damage to Host - Superinfections of these worms must injure these hosts. 

Many of these worms quickly leave the host after it dies. Therefore, the 

numbers recorded in each fish may be a small percentage of the worms that 

existed in the live host and the damage greater.


Harm to Humans - These worms pouring out of infected marlin should be 

treated with caution. Splattering or wiping these parasites on your mouth or 

nose could cause a direct infection. 

Significance to Sport Fishing - Heavily infected marlin shed these large 

roundworms from their mouths and gills after they are landed. This spectacle 

disgusts otherwise seasoned fishermen. It may cause perfectly safe and 

wholesome seafood to be unnecessarily discarded. 

Oncophora melanocephala (Rudolphi) 

This parasite sporadically occurs in yellowfin tuna 

and Atlantic and Mediterranean big game fishes. 

Name - The name "melanocephala" means black head. 

A questionable species, Oncophora neglecta Diesing, 

was inadequately described from the gall bladder of a 

bluefin tuna in Europe. 

Diagnostic Characters - It has a large, obvious, 

cuticulized buccal capsule that is shaped like a scallop 

shell and is golden in color. The ribs on the shell 

extend about half way down the capsule or less. The 

spines occur on the posterior 1/2 of the capsule. The 

capsule tapers and becomes more narrow before it 

attaches to the esophagus. The tridents which occur on 

either side of the buccal capsule are longer than the capsule and the center tine 

is shorter than the outer ones. The mature female has an enlarged posterior end. 

Records - Three occurred in an albacore from off Desecheo Island, Puerto Rico 

(USNPC), and 1-2 were found in 3 of 40 Atlantic blue marlin from various 

localities off Puerto Rico (USNPC). One was found in 2, and 7 in 1 of 10 

yellowfin tuna from the southern Gulf of Mexico; 1-5 in 13 of 303 swordfish 

from the northwest Atlantic (ARC 2415); and in Atlantic bonito, Atlantic 

mackerel, bluefin tuna, bullet tuna and frigate tuna from the North Sea and 


Geographic Range - Atlantic and Mediterranean. Our collections are the first 



Length - Female 111.0-150.0, male 10.5-12.6, immature female 14.2-16.1 mm. 

Host Specificity - It may be family specific to scombrids. Atlantic blue marlin 

and swordfish are probably false hosts. Albacore and Atlantic blue marlin are 

new hosts. 

Ctenascarophis lesteri Crites, Overstreet and Maung 

This is a cosmopolitan and ever present parasite of the economically 

important skipjack tuna. 

Name - It is named for the collector of this worm. 

Diagnostic Characters - This parasite can be distinguished by the obvious 

spines on the body which are in broken combs instead of continuous rings. 

151

152 


The only other member of the genus, C. 

gastricus Mamaev, appropriately occurs on 

another medium-sized big game fish, frigate tuna. 

It has only been reported in the Pacific, but we 

should be looking for it in this host in the 

Atlantic. Ctenascarophis gastricus differs from C. 

lesteri because the combs extend down 2/3 of the 

body length in the male and to the anus in the 

female, and it has a maximum of 32 instead of 8 

spines per comb. 

Records - Moderate infections occurred in 11 

skipjack tuna from various localities around Puerto 

Rico (USNPC). Moderate to very heavy infec- 

tions were found in 878 skipjack tuna from 14 

sites across the Pacific. 


Associations - This worm is always found with 

Prospinitectus exiguus. 



Host Specificity - It only occurs in skipjack tuna and is a characteristic 

parasite. 

Prospinitectus exiguus Crites, Overstreet and Maung 

This worm is a companion of the previous 

parasite species, and the little we know about their 

biologies is identical. 

Name - The name "exiguus" means small or short 

and refers to the size of the body. 

Diagnostic Characters - This parasite can be 

distinguished by the obvious spines on the body 

which are in continuous rings instead of broken 

combs. 

The only other member of the genus, P. mollis 

(Mamaev), occurs on another medium-sized big game 

fish, frigate tuna. It has only been reported in the 

Pacific, but we should be looking for it in this host 

in the Atlantic. Prospinitectus mollis differs from P. 

exiguus because its spines rings are interrupted rings, 

is less than half as long, has a proportionally longer 

esophagus, and fewer spines per ring (28-49 instead 

of 70-100). 

Records - Moderate infections occurred in 11 skipjack 

tuna from various localities around Puerto Rico (USNPC). Moderate to very 

heavy infections were found in 878 skipjack tuna from 14 sites across the 

Pacific.



Associations - This worm is always found with Ctenascarophis lesteri. 

Location in Host - Intestine especially near the pyloric ceca, and rarely in the 

stomach. 


Host Specificity - This worm is only found in skipjack tuna. It always occurs 

in this fish and is thus a characteristic parasite of this fish. 

Significance to Sport Fishing - Skipjack tuna are economically important and 

abundant throughout much of the world. This worm and the previous parasite 

species occur abundantly in every skipjack tuna on Earth. Yet, these 2 parasite 

species were only recently recognized and described, and almost nothing is 

known about their biology. This situation is unfortunately indicative of the state 

of knowledge of big game fish parasites. 

Miscellaneous Roundworms 

Genus Ascaris Linnaeus - These roundworms occur as adults in 

terrestrial mammals and the larvae are not found in big game fishes. This genus 

name was used in the early literature for many larval forms found in these 

fishes, and poorly studied larval forms are still, although incorrectly, attributed 

to this genus. 

Genus Contracaecum Railliet and Henry - Adults occur in fish- 

eating birds and marine mammals. All adult forms in fishes reported in this 

genus probably belong to the genus Hysterothylacium. Encysted larval forms 

in big game fishes may possibly belong in Contracaecum, but these are difficult 

to identify to species. Larval forms of members of this genus encyst in fish and 

are similar in appearance to Hysterothylacium spp. larvae. They differ by 

having the excretory pore at the level of the nerve ring in Hysterothylacium spp. 

and near the lips in Contracaecum spp. Contracaecum clavatum Rudolphi is a 

synonym of Hysterothylacium aduncum. Contracaecum papilligerum (Creplin) 

a larval form in Atlantic mackerel and bullet tuna in the Mediterranean is 

recognized as Ascaris papilligera Creplin by Bruce, Adlard and Cannon (1994). 

Cucullanus carangis (MacCallum) - A 10.0 mm long, obviously 

immature, worm from the intestine of a crevalle jack at the New York Aquarium 

was described as Dacnitis carangis. Neither the description nor the illustration 

was sufficient to establish a new species. This name is best forgotten. 

Cucullanus pulcherrimus Barreto - This worm was described from a 

black jack in Brazil, but has not been seen again in 75 years. Members of this 

genus are characterized by having relatively large broad mouths surrounded by 

a muscular pharynx with a club-shaped swelling posteriorly and no appendix. 

The male has a sucker anterior to the anus (preanal), 2 equal spicules and a 

153

154 


shorter accessory piece (gubernaculum) and no caudal alae. The female has 2 

ovaries and a vulva opening near midbody. 

Cystoopsis scomber Zlatev - This worm occurred in the outer layer 

(conjunctiva) of the eye in Atlantic mackerel from the Black Sea, but has not 

been found in the Atlantic. This worm was described so poorly that it might be 

an encysted tissue fluke instead of a roundworm. 

A very famous roundworm expert once screamed across a room, containing 

the bulk of USA parasitologists at an august national meeting, that she 

appreciated the "roundworm" specimens the poor fellow had sent her for 

identification, but if he could not tell a tissue fluke from a roundworm, then he 

did not belong at this meeting! 

Ichthyostrongylus thunni Nikolaeva - This worm was named from 

yellowfin tuna in the southern Gulf of Mexico, but the status of this species and 

genus and even its Family or Order is uncertain. 

Oncophora albacarensis Baudin-Laurencin - This worm was named 

from yellowfin tuna caught in the eastern Atlantic. It may not be very common 

as it was not seen by numerous other studies in the Atlantic. We should be 

looking for it in the western Atlantic. 

Parascarophis galeata (Linton) - This worm is apparently a parasite of 

sharks that was accidentally introduced with prey into a pompano dolphin off 

North Carolina, USA. Cystidicola galeatus (Linton) and Filaria galeata Linton 

are synonyms. The head has a distinctive cap-like cuticular expansion which 

extends posteriorly further on one side (dorsal) than the other (ventral). The 

mouth has 2 long, thin front-to-back (dorsoventral) lips and 4 papillae. The 

mouth (buccal) cavity widens front-to-back (dorsoventrally) anteriorly and then 

becomes cylindrical. The esophagus is not distinctly divided into muscular and 

glandular portions. The tail is elongate, conical and ends in a blunt point. 

Numerous fragments were found in the stomach. Recorded from the Atlantic 


Philometra sp. - See Philometroides sp. below. 

Philometroides sp. - We received a large fresh ovary of an Atlantic blue 

marlin caught off Puerto Rico that was so filled with squirming minute 

roundworms that most of its biomass was parasites. Raju (1960) reported 

68,200 larval roundworms in the ovary of a Pacific skipjack tuna. All the eggs 

in the left ovary had been destroyed except the small transparent ones along the 

periphery. Fewer, but similar roundworms have been reported from the ovary 

of bigeye tuna and yellowfin tuna in the Pacific. Simmons (1969) found 

Philometra sp. and an unidentified spiruroid infecting the ovaries of


approximately 90% of the mature skipjack tuna in the eastern and western 

tropical Atlantic Ocean. Young fishes lacking mature ovaries were not infected. 

The highest number of worms he found in a pair of ovaries was 75, and he 

found no obvious damage to the eggs. He also noted that the same 2 worms 

were found in skipjack tuna off New York, USA, and examined by Maybelle 

Chitwood (a famous roundworm expert). Ichthyonema globiceps Rudolphi, 

probably Philometra sp., was reported in the ovary of a Spanish mackerel from 

New Jersey, USA. All the worms reported above may not be the same species, 

but they all appear to be tissue-dwelling roundworms that were either larval or 

too undeveloped to identify. Superinfections of these worms destroy the ovaries. 

They must have a severe affect on the reproduction of these big game fishes. 

The other, equally parasitized, ovary from our Atlantic blue marlin case was 

cooked and eaten by humans before they received our diagnosis. This 

apparently caused no obvious deleterious effects, as the dish received rave 

reviews for flavor, appearance and texture. This may be the only case known 

of an entire family basing an evening meal almost exclusively on fish-parasitic 

roundworms. These parasites are a mystery clearly in need of investigation. 

A roundworm reported in the dorsal aorta of yellowfin tuna in Japan, has 

never been identified. It caused the vessel to thicken and become tough, and 

gave it a yellowish tint. Tissue roundworms have also been reported in the red 

muscle of bigeye tuna in the Pacific. Obviously, many tissue roundworms occur 

in big game fishes and are in need of study. 

Porrocaecum paivai Silva-Motta and Gomes - This parasite was 

described from king mackerel from Brazil, but has been noted as poorly 

described or of uncertain status by subsequent authors. Adults in this genus 

parasitize birds, not fishes, thus the genus of this worm is also in question. The 

type material needs to be re-examined. 

Prospinitectus mollis (Mamaev) - This worm has been found in frigate 

tuna and 2 species of little tunnies in the Pacific. Thus far it has not been found 

in frigate tuna or little tunny in the Atlantic, but we should be looking for it. 

Spinitectus mollis Mamaev, Spinitectus palawanensis Schmidt and Kuntz, are 

synonyms. See discussion of Prospinitectus exiguus for a comparison. Females 

are 11.0-15.4 mm and males 8.2-13.1 mm long. 

Raphidascaris anchoviellae Chandler - This worm was described from 

striped anchovies, Anchova hepsetus (Linnaeus) [as Anchoviella epsetus] and 

reported in other fishes from Galveston Bay, Texas, USA (USNPC 39537-8). 

This species appears to have been described from immature males 4.0-5.8 and 

females 4.0-6.0 mm long and must remain questionable (species inquirenda) 

until it can be correlated with adult forms. Yamaguti (1959) listed "Sphyraena" 

as a host, but this appears to be an error. 

spiruroids - See Philometroides sp. above. 

155

156 

ACANTHOCEPHALA 

(SPINY-HEADED WORMS) 

These worms form a small phylum in the Animal Kingdom. The name 

"acanthocephala" means spiny headed. All spiny-headed worms are permanent 

parasites in the intestine of most vertebrates, including humans. Over 1000 

species are known. Adult females vary from 1 mm to longer than 1 m, but are 

usually about 2 cm long. Males of the same species are typically smaller than 

females. They may be white, yel- 

low, orange or red in color (Pom- 

phorhynchus lucyae Williams and 

Rogers seems to absorb orange 

pigments from crayfish in the intes- 

tine of southeastern USA coastal 

fishes). They are bilaterally symmetrical 

and unsegmented. They attach 

in the gut of their host with a 

globular or cylindrical, protrusible, 

spiny proboscis. The proboscis pops 

out like an everting plastic glove, 

and the spines fold out and lock like 

a compact umbrella. Muscles invert 

the proboscis, and a hydraulic system 

(lemnisci) pop it back out. Some 

species have spines on the body as 

well. Sexes are separate, fertiliza- 

tion is internal, and embryos develop 

in the body of the female. Shelled 

larvae (acanthors) are shed into the 

intestine of the host, pass out with the fecal material, are eaten by a crustacean, 

insect, mollusk or fish intermediate host, and develop into an acanthella then to 

an encysted cystacanth larval stage in the second intermediate host. When the 

final host consumes an infected intermediate host, the cystacanth develops into 

an adult in the intestine. Big game fishes are final hosts for a few spiny-headed 

worms and are sometimes infected by larval stages of marine mammal spinyheaded 

worms. Adults absorb nutrients from the gut contents of their hosts. 

Proboscis spines (or hooks) cause some mechanical damage, but this is only 

serious in a heavy infection. Treatments seldom are necessary. Fortunately, 

natural infections in big game fishes usually consist of only a few worms per 

host. Some inshore, northern fishes are routinely infected with hundreds to 

thousands of spiny-headed worms. These worms rarely harm humans since they 

are usually discarded with the intestine and other internal organs when fish are 

cleaned, but contamination is possible. Thorough cooking kills these parasites. 

Tuna and other big game fishes are prized ingredients of Japanese raw-fish

ACANTHOCEPHALA (SPINY-HEADED WORMS) 

dishes, but most of these products are frozen long enough to kill parasites before 

they are served "fresh" in Japan. 

A necropsy is necessary to find these worms in the intestine. Spiny-headed 

worms in big game fishes can be identified in wet mounts for routine 

examinations. For more detailed study, the proboscis must be fully everted 

before preserving. Worms must be refrigerated in distilled or fresh water for 

12-24 hours before preserving in 5% formalin. The thick cuticle of these worms 

does not allow alcohol solutions or stains to readily penetrate. The cuticle must 

be pierced before dehydrating in alcohol solutions and staining. 


Phylum Acanthocephala - spiny-headed worms Page 

Class Palaeacanthocephala 

Order Echinorhynchida 

Family Rhadinorhynchidae 

Gorgorhynchoides elongatus ....................................................... 157 

Gorgorhynchus xiphias ............................................................... 162 

Rhadinorhynchus pristis .............................................................. 158 

Serrasentis sagittifer* .................................................................. 160 

Order Polymorphida 

Family Polymorphidae 

Bolbosoma vasculosum* ............................................................. 161 

Miscellaneous Spiny-headed Worms .......................................... 162 

________ 

*Adult and larval forms in big game fishes. 

Gorgorhynchoides elongatus Cable and Linderoth 

This parasite is a rare "taxonomic ghost" with virtually nothing known of 

its biology or affect on big game fishes. 

Name - Linton found this worm during his extensive collections at Woods Hole, 

recognized it as a new species and new genus, deposited type specimens 

(USNPC 8044, 8045), but died before he could publish his results. Cable and 

Linderoth (1963) described the genus Gorgorhynchoides based on the material 

deposited by Linton. Three similar species, G. elongatus, G. bullocki Cable and 

Mafarachisi, and G. lintoni Cable and Mafarachisi are recognized in western 

Atlantic jacks. These forms are similar in shape, size, hosts and locality. Their 

descriptions were based on few specimens (G. elongatus = 1_, G. bullocki = 

1_, 1_, 4 immature _ and G. lintoni = 2_, 3_) including only 3 males which are 

taxonomically more important than females in distinguishing spiny-headed worm 

species. These parasites are known to be highly variable in their characteristics. 

Examination of additional specimens may establish these forms as valid species, 

but we prefer to treat them as variants of a single, poorly known, species. 

157

158 


Diagnostic Characters - The proboscis is short, broad and 

bulbously expanded on the anterior end, and has numerous hooks. 

Proboscis hooks are larger anteriorly and smaller posteriorly (24- 

28 longitudinal rows of 16-22 hooks each). The neck has 

numerous spines followed by a hump-like dorsal swelling of the 

anterior trunk. The forms G. bullocki, G. elongatus, and G. 

lintoni largely differ by having no spines on the trunk swelling, 

few spines, or many spines, respectively. The constancy and 

separations in this gradation need to be substantiated with many 

worms to be considered diagnostic. 

Records - Four females and 2 males occurred in 2 of 7 bar jacks 

from Mona Island (Puerto Rico) (USNPC); 1 female in a blue 

runner from Curaçao (USNPC 60345); 1 female, 1 male and 4 

immature females in crevalle jack from the Gulf Coast of Florida 

(USNPC 62970); and 3 females and 2 males in almaco jack and 

greater amberjack from Woods Hole, Massachusetts, USA 

(USNPC 36619, 38644 and 38645) (forms G. elongatus, 

Curaçao; G. bullocki, Florida; G. lintoni, Woods Hole). 

Geographic Range - Western Atlantic. Our collections are the 

first in the insular Caribbean. 


Length - Female 31.3-76.0 mm, male 40.0-68.0 mm (forms G. 

elongatus _ 31.3; G. bullocki _ 53.0, _ 40.0; G. lintoni _ 49.5- 

76.0, _ 66.0-68.0). 

Host Specificity - It is family specific to jacks, but apparently 

occurs too sporadically to be a characteristic parasite of any species. The 3 

forms of this worm have been attributed to different species of jacks, but those 

records are based on few collections. Bar jack is a new host. 

Cable and Mafarachisi (1970) redefined the genus and suggested that the 

species were "rigidly" host specific to jacks, and that parallel evolution had 

occurred among these fishes and parasites. Their suggestions are intriguing but 

seem a bit bold considering the low number of available collections with so few 

specimens. They further noted "widely separate localities" as the "isolating 

mechanism" for these parasites to speciate. Actually the localities are not very 

widely separated when the wide ranging abilities of these hosts is considered. 

Rhadinorhynchus pristis (Rudolphi) 

These large, orange worms are obvious in the gut of many big game fishes 


Name - Rhadinorhynchus selkirki Van Cleave is a synonym; and R. katsuwonis 

Harada, R. ornatus Van Cleave, and R. seriolae Yamaguti could be synonyms. 

Any of these species noted from the Atlantic were probably R. pristis. Some of 

these species were placed in the genus Nipporhynchus which is a synonym of 

Rhadinorhynchus (the Japanese must be relieved at the demise of this genus 

which apparently means "Japanese nose"). The "R. trachuri Harada" of Chen


and Yang, 1973 in Pacific skipjack tuna and yellowfin tuna, were 

probably R. pristis. 

Diagnostic Characters -The proboscis is elongate, slender and 

cylindrical. Proboscis hooks are large and uniform in size (14-16 

rows of 26 hooks each or 22-25 longitudinal rows of 36-40 hooks 

each) with a row of longer hooks at the base. A few hooks are 

present on one side of the anterior body. Females are large and 

orange in color, males smaller and cream-colored. 

Records - We found 1 female in an albacore off La Parguera; 2- 

3 females in 2 of 3, and 1 female in 1 of 8 Atlantic blue marlin 

off western Puerto Rico (USNPC 84705); 1-16 females and 0-1 

males in 5 of 14 (USNPC 84707-9) and 1-5 females in 3 of 6 

dolphin, 1 female in 1 of 5 king mackerel and 2 males in a little 

tunny off La Parguera; numerous females and males in 2 skipjack 

tunas from Arecibo and 2 from south of Patillas and 9-12 females 

and 8-9 males in 2 off Ponce (USNPC); and 1 female in a white 

marlin off western Puerto Rico (USNPC 84704). One male and 

2 females occurred in a dolphin off Curaçao (USNPC 60341); 

and 18 females and 2 males in 5 of 9 little tunny and 151 females 

and 85 males in a skipjack tuna off Bermuda. It has also been 

found in Atlantic mackerel, chub mackerel, and frigate tuna in the 

western Atlantic; sailfish in the Gulf of Mexico; 1-2 females in 

4 of 303 swordfish from the northwest Atlantic (ARC 2413); 

albacore from Europe, and chub mackerel and skipjack tuna from 

Japan. The Rhadinorhynchus sp. reported from bullet tuna in the 

Pacific and the Rhadinorhynchus tenuicornis of Rego, Carvalho- 

V., Mendonca and Alfonso-R., 1985 in 24 of 80 Atlantic 

mackerel from Portugal could also be this worm. 


Ecology - Occurs in many offshore, but also in some inshore, 

fishes. We only found it in big game fishes in the Caribbean. 

Location in Host - Stomach, intestine. Throughout gut of 

skipjack tuna. 


Host Specificity - The incidence and sex ratios in skipjack tuna 

suggests that this worm is a characteristic parasite. It may also 

be a characteristic parasite of Atlantic mackerel. It is a secon- 

dary parasite in dolphin, at least in the Caribbean. Incidences in 

most other big game fishes are less certain. The occasional 

occurrence of low numbers of this worm in Atlantic blue marlin, 

king mackerel, little tunny and swordfish suggest that these are false hosts. This 

parasite may prefer big game fishes, but occurs in many other fishes. Atlantic 

blue marlin and king mackerel are new hosts. 

Damage to Host - No damage was found in the fishes examined in this study, 

but the heavy infections in skipjack tuna may cause problems in this host. 

159

160 


Detection - The large, orange-colored females are quite obvious in the gut of 

fishes. Unlike many parasites, they do not disintegrate in frozen hosts. 

Serrasentis sagittifer (Linton) 

This parasite is found in cobia and immature forms are 

encysted in a variety of marine fishes including dolphin. 

Name - Serrasentis chauhani Datta, S. giganticus Bilqees, S. 

longa Tripathi, S. longiformis Bilqees, S. scomberomori Wang, 

and S. socialis (Leidy) are synonyms. 

Diagnostic Characters - This parasite has distinctive rows of 

spines (combs) on the ventral surface of its body in adult and 

encysted stages. The proboscis is short, bulbous and expanded 

on the anterior end, and covered with numerous, uniform spines 

(16-24 longitudinal rows of 14-18 hooks each). 

Records - Adults occur in cobia along the USA Atlantic and 

Gulf coasts, off Europe and West Africa. Immature stages 

occur in a variety of fish host including dolphin. An 

unidentified spiny-headed worm encysted on the stomach wall 

of 1 dolphin off North Carolina (Rose 1966) was probably this 

parasite. Adult worms have also been reported in permit, 

Trachinotus falcatus (Linnaeus), from Miami, Florida, USA. 


Location in Host - Intestine and pyloric ceca; body cavity, 

mesenteries and external surfaces of internal organs (encysted 

forms). 

Length - Female 6.0-130.0 mm, male 8.6-75.0 mm, juvenile 

female 4.0-6.4 mm, juvenile male 2.6-4.2 mm. 

Host Specificity - This worm may be a characteristic parasite of cobia. 

Damage to Host - Heavy infections of this rather large worm could injure the 

host. Encysted forms damage the tissues of their intermediate fish hosts and 

should produce significant injury if they occur in heavy infections. Severe 

intestinal damage was caused in cobia by adult Serrasentis nadakali George and 

Nadakal from India. Although this parasite did not deeply penetrate the 

intestinal tissues enough to even cause discontinuity of the mucosa, it caused 

considerable histopathological alterations including connective tissue hyperplasia, 

epithelial metaplasia, muscle hypertrophy, mucus epithelium necrosis and 

degeneration and extensive destruction of intestinal villi. Host responses 

included mobilization of lymphocytes and macrophages, and excessive 

production of mucus in the sight of infection. 

Detection - The large, slender females are as long as your finger and are thus 

quite obvious in the gut of fishes.


Bolbosoma vasculosum Rudolphi 

Immatures of this marine mammal parasite are occasionally 

found in big game fishes. 

Name - Bolbosoma thunni Harada is a synonym. 

Diagnostic Characters - The body has a distinct bulge near the 

proboscis covered with 1 broad band of small hooks. A second 

band occurs on the body between the bulge and the proboscis. 

The proboscis is short, club-shaped and has numerous large, 

uniform hooks (16-20, usually 18, longitudinal (spiral) rows of 8- 

9 hooks each). Lemnisci (in adults) are elongate. The 

form B. thunni was distinguished because it had short lemnisci (0.8 

mm long). 

Records - We found 1 adult worm in 1 of 6 king mackerel off La 

Parguera, Puerto Rico. Immature forms have been found in the 

body cavity of albacore and jacks in the eastern Atlantic and 

yellowfin tuna off west Africa, and in the intestine of Atlantic 

mackerel off Russia and bluefin tuna in Japan. Adults occur in 

common or saddleback dolphin, Delphinus delphis Linnaeus, and 

North Sea beaked whale, Mesoplodon bidens (Sowerby) in the 

Atlantic, Mediterranean and Pacific. 

Bolbosoma spp. were reported from bullet tuna, chub 

mackerel and yellowfin tuna in the Pacific. These forms could 

have been B. vasculosum. 

Geographic Range - Worldwide (at least in the northern 

hemisphere). More commonly reported from the northern 

Atlantic and North Pacific and Mediterranean. Our collections are the first in 

the Caribbean. 

Life History - Crustaceans are the first intermediate hosts, fishes the second 

intermediate hosts for the encysted cystacanth larvae, but also serve as 

intermediary hosts for immature and young worms. Fish-eating marine 

mammals are the final hosts. 

Location in Host - Encysted forms occur in various fish tissues, immature 

forms are found in the intestine of fishes, and adult worms occur in intestine of 

marine mammals. 

Length - Immature females in fishes 8.0-13.5 mm, adult females in mammals 

12.0-15.0 mm. 

Host Specificity - Big game fishes with immature worms are probably false 

hosts (see Gorgorhynchus xiphias). Adults only occur in common dolphin and 

North Sea beaked whale, while cystacanths and immatures seem rather 

unselective occurring in almost any fish. The worldwide distribution suggests 

that the choice for a first intermediate host crustacean can also be diverse. King 

mackerel is a new host. 

We found Bolbosoma capitatum (Linstow) and Bolbosoma sp. in a shortfin 

pilot whale, Globicephalus macrorhynchus Gray, from Puerto Rico. Our 

161

162 


collection is the first in the Caribbean and shortfin pilot whale is a new host. 

Miscellaneous Spiny-headed Worms 

Gorgorhynchus xiphias Hogans and Brattey - Swordfish is probably 

a false or accidental host for these worms as they occur in low numbers as 

immature females. This new species was described by Hogans and Brattey 

(1982) in an unpublished report. It appears to be similar to Bolbosoma 

vasculosum which also occurs as immature females in big game fishes. This 

worm has a rather small, slender trunk which is expanded on the anterior end. 

the trunk is armed with 2 distinct bands of circular spine rows anteriorly. The 

proboscis has 14-18 longitudinal rows of recurved hooks. Light infections, 

almost always (98%) 1 per host, occurred in 295 swordfish from 4 localities in 

the northwest Atlantic (1-4 in 20 of 73 from Cape Hatteras, North Carolina, 

USA; 1 each in 6 of 89 from the Georges Bank; 1-3 in 12 of 70 from the 

Scotian Shelf; and 1 each in 9 of 63 from the Grand Banks). It occurs in 

moderate-sized hosts (115-174 cm FL). All the information above is from 

Hogans and Brattey (1982) 

Tegorhynchus pectinarium Van Cleave - This worm was reported in the 

stomach of a "Medialuna" or Seriola sp. from the Pacific coast of Costa Rica. 

The host was probably a half moon, Medialuna californiensis (Steindachner), a 

rudderfish (Kyphosidae), not a big game fish. 

Telosentis tenuicornis (Linton) - Cable and Linderoth (1963) listed 

Atlantic mackerel as a host for this worm, as Rhadinorhynchus tenuicornis, but 

these parasites were probably R. pristis.

CRUSTACEA (CRUSTACEANS) 

163 

Crustaceans are one of the phyla of animals with hard, segmented shells 

(exoskeletons) something like Medieval knights in armor. They are largely 

aquatic, while the insects and spiders and their allies (arachnids) are mostly 

terrestrial. All these groups are sometimes lumped together as Phylum 

Arthropoda, but we prefer to consider Arthropoda a subkingdom. Crustaceans 

generally have 2 pairs of antennae; respire through gills or the body surface; and 

anteriorly or throughout have paired, segmented, usually biramous appendages. 

More than 40,000 living species and many fossil species have been described 

including copepods, ostracods, fish lice, barnacles and isopods. A series of 

books defines crustaceans (Bliss 1982-85). 


Phylum (or Subphylum) Crustacea - crustaceans Page 

Class Maxillopoda 

Subclass Ostracoda - ostracods or seed shrimp ..................................... 163 

Subclass Copepoda - copepods ............................................................. 165 

Subclass Branchiura - fish lice .............................................................. 223 

Subclass Cirripedia - barnacles ............................................................. 224 

Class Malacostraca 

Order Isopoda - isopods ....................................................................... 228 

OSTRACODA (SEED SHRIMP) 

Seed shrimp, ostracodes or ostracods are a rather large subclass of small 

crustaceans. The name "ostraco" means shell, an appropriate descriptor. They 

are macho, with the longest sperm known, and have one of the best and longest 

continuous fossil records in the animal Kingdom (since the early Cambrian 550 

million years ago). Ostracodes are of value chiefly because their abundance, 

ubiquity, and exacting ecological requirements make them excellent 

environmental indicators today and for the distant fossil past. These qualities 

have been most utilized to locate petroleum deposits, but they might be used also 

to answer some fundamental questions about global changes. More than 10,000 

living and 40,000 fossil species are known and 1/4 of these (about 1/2 of the 

living) have been described in the overwhelming literature of the last 30 years 

or so. The spurt of interest, when most other taxonomies were in decline, was 

caused by financial support, due to their use in petroleum exploration, and the 

arrival of a new tool to define their tiny structures, the scanning electron 

microscope. Also, the general availability of computers and more advanced 

computer programs aided their study since the fossil record of ostracodes has 

been described as exceeding the comprehension of any one human. Some effort 

has been made to combine the fossil (paleontological) and living (biological)

164 


classifications into a single system. A "taxonomic inflation" joke among 

ostracodologoists is that if you can tell 2 ostracodes apart they are different 

genera; if you cannot, they are different species! Living adults range from 0.1 

to over 33 mm, with almost all tending toward the lower end of this scale; fossil 

forms can be over 80 mm long (leperditicopids), but most are in the range 0.3- 

3.0 mm. Almost all ostracodes are characterized by having a body which is 

completely enclosed in a clam-like (bivalve) shell (specialized head shield) which 

is usually calcareous and is hinged dorsally. The body is either unsegmented or 

the segments are obscure. Most ostracodes swim using their antennules and 

antennae. Their trunk regions are greatly reduced and have none to 2 

appendages, but the tail projections (caudal rami) often are well developed. The 

larval stages are encased in bivalve shells; except punciids which have a single 

headshield. All ostracodes, except punciids, brood their eggs between the upper 

(dorsal) part of the body and the shell. In some groups 1-2 larval molts occur 

before the larvae are shed. Most species lay eggs freely or attach them to 

something. One nauplius stage is followed by 5-8 metanaupliar molts (periodic 

shedding of the exoskeleton to allow enlargement and growth). Most adults do 

not molt. Sexes are separate. Most ostracodes are free living. Some are 

commensal on sea stars and their allies (echinoderms), sponges and other 

crustaceans, but only a very few occur on the gills and nares of sharks and rays 

or rarely bony fishes. Males and females occur together on hosts. Most 

ostracodes are detritus or filter feeders, a few scavengers, others parasitic. They 

are not known to transmit diseases; but some freshwater species prey on the 

juveniles of schistosoma-transmitting snails. 


Subclass Ostracoda - seed shrimp Page 

Order Myodocopida 

Family Cypridinidae 

Vargula parasitica ...................................................................... 164 

Vargula parasitica Wilson 

This unusual parasite of sharks 

has also been found on blue runner. 

It is only known from Jamaica. 

Name - The name "parasitica" is 

derived from it being the first 

parasitic ostracod described in fishes, 

although normally free-living, Vargula 

mediterranea (Costa), had 

previously been found in a fish from 

near Elba in the Mediterranean. Both ostracodes were originally placed in the 

genus Cypridina. Harding (1966) redescribed V. parasitica. Another cypridinid

OSTRACODA (SEED SHRIMP) 

species, Sheina orri Harding, was found in the gills and nares of sharks and rays 

from Heron Island, Australia. 

Diagnostic Characters - It has an oval shell with a notch near anterior end and 

relatively large eyes. The first antennae has 4 segments and 3 long setae. 

Records - Three males occurred in a blue runner from Jamaica (USNM 43604). 

Also 50, 50 and 12 males and females were found in 3 smooth hammerhead 

(USNM 43581, 43586, 43603) and 1 in a rock hind, Epinephelus adscensionis 

(Osbeck), (USNM 43599). 


Life History - Most female ostracodes do not swim and reside in the bottom 

sand or mud, but fish-parasitic females occur on the hosts with the males. 

Males and females scraped off fish gills into sea water were able to swim. 

Location in Host - Most specimens were attached to the gill filaments, but 12 

males in 1 smooth hammerhead were in the nares (USNM 43603). Oddly the 

2 sharks with 50 ostracodes each in their gills had none in their nares. All those 

in the gills were attached between the bases of the filaments in contact and at 

right angles to the axis of the gill arch. Only 1 ostracod was found between 

each set of filaments. The exact positioning, attachment, tissue reaction of the 

host, and sheer numbers in a host indicate that these ostracodes were parasitic. 


Host Specificity - This parasite is probably subclass specific to sharks and 

rays. Those in blue runner and rock hind may have been accidental. 

Damage to Host - The gill tissue appeared to surround each ostracod. This 

could have caused by erosion into the filament by the ostracod, minor cell 

proliferation around the attachment site, or both. Fifty ostracodes could impair 

a considerable area of gill tissue. 

Detection - The raised areas on the gill filaments should make these small 

animal a bit easier to find. More carefully-documented collections would be of 

considerable use in understanding this unusual association. 

COPEPODA (COPEPODS) 

Copepods form a class in the crustaceans. The common name copepod 

means "oar-foot." They are found in marine and fresh water, most are free- 

living and are very important food items for a variety of aquatic life. 

Approximately 10,000 species have been described and about 2000 of these 

parasitize fishes. Many fish-parasitic species remain to be named. Copepods 

range in length from 0.5-25 cm, but most are less than 1 cm long. Egg strings 

of some species may exceed 60 cm. Body shapes vary tremendously from a 

generally cylindrical shape to a flattened or saucer shape. The body is 

theoretically divided into 16 segments or somites (head or cephalic=5, 

thoracic=7, and abdominal=4), but most of these are fused together, combined 

or overlapped so they cannot be seen. The first 6-9 somites are fused into a 

expanded "head", cephalothorax or cephalosome; and the remainder are 

variously fused or separated into thorax and abdomen units. Appendages are 

165

166 


modified into mouthparts and other structures in the cephalosome, legs and other 

appendages in the thorax, and the abdomen has no appendages, and usually 

terminates in a bifurcate tail with projections (caudal rami). We illustrate 

females, use their morphological characters, and seldom mention males, because 

females are usually larger, more available, and have more distinct characters. 

In most parasitic forms, the life cycle is direct, but typically involves a 

series of free swimming, planktonic stages. Some have intermediary hosts, such 

as Pennella spp. on squids, and shark copepods embedded in coral reef fishes. 

Finding a final host a serious problem for parasites, especially those that 

parasitize offshore, big game fishes. The hatching nauplii can often be observed 

by merely holding egg strings in bowls of sea water until they hatch. Nauplii 

of many species have never been described or drawn. Raising the other 

planktonic stages of parasitic copepods is more complicated, but can be 

accomplished. (A newly hatched nauplius of Caligus elongatus; an early 

nauplius, later nauplius, and metanauplius of C. bonito; and chalimus of C. 

elongatus attached to a host scale, are illustrated above.) Sexes are separate 

with males often much smaller than females. The life cycle stage that first 

attaches to fishes (copepodid, sometimes chalimus) can be very damaging to 

young fishes. Copepods frequently occur on the gills or skin of fishes, but 

highly specialized species burrow into the flesh or head sinuses, or crawl into 

the nose (nares or nasal fossae or lamellae) or eyes (orbits). They also associate 

with or parasitize a variety of invertebrates. We found one damaging a basket 

sea star in Puerto Rico (Williams and Wolfe-Walters 1990). Large fish-parasitic 

copepods are capable of biting humans, but such injury has seldom been 

reported. Most fishermen would not admit being attacked by a mere copepod! 

They can be of value to humans. Eskimos eat a giant parasitic copepod from


the gills of Atlantic cod, Gadus morhua Linnaeus. Those on fishes are usually 

permanent parasites, feeding on mucus, sloughed epithelial cells and tissue 

fluids. Copepods can directly transmit microbial diseases. 

Copepods are the most diverse and among the most abundant parasites of 

big game fishes. Collected specimens can be preserved and stored in 70% 

ethanol. They can be examined in alcohol or in a mixture of alcohol and 

glycerine to clear some structures. Smaller specimens or parts can be mounted 

in glycerine jelly for convenience in handling. 

Popular Reference - Kabata (1970) "Crustacea as enemies of fishes." 


Subclass Copepoda - copepods Page 

Order Poecilostomatoida ..................................................................... 169 

Family Bomolochidae 

Ceratacolax euthynni ................................................................. 170 

Holobomolochus asperatus ........................................................ 171 

Holobomolochus crevalleus ....................................................... 172 

Holobomolochus divaricatus ...................................................... 172 

Pseudoeucanthus uniseriatus ..................................................... 173 

Unicolax anonymous .................................................................. 174 

Unicolax collateralis .................................................................. 174 

Unicolax mycterobius ................................................................ 175 

Family Shiinoidae 

Shiinoa inauris ........................................................................... 176 

Family Philichthyidae 

Philichthys xiphiae .................................................................... 177 

Order Siphonostomatoida 

Family Caligidae 

Anuretes heckelii ....................................................................... 218 

Genus Caligus ................................................................................ 178 

Caligus balistae ....................................................................... 179 

Caligus bonito .......................................................................... 180 

Caligus chelifer ........................................................................ 219 

Caligus chorinemi .................................................................... 181 

Caligus coryphaenae ................................................................ 182 

Caligus curtus ........................................................................... 219 

Caligus elongatus ..................................................................... 183 

Caligus haemulonis .................................................................. 219 

Caligus isonyx .......................................................................... 184 

Caligus lobodes ........................................................................ 185 

Caligus longipedis .................................................................... 186 

167

168 


Caligus mutabilis ....................................................................... 187 

Caligus patulus .......................................................................... 220 

Caligus pelamydis ..................................................................... 188 

Caligus praetextus ..................................................................... 220 

Caligus productus ..................................................................... 189 

Caligus quadratus ..................................................................... 190 

Caligus robustus ....................................................................... 191 

Caligus spinosus ...................................................................... 191 

Caligus wilsoni ........................................................................ 192 

Lepeophtheirus bermudensis ................................................... 193 

Lepeophtheirus dissimulatus ................................................... 194 

Lepeophtheirus edwardsi ......................................................... 195 

Parapetalus occidentalis .......................................................... 196 

Tuxophorus caligodes .............................................................. 197 

Tuxophorus collettei ................................................................. 198 

Family Euryphoridae 

Genus Alebion ................................................................................ 217 

Alebion carchariae ................................................................... 217 

Alebion glaber .......................................................................... 218 

Alebion gracilis ........................................................................ 218 

Euryphorus brachypterus ......................................................... 198 

Euryphorus nordmanni ............................................................ 200 

Genus Gloiopotes ........................................................................... 201 

Gloiopotes americanus ............................................................. 201 

Gloiopotes hygomianus ............................................................ 202 

Gloiopotes ornatus ................................................................... 203 

Family Hatschekiidae 

Hatschekia amplicapa ............................................................. 203 

Family Pandaridae 

Pandarus sinuatus ................................................................... 222 

Family Cecropidae 

Cecrops latreillii ....................................................................... 220 

Family Pseudocycnidae 

Pseudocycnus appendiculatus .................................................. 204 

Pseudocycnoides buccata ......................................................... 205 

Family Lernanthropidae 

Lernanthropus giganteus .......................................................... 206 

Lernanthropus hiatus ................................................................ 221


Family Pennellidae 

Lernaeenicus longiventris ........................................................... 206 

Lernaeolophus hemiramphi ........................................................ 220 

Lernaeolophus striatus ............................................................... 208 

Lernaeolophus sultanus .............................................................. 221 

Genus Pennella ................................................................................ 209 

Pennella filosa ........................................................................... 211 

Pennella instructa ...................................................................... 212 

Pennella makaira ...................................................................... 213 

Pennella sp. ............................................................................... 214 

Family Lernaeopodidae 

Brachiella elegans ..................................................................... 218 

Brachiella thynni ........................................................................ 214 

Charopinopsis quaternia ........................................................... 215 

Clavellisa scombri ..................................................................... 216 

Thysanote longimana ................................................................. 222 

169 

Miscellaneous Copepods ........................................................... 217 

Order Poecilostomatoida 

Most of these copepods in big game fishes are modern discoveries. Their 

small size, ability to move rapidly on the body of the host and their attachment 

positions in the orbits and nares of fishes probably contributed to their being 

overlooked. Thus, we know very little about these parasites. Many have been 

described and never seen again, not from rarity but neglect. In this group, a 

prime opportunity exists for anyone to collect new and interesting information. 

Many of these parasites look like free-living copepods. The bomolochids 

are among the least modified, most generalized and smallest of the fish-parasitic 

copepods, yet others in this order are modified, Shiinoa inauris, or highly 

modified, Philichthys xiphiae, from the free-living form. Von Nordmann must 

not have been very impressed with these copepods as he named them 

"bomolochos" which means toady, beggar or buffoon. The Family Bomolo- 

chidae was established by Claus in 1875 [Not Sumpf in 1871 as noted in 

Yamaguti (1963)], but was generally ignored until the 1930's. Many of the 

genera, much less species, of bomolochids appear remarkably similar, and these 

similarities have resulted in great difficulties in the taxonomy and incorrect 

(synonymous) genera (Bomolochoides Vervoort, Parabomolochus Vervoort, 

Pseudobomolochus Wilson). None of these tiny bomolochids are easily 

distinguished. Body shapes and even proportions vary with swelling due to the 

state of maturity. The only reliable characters are the structure and appendages 

of the first antennae, mouthparts, and legs of the female. These minute parts 

must often be miraculously detached and mounted on microscope slides, and 

examined with a compound microscope. Attempting this process gives one a

170 


much greater admiration of "copepod-ologists" (carcinologists). In most groups 

of parasites, we have been able to suggest methods for separating species 

without resorting to intricate dissections. For bomolochids, we suggest some 

identification tricks, but reluctantly provide the more detailed and technical 

characters necessary for absolutely confirming species. If all else fails, send 

these tiny copepods to us. 

None of these lively creatures is known to harm their hosts to any noticeable 

extent, much less to hurt humans, or have any obvious effect on the sport 

fishing for their hosts. Just because these copepods are minuscule in size and 

occur in light infections does not necessarily mean they are unimportant. They 

occur in the highly sensitive nares and nasal passages of their hosts. These 

copepods can cause extreme irritation and can probably affect the health of the 

host. They may also be valuable as biological tags to distinguish stock, indicate 

migration, or distinguish host species. These eye and nose parasites are host 

specific. This is unusual among the copepod parasites of big game fishes, but 

is not surprising. Anything this minuscule in such a great ocean is either closely 

tied to the specific habits of its host or is lost. Bomolochids can only be 

detected with a careful host dissection. They appear only as tiny specks of 

discoloration in the mucus to the unaided eye. Just looking into, opening or 

scraping materials from the nares may not be sufficient. Tissues and mucus 

from the nasal areas and sinus or mucus canals of the head should be removed, 

placed in a small dish of sea water and examined with a dissection microscope. 

A few other fish-parasitic families and numerous genera and species, that 

do not reside in the sinuses of fishes, are found in this order, but none of them 

have infiltrated the open ocean world of big game fishes. This suggests that the 

sinuses of fishes is a pioneer niche that is being used by this order to invade a 

new habitat - the open ocean. Such an invasion should begin at the margin and 

spread out with the near-shore pelagic hosts having the most species of these 

copepods. Four out of the 10 species parasitize inshore pelagics, 4 parasitize 

both inshore and offshore pelagics, and 2 species only parasitize offshore 

pelagics. Interestingly, 4 of these species occur on little tunny. This inshore 

pelagic appears to be a real "stepping stone" to the offshore hosts. Two of these 

copepods occur on all 3 species of near shore Spanish mackerels, but not on the 

offshore king mackerel. This also indicates some degree of habitat 

differentiation between inshore and offshore pelagics. The one species that does 

occur on the king mackerel is not found on the other 3 species of hosts. These 

2 species of Holobomolochus on inshore and offshore Spanish mackerels are 

very similar and probably enjoyed a common ancestor. A perfect example of 

the inshore to offshore transfer. 

Ceratacolax euthynni Vervoort 

This nasal copepod occurs across the entire Atlantic in the Atlantic bonito, 

but in a more restricted distribution in the little tunny. 

Name - It is named for the genus of one of its hosts.


Diagnostic Characters - The thorax has 2 distinct 

saddle-shaped segments. Microscopically, the first 

antennae of the female has a long, heavily sclerotized 

hook between the first and second segments. 

Records - One to 10 females (average 3) and 0-5 males 

(average 2) occurred in little tunny from the Atlantic and 

Gulf coasts of the USA and Atlantic bonito throughout 

the Atlantic. Both nares were found infected in each of 

226 little tunny from west Africa. It has also been 

recorded from Venezuela, Brazil, Spain and South 

Africa. 


Ecology - This copepod parasitizes hosts in near-shore 

pelagic to offshore pelagic areas. 

Associations - Ceratacolax euthynni and Unicolax 

anonymous occur together in the nares of little tunny. 

Unicolax collateralis often occurs with U. mycterobius in little tuna in the same 

geographic regions known for C. euthynni, but neither has been found with C. 

euthynni. 

Location in Host - Nares (nasal lamellae). 

Length - Female 3.7-4.2 mm, male 1.3-1.6 mm 

Host Specificity - This parasite only occurs in the Atlantic bonito and little 

tunny. These hosts are both scombrids, but are not otherwise related. 

Significance to Sportfishing - If the distribution of this copepod on little 

tunny is limited to the northern Gulf of Mexico and Atlantic coasts of the USA, 

it may be of value as a biological tag. We expect this odd distribution is an 

artifact of limited collecting instead of a restricted host range, particularly since 

it has also been reported from west Africa. 

171 

Holobomolochus asperatus Cressey and Cressey 

Light infections occur on king mackerel 

throughout the southern half of the western North 

Atlantic. 

Name - The name "asperatus" refers to the patches of 

hairs on the abdomen and caudal rami. 

Diagnostic Characters - This copepod is most easily 

distinguished by its host, and location in the host. 

Microscopically, females of this genus have first 

antennae with unmodified (not hooked) setae. Homobomolochus 

asperatus can be distinguished from other 

members of this genus because of the patches of hairs 

on the caudal rami. 

Records - One to 2 females occurred in king mac- 

kerel off Cuba (USNM 172242-3). It has also been

172 


found in this host from Georgia, Florida, and Texas in the USA, and Trinidad 

and northern Brazil. 


Ecology - This copepod occurs on the offshore king mackerel, but not on 3 

inshore host species in the same genus and in the same regions. 


Length - Female 1.8 mm, male unknown. 

Host Specificity - This parasite only occurs on king mackerel. It is probably 

a characteristic, or at least, a primary parasite of this host. 

Holobomolochus crevalleus Cressey 

This small nasal and gill chamber copepod occurs in 

light infections on crevalle jacks. 

Name - "Crevalleus" refers to the common name of the 

host, crevalle jack. 

Diagnostic Characters - It is most easily distinguished 

by its host. Microscopically, female members of the 

genus have the setae of the first antennae unmodified 

(not hooked). Holobomolochus crevalleus differs from 

H. asperatus by having large patches of spinules on the 

endopod segments of the second and third legs; and from 

H. divaricatus by having ornamentation on the ventral 

surface of the caudal rami. Egg strings are not shown 

in the accompanying figure because they have not been 

observed or described. 

Records - One to 2 females in each of 4 crevalle jacks, 

and 14 other females from an undetermined number of 

this host, from the west central Gulf coast of Florida. 


Location in Host - Gill chambers and nares (nasal lamellae). 

Length - Female 1.5 mm; male unknown. 

Host Specificity - This copepod is only known from the crevalle jack. 

Holobomolochus divaricatus Cressey and Cressey 

Light infections of this small copepod occur in the noses of all 3 inshore 

western Atlantic mackerels, but not on the offshore king mackerel. 

Name - Named for the "spread-legged" appearance of the caudal rami. 

Diagnostic Characters - It is most easily distinguished by its hosts, and 

location in the host. Only Shiinoa inauris shares the nares of these mackerels, 

and its body is cylindrical. Microscopically, female members of the genus have 

the setae of the first antennae unmodified (not hooked). Holobomolochus 

divaricatus can be distinguished from other members of this genus because the 

distal-most spine of the exopod on the third leg is shorter than the preceding 2. 

Records - We found 3-4 each in 35 cero from various localities around Puerto 

Rico; and in 10 Spanish mackerel from Dauphin Island, Alabama, USA. It is


apparently found in cero, Spanish mackerel and serra 

Spanish mackerel throughout the western Atlantic. 


Ecology - This parasite occurs on 3 species of inshore 

Spanish mackerels, but not on the offshore king 

mackerel in the same regions. Only a few copepods 

occur on each host. Females are usually more 

numerous (approximately 3:1), but are sometimes 

equal in numbers with the males. 

Associations - Another copepod, Shiinoa inauris, 

occurs in the nares of these Spanish mackerels, but 

these 2 parasites have not been reported from the same 

host specimen. 


Length - Female 2.4-2.8 mm; male 1.1-1.3 mm. 

Host Specificity - This copepod is a characteristic 

parasite of these 3 species of Spanish mackerels. It is also genus specific 

(Scomberomorus). 

173 

Pseudoeucanthus uniseriatus Wilson 

This is a small and little known parasite of the blue 

runner in Jamaica. 

Name - "Uniseriate" refers to the eggs that occur in single 

rows in the egg strings (although some multiserate do 

occur, see figure). The genus was named 

"Pseudoeucanthus" because it was similar to the genus 

Eucanthus, but this name had already been used for a genus 

of beetles. Thus the copepod genus Eucanthus was 

changed to Anchistrotos (but Pseudoeucanthus was not 

changed), and now we are left with a copepod genus name 

that means "false some kind of beetle!" 

Diagnostic Characters - The cephalosome is round. 

Body segments 2-6 are more narrow than the cephalosome 

and are all of equal width. The pair fifth legs are 

rudimentary and sixth legs merely small setae. The 

abdomen is divided into 3 segments and is about 1/3 the 

width of the genital complex. The caudal rami are longer 

than the last abdominal segment. Eggs occur in a single 

row (uniseriate) in the egg strings except for short segments 

with 2 rows. The egg strings are longer than the body. 

The body is dark gray. 

Records - Three females with egg strings occurred in a 

blue runner from Jamaica (USNM 43510, 42256). 


Life History - Eggs strings contain 30-35 eggs each.

174 


Location in Host - Mouth. 

Length - Female 1.2-1.3 mm, male unknown. 

Host Specificity - Only known from the blue runner. 

Unicolax anonymous (Vervoort) 

This rather small, enigmatic nasal copepod 

has only been collected in little tunny from 2 

localities on either side of the Atlantic. 

Name - This copepod was originally described in 

the genus Parabomolochus. 

Diagnostic Characters - These are minuscule 

nasal copepods. Females have 3 pairs of exposed 

legs in the mid-section of the body with an oblong 

shield over each pair. Microscopically, this 

copepod differs from U. mycterobius by having 

exopod spines of legs 2-4 serrated instead of edged 

with fine hairs. 

Records - Fifteen females and a male were found 

in a little tunny off Alabama, USA. Previously, 

it was reported from an undisclosed locality in the 

Gulf of Mexico and off Ghana, West Africa. This 

small copepod can easily be overlooked and 

probably occurs over a broader geographic range 

than the collections indicate. 


Life History - Males are rarely found. 

Associations - See Ceratacolax euthynni. 


Length - Female 0.9-1.0 mm, male 0.8 mm. 

Host Specificity - This parasite only occurs on little 

tunny. It is probably a characteristic parasite, or at 

least, a primary parasite of this host. 

Unicolax collateralis Cressey and Cressey 

This nasal parasite occurs in a variety of small tunas, 

bonitos and mackerels in both inshore and offs- 

hore pelagic habitats. 

Name - "Collateralis" means standing side-by-side, and 

refers to its occurrence with U. mycterobius. 

Diagnostic Characters - The thorax has 2 distinct 

saddle-shaped segments. Microscopically, the first 

antennae of the female has a long, heavily sclerotized 

hook between the first and second segments. 

Records - One to 12 females (average 2) and 0-5 

males (average 2) occurred in little tunny from the


Caribbean between Colombia and Panama; and Brazil. It was also found in 

bullet tuna and frigate tuna from Massachusetts, USA, and in these hosts in the 

Mediterranean. This copepod occurs on other species of bonitos and little tunas 



Ecology - It parasitizes hosts in near-shore pelagic and in offshore pelagic 

habitats. 

Associations - This copepod often occurs with Unicolax mycterobius in little 

tunny, bullet tuna and frigate tuna, but their interactions have not been studied. 

Ceratacolax euthynni and U. anonymous occur in the nares of little tunny in the 

same geographic regions, but have not been found with U. collateralis. 


Length - Female 1.5 mm, male 1.1 mm. 

Host Specificity - This parasite occurs in little tunas of the genera Euthynnus 

(all 3 species), Auxis (both species), and bonitos. This pattern of hosts suggest 

that little tunas (Katsuwonini) and bonitos (Sardini) might be more closely 

related than other scombrids. Why this copepod parasitizes bonitos in the 

Pacific, but not Atlantic bonito, might be a question worthy of further 

investigation. 

Unicolax mycterobius (Vervoort) 

This copepod occurs in little tunas worldwide. 

Name - This copepod was originally described in the 

genus Parabomolochus. 

Diagnostic Characters - These minuscule nasal 

copepods have 3 pairs of exposed legs in the mid- 

section of body with an oblong shield over each pair. 

Microscopically, it differs from U. anonymous by 

having exopod spines of legs 2-4 edged with fine hairs 

instead of being serrated. 

Records - One female and 2 males occurred in a 

frigate tuna from Massachusetts, USA; and 1 female in 

a little tunny from Pensacola, Florida, USA. Off west 

Africa, 63 of 72 frigate tuna were infected, usually in 

both nostrils. The examiner suggested that all host 

specimens were probably infected, but the smaller 

copepods were overlooked. It was also found in little 

tunas from the Mediterranean, Hawaii, Japan and the 

Philippines. 


Ecology - Males are found on the host as often as females. 

Associations - Ceratacolax euthynni and U. anonymous also occur in the nares 

of little tunny, but they have not been found in the same host specimen with N. 

mycterobius. 


175

176 



Host Specificity - This parasite is tribe specific to little tunas (Katsuwonini). 

It is probably a characteristic parasite of frigate tuna. 

Shiinoa inauris Cressey 

This tiny, cylindrical copepod occurs 

in very light infections in the nares of all 

3 inshore Spanish mackerels, but not 

offshore king mackerel. The piggy-back 

attachment of the male is unusual among 

copepods. 

Diagnostic Characters - It has a 

cylindrical body and a long curved 

"nose" (rostrum). The male is often 

attached on the dorsal surface posterior of 

the rostrum of the female (as illustrated). 

The rostrum and second antennae form a 

loop which anchors the copepod through 

a hole in the nasal lamella. This highly 

modified copepod is distinct from any 

other copepods found in the nares of 

western Atlantic big game fishes. A 

broken egg string is figured because an 

intact one has never been found. 

Records - One to 3 (usually 1) females, often with attached males, were found 

from cero in Puerto Rico, Cuba, Surinam and Venezuela; serra Spanish 

mackerel from Brazil; and Spanish mackerel from Florida, Texas and 

Massachusetts, USA. Cressey (1975) reported this copepod from Argentina, but 

later limited its southern range to Brazil (Cressey and Cressey 1980). 


Ecology - This copepod is found on inshore but not offshore Spanish mackerels. 

Associations - Another copepod, Holobomolochus divaricatus, occurs in the 

nares of these mackerels, but these 2 parasites have not been reported from the 

same host specimen. 


Length - Female 3.4-3.7 mm; attached male 1.9 mm. 

Host Specificity - The 3 inshore species of Spanish mackerels were infected 

but not the offshore king mackerel. 

Detection - The nares must be carefully opened to search for these small 

copepods. Egg sacs may be more easy to see than the body, they are usually 

broken during the examination and may not remain attached to the copepod.


Philichthys xiphiae Steenstrup 

This most unusually shaped copepod is hidden in 

the head sinuses canals of swordfish around the world. 

Name - The genus name means a friend of fish 

("phil"=friend, "ichthys"=fish), a slight misnomer. 

The specific name refers to swordfish. Yamaguti 

(1963) placed this copepod in its own order 

(Philichthyidea), and this copepod and bomolochids 

have traditionally been placed in separate orders at 

extremes of the copepod phylogenetic scheme. 

Modern classification unites these seemingly different 

forms through similarities in less modified structures. 

This scheme does unite all the nasal copepods of big 

game fishes into a single order which seems 

appropriate. 

Diagnostic Characters - This copepod is considerably 

larger than any other sinus copepod and the only 

one noted from swordfish. The highly modified body 

of the female resembles a twisted mass of PVC plastic 

tubing. The swimming legs, obvious in most other copepods, are absent. The 

body is white and eggs olivaceous. 

Records - Reported in swordfish from the New England coast of the USA. 

There are also a few other widely dispersed records but none have been reported 

from the tropical and subtropical regions. It has also been reported from striped 

marlin, Tetrapterus audax (Philippi), from the eastern Pacific. 


Life History - Younger females are smaller and also have less elaborate and 

convoluted projections. 

Ecology - It is only found in the offshore and pelagic habitat. 

Location in Host - This parasite occurs in the mucus canals in the head of 

swordfish and, oddly, embedded in the opercle bone of striped marlin. 


Host Specificity - This parasite has only been found in swordfish and striped 

marlin. It may be a characteristic parasite of swordfish. The location of this 

copepod in striped marlin was unusual, and may have been an accidental host. 

Damage to Host - The possibility of heavy infections was suggested but not 

confirmed by Wilson (1932). High numbers of this large copepod in the mucus 

canal would probably injure swordfish. It produces swelling in the host tissues. 

Detection - Unlike most tiny sinus copepods, this large copepod is easy to find. 

Opening and examining the mucus canals requires more dissection than just 

looking in the nares. 

177

178 


Genus Caligus Müller 

A Caligus was the second species of fish-parasitic copepods ever mentioned 

in the scientific literature and the genus was established in 1785. Chronologically 

they were number 2, but in big game fishes, they are number 1 - the most 

common and diverse genus of parasites. 

Almost everyone who has ever seen a freshly caught big game fish is 

familiar with these copepods. This genus is the stereotypical fish copepod that 

everyone envisions (what the Shell Oil sign is to mollusks, this genus is to 

copepods). The characters that are so distinctive for this genus include: having 

lunules (not the big headlights that they appear, but attachment devices) on the 

front of the cephalosome; H-shaped grooves on top of the cephalosome; the 

thorax usually cylindrical; the male usually smaller than the female; and egg 

strings that are usually longer than the body. They attach to the host with 

prehensile appendages and are capable of sometimes surprisingly rapid 

movement over the surface of a host. This large (200+ species) and important 

genus of skin-scurrying copepods has been the basis for various ill-fated orders, 

and a family of copepods that still survives. It has more species (at least 20) on 

western Atlantic big game fishes than any other genus of parasites. 

There are a lot of these creatures on offshore big game fishes, sometimes 

multiple species on a single host (although this is seldom reported), and some 

are a little tricky to tell apart. These parasites are some of the larger copepods 

on fish, but you may need a good dissection and possibly a compound 

microscope to tell all of them apart. Much of the confusion in identifying these 

copepods has come from the use of variable or undependable characters such as 

hosts, size, body width, etc. We try to use dependable but more easily seen 

characters, such as the separation of lunules on the front of the cephalosome. 

We define 3 categories of separation: (1) widely = separated by 3 or more 

diameters of the lunules, (2) moderately = 2 diameters more or less, (3) 

narrowly = 1 diameter or less. All of our characters sound good on paper, but 

may fall apart on a pitching deck, when all you have is a hand full of copepods 

and a hand lens. Let us know which characters fail field tests. Some species 

are so similar that they can only be separated by using microscopic structures. 

We have tried to avoid using these "third hair on the fifth toe" characters, but 

sometimes have to resort to the technical and tiny. 

These relatively large copepods are not known to bite or otherwise harm 

humans. They move around quickly, thus a host must be examined immediately 

after capture to determine the correct positions and numbers of these parasites. 

These copepods may leave or be knocked or washed off the host and attach to 

anything available, even your fingers. Many erroneous host records have been 

established by mixing sport fishes on the decks of vessels. 

Hundreds of these copepods can occur on a single fish. They often 

congregate in small areas that provide protection or easy feeding and produce 

open sores or lesions. The feeding of some species penetrates into muscle tissue 

of the host and leaves obvious scars. Wilson (1932) perceived caligoids as


threats to food fishes because of their ability to move about on the host and 

swim to and congregate on weakened hosts. Most copepods are permanent 

parasites that do not change hosts in the wild, but some in this genus apparently 

swim between hosts. His vision adequately describes some of the copepod 

problems in modern cage culture of marine fishes, particularly salmon, 

yellowtail, amberjack and possibly dolphin. 

All of these copepods on big game fishes are sufficiently large to cause 

damage, but seldom occur in numbers higher than a few on a host. Most do not 

attach in one place and erode a site, but move around on the host. We know of 

no cases of obvious damage to big game fishes, although all parasites cause 

minor damage. They could be of more significance to sport fishes by their use 

as biological tags. Unfortunately, the early literature is so confused, and 

examinations by modern copepod people are so limited, that sufficient 

information is lacking to properly understand these "biological tags." We 

expect that few are strictly host, genus, family or region specific, but some may 

be close enough to use as biological tags. 

Margolis, Kabata and Parker (1975) provided a synopsis of the world literature 

on this genus and Cressey (1991) surveyed those in the Gulf of Mexico. 

Caligus balistae Steenstrup and Lütken 

This copepod is rarely transmitted to dolphins as 

a false host from sargassum triggerfish that they eat. 

Beyond this curious example of prey-to-predator 

transfer, it has no significance to big game fishes. 

Name - The species name would more logically 

represent the family of the principle hosts 

(Balistidae), but it was actually named for the genus 

of the unknown original host (Balistes sp.). Caligus 

alatus Heegaard, C. canthidermis Yamaguti and 

Yamasu, C. polycanthi Gnanamuthu, and C. sensilis 

Kabata and Guzev are synonyms. 

Diagnostic Characters - It has a series of wrinkles 

in the dorsal cephalosome between the lunules. The 

cephalosome is about 1/2 of the body length, the 

lunules are moderately separated. The genital 

complex is rectangular, about 1/2 to 2/3 as wide as 

the cephalosome. The 1-segmented abdomen is short, less than 1/2 the length 

of the genital complex. The orange egg strings are longer than the body. 

Records - One female occurred on 1 of 20 dolphin off La Parguera, Puerto 

Rico. Cressey (1991) also found a female on a dolphin from either the southern 

USA or the northeast coast of South America. It was also reported once from 

bluefin tuna (as C. calistae, a misspelling). 

Geographic Range - Worldwide. Puerto Rico is a new locality and Williams, 

Bunkley-Williams and Rand (1994) noted this copepod for the first time off 

Bermuda. 

179

180 


Life History - A few chalimus stages were found attached by frontal filaments 

to the bodies of West Indian filefishes (Wilson 1905). 

Ecology: - This copepod occurs on hosts in both inshore benthic and offshore 

pelagic habitats. 

Associations - This copepod is very likely transmitted to the dolphin from prey 

items. Sargassum triggerfish, Xanthichthys rigens (Linnaeus), are an important 

food item for dolphin. We find them in almost every dolphin stomach we 

examine. 

Location in Host - Fins and body. 

Length - Female 4.0-4.5 mm, egg strings 4.6 mm; male 3.5-4.0 mm. 

Host Specificity - This parasite is family specific to triggerfish and filefishes 

(Balistidae). It is a characteristic parasite of some triggerfishes, but dolphin is 

probably a false host. 

Significance to Sport Fishes - This rare parasite has little effect on dolphin. 

Other big game fishes that consume sargassum triggerfish may also become 

infected. This transfer of a normally family-specific copepod could be a 

pathway for eventually forming new species on the predator species. Big game 

fishes may have more open niches for parasites than coastal hosts. 

Caligus bonito Wilson 

This ubiquitous copepod is a common parasite of bonitos 

and tunas around the world, but also occurs on a great variety 

of other fishes. Heavy infections probably injure bonitos. 

Name - It is aptly named for bonitos, its most common host, 

and was redescribed by Lewis (1967) and Pillai (1971). 

Caligus auxisi Pillai, C. kuroshio Shiino and C. sarda Pearse 

are synonyms. 

Diagnostic Characters - It has an oval cephalosome and 

narrowly separated lunules. The abdomen is elongate and 1segmented. 

It is similar to C. productus but can be 

distinguished for having 7 instead of 4 setae of the last segment 

(exopod) of leg 1. 

Records - One to 6 females occurred on horse-eye jack and 

coral reef fishes from La Parguera, Puerto Rico. It was also 

found on Atlantic bonito, bluefin tuna, cero, dolphin, little 

tunny and skipjack tuna in the Atlantic, and a variety of 

bonitos and tunas in the Indo-Pacific. It was reported on king 

mackerel and Spanish mackerel in the Atlantic, but these 

records need to be re-confirmed. Wilson (1905) found as 

many as 100 of these copepods on a single Atlantic bonito. Silas and 

Ummerkutty (1962) found 2-10 copepods on all 50 adult to juvenile striped 

bonito, Sarda orientalis (Temminck and Schlegel), they examined off India. 

Geographic Range - Worldwide (except polar oceans). 

Life History - Silas and Ummerkutty (1962) found chalimus stages and 

immatures on bonitos, but did not describe them.


Ecology - This copepod is more common offshore, but occurs in a variety of 

inshore habitats. 

Associations - Males and females of this parasite are in turn hyperparasitized 

by the copepod worm, Udonella caligorum, and covered with the attached eggs 

of this worm. We have only seen this worm and copepod combination on a 

large inshore fish, dog snapper, Lutjanus jocu (Schneider), but the association 

may occur on offshore fishes as well. 

Location in Host - Most attach to the roof of the mouth and the bases of the 

branchial arches. A few occur on the gill filaments and inside the operculum. 

Length - Females 4.9-8.5 mm, egg strings 8.0 mm, males 4.0-6.5 mm. The 

smallest occurred in the warmest waters and the largest in the coldest waters. 

Host Specificity - Both the number of hosts infected and the number per host 

indicate that bonitos are the preferred hosts of this parasite. It occurs 

commonly, but less often on tunas, rarely on Spanish mackerels and jacks, and 

also occurs on a wide variety of hosts. It may be a characteristic parasite of 

Atlantic bonito. We found it in 5 new hosts including horse-eye jack. 

Damage to Hosts - One hundred of these large copepods, as noted by Wilson 

(1905), could stunt or kill hosts as large as Atlantic bonito. 

Caligus chorinemi Krøyer 

This is a much misunderstood parasite of 

jacks in the western Atlantic. 

Name - This copepod was named for the genus of 

the original host, Chorinemus saliens (=leatherjacket), 

in 1863. Caligus germoi Pearse, C. rectus 

Pearse and C. tenax Heller are synonyms. 

Diagnostic characters - It has widely spaced 

lunules. The 1-segmented abdomen is about as 

long as the genital complex. The caudal rami are 

small (less than 1/10 the length of the abdomen). 

It is similar to C. isonyx and Caligus spp. (some 

called C. tenax) from Indo-Pacific jacks. It differs 

from C. isonyx by hosts and, microscopically, by 

having the terminal-most seta of leg 4 in the 

female twice as long as the other 2 terminal seta, 

instead of being similar in length; and the 

spiniform process of the first maxilla bearing an 

accessory process. 

Records - One female occurred in an albacore from Bimini, Bahamas (USNM 

88569); and in blue runner and yellow jack from Belize (USNM). Eleven 

females were found in 7 crevalle jack from Port Aransas, Texas (USNM 

92665); and in crevalle jack from Florida, USA, and Brazil. 

Geographic Range - Western Atlantic. Margolis, Kabata and Parker (1975) 

suggested that it occurs in the New World tropics in the western Atlantic and 

eastern Pacific. Despite a confused history under many names with various 

181

182 


Indo-Pacific records, Cressey (1991) found this copepod only in the Western 

Atlantic. 

Life History - The egg strings contain 105 eggs. 

Location in Host - Gills. 

Length - Female 2.6-5.1 mm, egg strings about 3/4 body length 3.4 mm; male 

2.9 mm. 

Host Specificity - This parasite is genus specific to jacks (Caranx spp.). The 

single record from an albacore may have been a false or accidental host or just 

contamination. 

Caligus coryphaenae Steenstrup and Lütken 

This is a ubiquitous parasite of tunas, little tunas 

and dolphins everywhere but polar seas. It is probably 

important but is virtually unstudied. 

Name - Despite being named for dolphins, this 

copepod is as often found on tunas. Caligus aliuncus 

Wilson, C. elongatus Heegaard, C. tesserifer Shiino 

and C. thymni Dana (sometimes spelled C. thynni) are 

synonyms. 

A record of this copepod on cobia by Causey 

(1953b) was actually Tuxophorus caligodes. A similar 

record on crevalle jack in Texas, USA, by Causey 

(1953b) could have been C. bonito or T. caligodes as 

this specimen was not available for examination, and 

many of Causey's copepod identifications were 

incorrect (Cressey and Nutter 1987). 

Diagnostic characters - All species in this genus 

have an H-shaped series of grooves on top of cephalosome, 

but it is easily distinguished by an additional 

groove across the top of the H. It has moderately 

separated lunules. The cephalosome is about 1/2 the 

body length; the genital complex and abdomen are 

about 1/4 each. The 3-segmented abdomen (sometimes looks like 2 segments, 

and has been called 5). Microscopically, it differs from other members of the 

genus, except Caligus regalis Leigh-Sharpe, by having a spine below the 

antennae, and a sclerotized process near the base of the sternal furca. These 2 

species may belong in a new genus (Cressey 1991). 

Records - Three to 20 males and females occurred on blackfin tuna, dolphin, 

little tunny, pompano dolphin and skipjack tuna from various localities off 

Puerto Rico. Females were found in little tunny off the Dry Tortugas, Florida, 

USA (USNM 64044); 25 females on 1 and 1-24 on 22 of 30 bluefin tuna off 

Prince Edward Island, Canada (ARC 2488-83); and 2 females in a swordfish 

from off Massachusetts, USA (USNM 54103). It has also been noted on 

albacore, bigeye tuna, bluefin tuna, skipjack tuna and yellowfin tuna from the 

western Atlantic; and bullet tuna, frigate tuna and wahoo from the Pacific.


Geographic Range - Worldwide (except in polar oceans). 

Life History - Male copepods occur on the host almost as commonly as 

females. Egg strings contain about 40 eggs each. 

Location in Host - Body surface and sometimes gills. 

Length - Females 4.5-8.0 mm; male 3.5-5.6 mm. The egg strings are longer 

than the body. Atlantic specimens on scombrids tend to be a bit larger (5.8-6.5 

mm) than those from the Indo-Pacific (4.5-6.0 mm)(Cressey and Cressey 1980). 

Host Specificity - Little tunas (Katsuwonini), dolphin (Coryphaenidae) and 

tunas (Thunnini) are the preferred hosts of this parasite. Its preference for this 

family and tribes and avoidance of similar and numerous species in other 

scombrid tribes is difficult to explain. The record from a swordfish could have 

been an accidental or false host. 

Caligus elongatus Nordmann 

This generalist has the least host specificity of all the fishparasitic 

copepods, attacking the world's fishes at random. It 

may be more important in temperate and colder waters. 

Name - This name has been used 3 times (Nordmann, 1832; 

Edwards, 1840; Heegaard, 1943) for 3 different species of 

copepods. Nordmann failed to note the host and locality for 

this parasite and did not publish the plate figuring this 

copepod, but the species is inexplicably credited to him. It 

has been confused with C. rapax Edwards since 1850 and 

recorded numerous times around the world under that name. 

This copepod was redescribed by Parker (1969). 

Diagnostic Characters - It has widely separated lunules. 

The fourth leg-bearing segment, genital complex and abdomen 

appear to be continuous, without segments. The posterior 

corners of the abdomen have or do not have gently rounded 

posterior lobes. The abdomen is 1-segmented. Microscopi- 

cally, the terminal-most seta on the end of the fourth leg is 

twice as long as the other 2 terminal setae. Extreme variation 

in parts, appendages and color have been noted. 

Records - This copepod occurred in Atlantic mackerel (USNM 12620-2), blue 

runner, crevalle jack, inshore remora, shark remora and swordfish off the 

northern USA Atlantic coast. Reports from crevalle jack, king mackerel and 

Spanish mackerel off Texas (Causey 1953b) are in doubt because Cressey and 

Nutter (1987) identified the copepod collected by Causey from the Spanish 

mackerel as a new species of Caligus [although this was not mentioned in a 

subsequent summary Gulf of Mexico Caligus spp. (Cressey 1991)]. 

Geographic Range - Worldwide. Possibly not as common in the tropics. 

Life History - The first larval stage to hatch out of eggs (nauplius) and the 

stage that develops after settling onto the host (chalimus) were described and 

figured by Wilson (1905). 

183

184 


Ecology - This copepod appears to parasitize largely inshore fishes (shark 

remoras and ocean sunfish are exceptions), even the big game fishes infected 

are largely coastal and none of the oceanic tunas are parasitized. Wilson (1905) 

stated that this copepod was the most abundant one on the northeast Atlantic 

coast of the USA, and Kabata (1979) said much the same about Britain. Even 

hosts which bury themselves in the sand or mud cannot dislodge this parasite. 

Both sexes occur on the host and sometimes swim off the host and are caught 

in plankton tows. They are more likely to be off the host at night. This may 

help to explain their great range of hosts. Wilson (1905) suggested that many 

hosts were only temporary resting places (intermediary hosts) between 

preferred hosts. They appear to be more active than other caligoids. Because 

of color variations observed in this parasite, it is reputed to be able to change 

color to agree with the color of the host. Attachments to remoras and ocean 

sunfish probably transmit these copepods out to the open ocean. 

Location in Host - This copepod attaches anywhere on the external surfaces 

of the fish host, but may prefer the dorsal fin. 

Length - Female 5.0-7.0 mm, egg strings 2.6-3.0 mm; male 4.0-5.2 mm; nauplius 

larva 0.4 mm, chalimus stage 2.0 mm increasing gradually to 3.5-4.0 mm. 

Host Specificity - This parasite has no specificity, being recorded from more 

than 100 species of bony fishes and elasmobranchs in 17 orders and 45 families. 

Caligus isonyx Steenstrup and Lütken 

These copepods are so obvious scurrying around 

on the body of great barracuda that you can see them 

while snorkeling. It is a characteristic parasite of this 

host. 

Name - This copepod was described from a single 

female taken from a great barracuda at an unknown 

locality in the West Indies in 1861. Wilson (1905) 

translated the original description and refigured this 

copepod. Cressey and Cressey (1980) synonymized 

it with C. diaphanus Nordmann, but Cressey (1991) 

redescribed and re-established this species. 

Diagnostic Characters - It has widely separated 

lunules. The genital complex is wide (approximately 

80% of cephalosome width) and broadly triangular. 

The abdomen is approximately equal in length to the 

genital complex. 

Records - We found 3-15 females on 28 of 47 great 

barracuda from various localities around Puerto Rico; 

and 9 on this host from off Matthew Town, Great 

Inagua, Bahamas (USNM). It also occurred in Belize 

(USNM); Biscayne Bay, Florida, USA; and Jamaica. 

As in other commercially handled fishes, these 

parasites may be knocked off.


Geographic Range - West Indies. Margolis, Kabata and Parker (1975) 

suggested that it occurs in the New World tropics in the western Atlantic and 

eastern Pacific. 

Associations - Isopods, Excorallana tricornis and Gnathia sp. often occur with 

this copepod. We collected a small inshore remora from a great barracuda 

infected with all 3 of these crustacean parasites. The remora had consumed 

gnathid isopods from the host, but had ignored numerous Caligus isonyx and 

Excorallana tricornis. 

Location in Host - Body, rarely in the gill chambers or mouth. When 

approaching this fish underwater, you can see these parasites scurrying around 

on the outside of a hovering barracuda. 

Length - Female 4.5-5.4 mm; male unknown. 

Host Specificity - This copepod is only known from the great barracuda, and 

it is a primary parasite of this host. A single record from a flyingfish in the 

Galapagos Islands seems unlikely to be the same copepod. 

Significance to Sport Fishing - Back in our young-and-foolish days, we 

speared and ate barracuda. Despite what Anonymous (1992) may say, this is 

a delicious fish. Unfortunately, it is also a dangerous fish to eat because it is 

a well known carrier of ciguatera toxins. The absence of isopods has been 

suggested as a possible indicator of ciguatera fish poisoning. Copepods might 

also serve in this role. 

Caligus lobodes (Wilson) 

Its lobes are so distinctive that it was once placed 

in its own genus based on this character. It 

occasionally occurs at injurious levels on great 

barracuda. 

Name - It is aptly named for the abdominal lobes 

characteristic of this species. It was placed in a new 

genus Midias (for the golden-touch King Midas?) by 

Wilson (1911), but removed to Caligus by Kabata 

(1979). Lewis (1967) erected type specimens from 

Wilson's materials. This copepod should not be 

confused with C. lobatus (=C. productus) named by 

Wilson (1935a) from a pilotfish off the north coast of 

Puerto Rico. 

Diagnostic Characters - This copepod is unique in 

this genus by having small lobes on the first abdominal 

segment. It has small, widely spaced lunules. The 

abdomen (with the lobes) is as wide as the genital 

complex. 

Records - Over 100 females and males occurred under the jaw of a great 

barracuda off Salinas, Puerto Rico; and 2-20 females on 19 of 47 of the same 

host from various localities around Puerto Rico. Forty were found on a great 

barracuda from off Montego Bay, Jamaica (USNM 39613, 112846-7). It 

185

186 


occurred on this host from Florida and Texas, USA; Hawaii (USNM 112888; 

and from the Indian Ocean. 


Associations - Two Excorallana tricornis occurred in the mouth of a host 

which was infected with 2 males of this copepod on the body. 

Location in Host - Body. 

Length - Female 12.0 mm, male 6.3-7.1 mm. 

Host Specificity - This copepod is only known from great barracuda, but is 

only a secondary parasite of this host. The record from the Indian Ocean was 

on an unidentified species of barracuda. 

Damage to Host - The hundred copepods we found on a single host could 

probably cause problems. The fish appeared otherwise healthy and the 

congregation of large copepods had produced no obvious lesions. 

Significance to Sport Fishing - see Caligus isonyx. 

Caligus longipedis Bassett-Smith 

This little known, external copepod parasitizes 

jacks around the world. 

Name - Caligus amplifurcus Pearse is a synonym. 

Diagnostic Characters - It has moderately 

separated lunules. The cephalosome is more than 1/2 

of the total body length. The genital complex is 

wider than long and much longer (more than 4 

times) than the abdomen. The caudal rami are as 

long as the abdomen. Microscopically, it differs 

from all other members of the genus except C. 

rubustus by having crescent-shaped sclerotized 

areas on the last segment of the inside branch 

(endopod) of leg 2. 

Records - One to 26 females and 1-3 males occurred 

on blue runner from Florida, USA, and Belize, and 

on crevalle jack from Florida. It was found on a 

number of Indo-Pacific jacks. 

Geographic Range - Worldwide. We have not found it in the Caribbean. 

Ecology - This parasite occurs on a number of inshore fishes. Its abundance 

offshore is less certain. 

Location in Host - External surface. 


Host Specificity - Jacks appear to be the dominant hosts for this parasite. 

Cressey (1991) suggested that this copepod had only previously been found on 

1 species of Pacific jack, but it had been noted on several species of jacks 

including a blue runner from the Gulf of Mexico (Lewis 1967). Cressey (1991) 

found it on 9 species of coral reef fishes in Florida and Belize, but it appears to 

occur most abundantly on jacks.


Detection - This copepod moves about very actively on the host and may be 

lost from fishes that are not examined soon after capture. 

Caligus mutabilis Wilson 

This unusual parasite uses Spanish mackerels as 

intermediary hosts. It prefers inshore big game fishes. 

Name - "Mutabilis" means changeable and refers to the 

shape of the genital complex. 

Diagnostic Characters - It has an oval cephalosome, 

less than 1/2 the total body length, and with closely 

spaced lunules. The genital complex is longer than 

wide. The abdomen is about twice as long as wide and 

incompletely divided into 2 segments. The egg strings 

are about 1/2 as long as the body. Some variation in 

ornamentation occurred on copepods collected in the 

Atlantic versus those collected in the Gulf of Mexico and 

further south. 

Records - One to 6 immature females occurred on king 

mackerel from Surinam; serra Spanish mackerel from 

Brazil and Costa Rica; and Spanish mackerel from the 

west coast of Florida, USA; and 1 female on a chub 

mackerel from Campeche, Mexico. Cressey and Cressey 

(1980) found immature copepods in 3% of the western 

Atlantic Spanish mackerels they examined. Adults were found on Atlantic 

bonito and skipjack tuna off the Atlantic coast of the USA. Records of this 

copepod in the Eastern Pacific were probably C. omissus Cressey and Cressey. 


Life History - Egg strings have about 50 eggs each. Cressey and Cressey 

(1980) suggested that this copepod utilizes Spanish mackerels (Scomberomorus 

spp.) for its immature forms (intermediary hosts) and other hosts for its adults. 

They found a similar situation with C. biseviodentatus Shen which uses Indo- 

Pacific Spanish mackerels as intermediary hosts and bullet tuna and frigate tuna 

for final hosts. This is a very interesting idea, but the limited data on C. mutabilis 

could as easily suggest that Spanish mackerels are such poor or accidental 

hosts that this copepod cannot develop into adults. Young Spanish mackerels 

might serve as decoy hosts. 

Ecology - This parasite only occurs inshore. 

Location in Host - Gill filaments, gill chamber and mouth. 

Length - Female 5.0-5.8 mm, egg strings 2.0-2.5 mm; male 3.0-3.5 mm. 

Cressey and Cressey (1980) found females 2.8 mm long. 

Host Specificity - This parasite is almost habitat specific to tunas and jacks 

found inshore and on similar, fast-swimming inshore fishes (Cressey 1991) but 

also occasionally occurs on a variety of other fishes. 

187

188 


Caligus pelamydis Krøyer 

This is an important parasite of bonitos worldwide, 

it also occurs on other scombrids. 

Name - This copepod was appropriately named for one 

of its preferred hosts, Atlantic bonito (formerly called 

Pelamys sarda), and not for skipjack tuna (formerly 

Gymnosarda pelamis) as Wilson (1905) stated. Cressey 

and Cressey (1980) redescribed it. Parapetalus sp. of 

Silas and Ummerkutty (1967) is a synonym. 

Diagnostic Characters - The cephalosome has widely 

spaced lunules and is 40-45% of the body length. The 

1-segmented abdomen is 23-25% of the body length. 

The genital complex is roughly triangular. 

Records - One to 11 females with a male occasionally 

associated were found on Atlantic bonito, Atlantic mackerel, 

chub mackerel, and little tunny in the Atlantic; 

and bullet tuna and frigate tuna in the Pacific. Causey 

(1953a,b, 1960) reported this copepod from a wide 

variety of fishes in the Gulf of Mexico, but Cressey and 

Nutter (1987) re-examined his copepods and found all he identified as C. 

pelamydis were other species. 

Geographic Range - Worldwide. Cressey and Cressey (1980) suggested that 

it does not occur in the Indian Ocean, but they also noted that Parapetalus sp. 

from India was the same as this copepod(?), and later Cressey (1991) noted that 

it occurred in all major oceans. It has not been reported from the Caribbean 

possibly because many of its preferred hosts do not occur in this region. This 

may be another copepod that is limited to temperate and cooler waters. 

Life History - The egg strings contain 30-50 eggs each. 

Ecology - Males are seldom found on the host. Scott and Scott (1913) found 

1 male on 1 of 1500 Atlantic mackerel, but Cressey and Cressey (1980) found 

males about 4% as often as females on bonitos and other scombrids. Males 

were always associated with females. 

Associations - Sumner et al. (1913) found that this copepod usually associated 

with C. bonito off the Atlantic coast of the USA. 

Location in Host - Gill filaments, arches or on inner wall of operculum. 

Length - Female 3.0-4.6 mm, egg string 2.0 mm, juvenile female 1.9 mm; 

male 1.9-2.9. Differences in sizes are correlated with water temperature and 

not with the species of host. Larger specimens of this copepod are found in the 

coldest waters, and smaller ones in the warmer waters. 

Host Specificity - This copepod is almost genus specific to bonitos (Sarda), 

a secondary parasite of bonitos, and occurs approximately 13% of the time in 

bonitos around the world. It occurs in lower numbers on other scombrids and 

occasionally on many non-scombrid fishes. Most records from other hosts may 

be erroneous or accidental, but it may occasionally occur on non-scombrids. 

Silas and Ummerkutty (1962) found it was "relatively rare" even on scombrids.


Caligus productus Dana 

This common and important parasite of scombrids 

around the world is remarkably consistent in size. 

Name - Much confusion exists over this name because 

another species of copepod from sharks was accidentally 

given the same name (C. productus Müller). This naming 

error creates what is called a "homonym." Fortunately the 

shark parasite was transferred to another genus and is now 

called Dinematura producta, but these copepods have often 

been confused in the scientific literature. Caligus qalalongae 

Krøyer, C. katuwo Yamaguti, C. lobatus Wilson 

named from Puerto Rico (Wilson 1935a), C. microdontis 

Heegaard, and C. mirabilis Leigh-Sharpe are synonyms. 

Diagnostic Characters - The round cephalosome, with 

moderately separated lunules, is about as wide as long and 

is less than 1/2 of the body length. The posterior 

projections of the genital complex are rounded and extend 

well beyond the beginning (insertion) of the abdomen. The 

abdomen has 1 segment. 

Records - We found 1-20 females and 1-7 males in cero, dolphin, skipjack 

tuna, wahoo, longbill spearfish and white marlin from various localities around 

Puerto Rico (USNM); and 1 female in 1 of 5 dolphin, and 2 females in 1 of 2 

yellowfin tuna off Dauphin Island, Alabama, USA. Twelve females and 2 males 

occurred in a pilotfish off the north coast of Puerto Rico (USNM 64059-60); in 

blackfin tuna from several localities in the West Indies and Nicaragua; cero from 

the Dominican Republic and the U. S. Virgin Islands; little tunny from the Anagada 

Passage and Bermuda; skipjack tuna from the Dominican Republic, Venezuela 

and Brazil; and in yellowfin tuna from the Dominican Republic, Bermuda 

and Brazil. It was found in albacore from North Carolina, USA; skipjack tuna 

from Alabama and New Jersey, USA; bluefin tuna from the Gulf of Mexico; 

and bluefin tuna, dolphin, king mackerel and yellowfin tuna from several localities 

on the east coast of USA. This copepod occurred in bigeye tuna, bullet 

tuna, frigate tuna, other scombrids and great barracuda from the Indo-Pacific. 

Cressey and Cressey (1980) found this copepod on 14 species of scombrids, 

but only commonly on blackfin tuna (92%), skipjack tuna (52%), yellowfin 

tuna (45%), and bluefin tuna (28%). They did not find it on 40 bigeye tuna. 

The records of this copepod from Atlantic bonito are errors and those from 

Spanish mackerel are questionable. Many of the records of this copepod in the 

literature were based on misidentifications. 


Life History - The egg strings contain 30 eggs each. Chalimus stages were 

described by Shiino (1959). 

Ecology - Adults have occasionally been found free swimming. Females are 

3-4 times more abundant than males on the host. 

Location in Host - Mouth, occasionally gills, rarely body. 

189

190 


Length - Female 3.8-6.0 mm, egg string 2.2 mm; male 3.8-4.7 mm. 

Numerous specimens from the Atlantic, Pacific and Indian Ocean were 

remarkably consistent in size. 

Host Specificity - It is almost family specific (Scombridae). This copepod 

infects most western Atlantic species of scombrids except mackerels (Scomber 

spp.). Most records from non-scombrid hosts may be errors, but it may occasionally 

occur on these fishes. This copepod is a characteristic parasite of blackfin 

tuna, a primary parasite of skipjack and yellowfin tunas, and a secondary 

parasite of bluefin tuna. Longbill spearfish and white marlin are new hosts. 

Caligus quadratus Shiino 

This worldwide parasite on all sizes of dolphin is 

occasionally found in other big game fishes. 

Name - It is named for its quadrate genital complex. 

It has been confused with C. coryphaenae, C. productus 

and other species in this genus. 

Diagnostic Characters - It has a round to oval (as 

wide as long to longer) cephalosome with moderately 

separated lunules. The square genital complex has 

parallel sides. The 1-segmented, elongate abdomen is 

longer than the genital complex and about 1/3 as wide. 

Records - Two females occurred in 1 of 5 dolphin 

from off Dauphin Island, Alabama, USA; 3-5 females 

and 1-2 males on dolphin off south Florida and off the 

Atlantic coast of the USA; and in 126 of 145 dolphin 

from the Straits of Florida. It also occurred in 

dolphin, Indo-Pacific sailfish, skipjack tuna and 

yellowfin tuna in the Indo-Pacific. 


Life History - The egg strings contain 30 eggs each. 

Ecology - This parasite seems to infect all sizes of 

dolphin from 3.2 mm juveniles to 1.4 m adults (SL). 

Mean numbers of copepods per host increased 

gradually from 0.2 in the smallest to 38.0 in the largest 

fish. It parasitized the broadest range of sizes with the 

highest numbers of any of the copepods of dolphins. 

Location in Host - This copepod is usually found inside the operculum, but 

occasionally in the mouth, on the body, out side the gill chamber or in the gills. 


Host Specificity - Dolphin is the dominant host of this copepod. It is a 

primary parasite of dolphin, at least along the Atlantic coast of the USA. 

Damage to Host - The ability of this copepod to parasitize all sizes of dolphin 

and to maintain relatively high numbers may allow it to cause disease problems 

in dolphin aquaculture.


Caligus robustus Bassett-Smith 

This is a common parasite of jacks and tunas 

worldwide. 

Diagnostic Characters - It has a round cephalosome 

that is about as long as wide and less than 1/2 the 

length of body. The lunules are moderately 

separated. The fourth leg is robust (the width of the 

first leg segment is more than 1/2 the length). The 

genital complex is roughly triangular and about as 

long as the abdomen. The caudal rami are almost 

square and almost touching. 

Records - Two females and a male were found on a 

bar jack off La Parguera, Puerto Rico (USNM); 2 

females on a blue runner from Jamaica (USNM 42268); 

on bar jack, blue runner and yellow jack from Belize 

(USNM); crevalle jack from Florida and Texas, USA; 

and jacks, yellowfin tuna and other scombrids in the 

Indo-Pacific. 


Location in Host - Body and gill chamber. 


Host Specificity - Jacks are the preferred hosts for this copepod, but it also 

parasitizes scombrids. A few records from fishes other than jacks and tunas 

have been reported. It was oddly ignored in the Silas and Ummerkutty (1962) 

summary of scombrid parasites. Many parasites tend to 

favor tunas or jacks, but this is an unusual host 

combination. 

Caligus spinosus Yamaguti 

This distinctive copepod is a rare parasite of Atlantic 

jacks, but occurs commonly enough to damage Pacific 

amberjacks. 

Name - Both Caligus spinosurculus Pearse and C. 

spinosus were named for the unusual spiny structures 

on either side of the base of the cephalosome. Shiino 

(1960) suggested that C. spinosurculus was a synonym 

of C. spinosus, but did not formally synonymous them. 

The unique and distinctive spiny structures on the 

cephalosome and third leg of C. spinosus and C. spinosurculus 

strongly suggest that they are the same species. 

Most of the other structures of these 2 forms are similar, 

with the exception of slightly closer spaced lunules of C. 

spinosus and other minor differences. Pearse (1951) was 

aware of Yamaguti's description because he quoted his 

191

192 


paper. Why he did not compare these 2 forms is inexplicable. Caligus 

spinosurculus is a synonym of C. spinosus. 

Diagnostic Characters - The round cephalosome (as wide as long) is about 

1/2 as long as the whole body, has moderately separated lunules and distinctive 

spiny structures at the base. The genital complex is vase-shaped and about 1/3 

of the body length. The 1-segmented abdomen is short (about 1/3 of the genital 

complex length). Caudal rami are as wide as long and small (1/5 or less of the 

abdomen length). 

Records - One female each occurred in a crevalle jack and a yellow jack from 

Bimini, Bahamas (USNM 88566). A mutilated copepod, of questionable 

identity, from a rock hind was also reported as this copepod by Pearse (1951). 

It occurs on the commercially important yellowtail and other amberjacks in 

Japan. There is a questionable record in a barracuda from India. 


Location in Host - Gills. 


Host Specificity - The only reliable records of this copepod have been from 

jacks. It could be family specific to jacks. 

Damage to Host - Heavy infections develop in greater amberjack and yellowtail 

held in confinement and culture. This copepod injures and occasionally kills 

these aquaculture fishes. 

Treatment - Masoten has been use to effectively control heavy infections of 

this copepod on cultured yellowtails. 

Significance to Sportfishing - Greater amberjack and yellowtail aquaculture 

is practiced in Japan and Hawaii and may eventually be conducted in other 

areas. This copepod is supposed to be worldwide in distribution, but the forms 

in Japan appear to be particularly damaging. Great care should be taken to 

avoid the dispersion of this copepod to regions where it may not occur. A 

gillworm, causing devastating diseases in cultured fishes, has similarly been 

exported from the USA west coast to Hawaii, Japan and China in greater 

amberjack. 

Caligus wilsoni Delamare-Deboutteville and Nunes-Ruivo 

This practically unknown copepod rarely parasitizes dolphins off the Atlantic 


Name - Wilson (1905) redescribed and refigured copepods from pompano 

dolphin as C. belones Krøyer, and later noted it from dolphin (Wilson 1932). 

Delamare-Deboutteville and Nunes-Ruivo (1958) distinguished the original 

needlefish copepod (C. belones) from the dolphin copepod and named the latter 

for Wilson (C. wilsoni). Yamaguti (1963) re-clouded the issue by noncommittally 

retaining Wilson's pompano dolphin host for both C. belones and 

C. wilsoni. This treatment perpetuated the use of C. belones and neglect of C. 

wilsoni for this copepod of dolphins in the modern literature, to the extent that 

Burnett-Herkes (1974) called dolphin a new host for C. belones, and Palko et 

al. (1982) noted dolphin and pompano dolphin as hosts for C. belones and not


C. wilsoni in a checklist that quoted Delamare- 

Deboutteville and Nunes-Ruivo (1958) as a source. 

Cressey and Collette (1970) did not find any C. 

wilsoni in their survey of copepods of needlefishes, 

although they mention this species. Later, in a paper 

concerning Caligus spp. from the Gulf of Mexico, 

Cressey (1991) decided he did find C. wilsoni in 

needlefish in the north Atlantic. This name has been 

treated in a most confusing and unsatisfactory manner. 

We call the form in dolphins C. wilsoni out of respect 

for the opinions of Dr. Cressey, but we suspect that 

it is a synonym of C. belones. 

Diagnostic Characters - The round cephalosome (as 

wide as long) is about 1/2 as long as the whole body, 

and has moderately separated lunules. The genital 

complex is longer than wide, and longer than the 

abdomen. The 1-segmented abdomen is about 3 times 

as wide as the constricted attachment to the genital 

complex. The caudal rami are short and as long as 

wide. It only differs microscopically from C. belones 

by lacking lateral setae on the last exopod segment 

(end of outer branch) of leg 4. 

Records - Two females occurred on a pompano dolphin from the North Atlantic 

(USNM) and 2 females from a "small" dolphin near Woods Hole, Massachusetts, 

USA; 1 female each in 2 of 145 dolphin from the Straits of Florida; on 

gray snapper, Lutjanus griseus (Linnaeus), from Florida and needlefish, Belone 

sp. in the north Atlantic (the host and original locality for C. belones). 

Only females of this copepod have been collected, but Wilson (1932) 

described the male. He tended to blend old and new information without 

explanation. The description of the male may have been from a previously 

published description of C. belones. Only 9 specimens of this apparently rare 

copepod have ever been recorded. 




Length - Female 4.8-5.0 mm, egg strings 3.0 mm; male unknown, but 

erroneously listed as 4.0-5.0 mm by Wilson (1932). Host Specificity - This 

copepod was known only from dolphins until Cressey (1991) reported it from 

snappers and needlefish. 

Lepeophtheirus bermudensis (Heegaard) 

This rare, odd-shaped copepod on skipjack tuna near Bermuda is virtually 

unstudied. 

Name - This copepod has been placed in 2 other genera (Homoiotes and 

Dentigryps). It is named for the locality in which it was found. 

193

194 


Diagnostic Characters - It has a caligoid cephalosome 

without lunules that is about 3/4 the total 

length. The other segments of the body are reduced. 

The fourth segment is fused with the genital 

complex and covered by the same plate. The 

genital complex is about twice as wide as long. 

The tiny abdomen is shorter than the caudal rami. 

Records - This copepod occurred on skipjack tuna 

from Bermuda. 



Length - Female 3.0 mm, egg strings 1.2 mm; 

male 3.7 mm. 

Host Specificity - This parasite is only known 

from skipjack tuna. 

Lepeophtheirus dissimulatus Wilson 

This copepod is largely a parasite of groupers, but 

is sometimes found on other hosts. It probably only 

accidentally occurs on skipjack tuna and chub 

mackerel. 

Name - The genus name means scale louse. The 

name is from "dissimulo" meaning to conceal what 

exists, and refers to the hidden abdomen. 

Diagnostic Characters - It has a caligoid body 

without lunules, and an oval cephalosome which is 

more than 1/2 of the total length of the body. The 

oval genital complex is wider than long and has a 

series of short projections along its posterior margin. 

The short abdomen is almost entirely concealed by the 

genital complex. The egg strings are not as long as 

the body. The female and male are dark yellow and 

lack pigment spots. 

Records - This copepod occurred on skipjack tuna off 

Bermuda and chub mackerel in the eastern Pacific. It 

has been found on a number of species of groupers and other fishes around the 

world. 



Ecology - This copepod is most often found on benthic groupers. 

Location in Host - Gill cavity or body. 

Length - Female 2.5-3.5 mm, egg strings 1.0-2.0 mm; male 2.0-2.5 mm.


Host Specificity - Groupers (Serranidae) may be the preferred hosts, but this 

copepod has been found on a variety of other bony fishes. Chub mackerel and 

skipjack tuna may be accidental hosts. 

Detection - The color of this copepod blends with that of the host, thus careful 

examination is necessary. 

Lepeophtheirus edwardsi Wilson 

Edward's scale louse has a most peculiar geographic 

distribution and a wide range of hosts. It 

probably only accidentally occurs on crevalle jack. 

Name - This copepod was first found on flounders 

from Woods Hole in 1875, but was not described 

until 1905. The genus name means scale louse, thus 

the name "Edward's scale louse" is intended to 

bestow honor. It is not named for the famous Milne- 

Edwards, as one would suspect, but for the collector 

of the copepods. 

Diagnostic Characters - It has a caligoid body 

without lunules, and an oval cephalosome which is 

more than 1/2 the total length. The oval genital 

complex is longer than wide and has no projections 

along the posterior margin. The abdomen is short 

and not concealed by the genital complex. Egg 

strings are not as long as the body. The female is 

pinkish yellow in life, and not speckled with pigment 

spots like the male. 

Records - One copepod occurred on a crevalle jack 

off Woods Hole, Massachusetts, USA. It has also been found on various hosts 

from the northern Atlantic coast of the USA from Massachusetts, USA, to 

Charles Island, Northwest Territories, Canada; off Bermuda; and China. 


Life History - The egg strings contain 75-80 eggs each. Typical nauplius and 

chalimus stages were described and illustrated by Wilson (1905) and figured by 

Yamaguti (1963). 

Ecology - It is largely a parasite of benthic, inshore flounders, and also occurs 

on groupers, skates and goosefish. When disturbed, males scuttle about over the 

body of the host, while females remain motionless. When placed in an 

aquarium both sexes swim about, but males are more energetic and usually live 

longer. They live for greater periods of time off the host than most other fish 

copepods. Egg strings are not dislodged from females even during rough 

handling, and do not darken when ripe because there is little pigment in the 

larvae. The nauplii are easily hatched in aquaria. 

Location in Host - Dorsal side of the body. 

Length - Female 6.5-7.5 mm, egg strings 5.5-5.7 mm; male 3.0-3.8 mm; 

chalimus stage 2.6-2.8 mm, nauplius stage 0.5 mm. 

195

196 


Host Specificity - Flounders (Bothidae) are the dominant host of this parasite 

(Wilson 1932), but it is occasionally found on crevalle jack and a variety of 

other bony fishes and rays. 

Detection - The color of this copepod blends with that of the host, thus careful 

examination is necessary. 

Parapetalus occidentalis Wilson 

This menacing, caped creature hides in the gill 

chambers in cobia and injures the gill cover. 

Name - "Occidentalis" refers to the western 

hemisphere to contrast with P. orientalis Steenstrup 

and Lütken from the east. Parapetalus gunteri Pearse 

is a synonym. Many of the Indo-Pacific species in 

this genus appear rather similar to this copepod and 

could be synonyms. None of these questionable 

species have been reported from our big game fishes, 

but the genus is in need of revision. 

Diagnostic Characters - The caligoid cephalosome 

with moderately separated lunules is wider than long 

and 25-33% of the length of the body. The genital 

complex is circular and almost as wide as the 

cephalosome and has broad membranous wings 

extending back almost to the end of abdomen. The 1segmented 

abdomen is shaped like an inverted heart 

with membranous wings, similar to those of the genital 

complex, extending posteriorly as far as the caudal 

rami or further. The body is milky white and the egg 

strings develop reddish purple spots when mature. 

Pearse (1952a) used the length of the egg strings 

as a character to separate his species although he 

showed more than 100% variation in this character in 

his few specimens. 

Records - Numerous females occurred in a cobia from North Carolina, USA; 

and 11 females in 8 cobia from Texas, USA. 

Geographic Range - Atlantic coasts of the USA. 

Life History - The egg strings hold 60-100 eggs each. The first larval stage 

(nauplius) that emerges from the egg was described and figured by Wilson 

(1908). 

Ecology - The females swim actively when removed from the host, but not as 

rapidly as Caligus spp. or Lepeophtheirus spp. They swim to the surface and 

float with the aid of a little air under the membranous wings, possibly waiting 

for another host. The wings become easily wrinkled or snarled while 

swimming and floating. 

Location in Host - Gill chamber, particularly inside the gill cover in the dorsal 

corner where 4-5 may congregate close together.


Length - Female 6.0-8.2 mm, egg strings 6.0-13.0 mm; male 4.2 mm; nauplius 

0.4 mm. Wilson (1908) reported that the egg strings were shorter than the 

body, but in his figure they are obviously longer than the body. 

Host Specificity - This copepod is only known on cobia and it is a characteristic 

parasite of this host. 

Damage to Host - This copepod congregates in the upper corner inside the gill 

cover and may produce wounds. 

Tuxophorus caligodes Wilson 

This little collected, external copepod of cobia may 

be spread by intermediate hosts, prey-predator and fishassociate 

transfers. 

Name - Wilson (1908) created a new genus for this 

species. Many of the species subsequently placed in this 

genus have been from big game fishes: T. cybii Nunes- 

Ruivo and T. solandri Kurian (=T. cybii) from Indian 

Ocean wahoo, T. cervicornis Heegaard from Indo-Pacific 

scombrids, and T. collettei. The genus name means to 

bear a plate and refers to the 2 plates over the fourth 

thoracic segment. Caligus aliuncus reported from cobia 

by Causey (1953b) was actually T. caligodes (Cressey 

and Nutter 1987), and the C. aliuncus he reported from 

a crevalle jack was probably also this copepod. Caligus 

remorae Brian from an unidentified remora in the Red 

Sea (not Mediterranean as stated in Yamaguti, 1963) is a 

synonym of this copepod. 

Diagnostic Characters - It has a caligoid body. The 

cephalosome has moderately spaced lunules, is wider than 

long, and about 1/2 the body length. The dumbbell- 

shaped plates on the fourth thoracic segment cover the bases of the fourth legs 

and extend posteriorly over part of the genital complex. The genital complex 

is almost square but has broadly rounded projections of the posterior corners. 

The abdomen is 2-segmented and shorter than the genital complex. The body 

is pale brownish gray with brown or purple spots. 

Records - Females and males occurred on cobia (USNM 32805), females on 

inshore remora (USNM 32806), young females on a pilotfish and chalimus 

stages on a needlefish from North Carolina, USA; both sexes on cobia and 

probably a crevalle jack from Texas, USA. It was also found on jacks and 

other fishes and the Red Sea. 


Life History - The occurrence of chalimus stages on the fins of a needlefish 

could suggest that this copepod settles out of the plankton, attaches and develops 

on an intermediary host, and then moves to a final host. This could be a decoy 

host. Both inshore remoras and pilotfish associate with a variety of big game 

fishes and might transfer this parasite. Each egg string contains about 50 eggs. 

197

198 


Ecology - Females removed from the host swim little and usually remain 

inactive near the bottom of the aquarium or floating at the surface (a possible 

host-seeking behavior). Males are active and constantly swim. 

Location in Host - Body. Five chalimus stage copepods were attached by 

frontal filaments to the anal fin and 1 each to the caudal, dorsal and pectoral 

fins. 

Length - Female 5.7, egg strings 4.3; male 3.6; chalimus 0.8 mm. 

Host Specificity - Cobia is the preferred host, but inshore remoras and pilotfish 

are also hosts for this copepod and associate with cobia. These possible 

transfers from intermediary and decoy hosts suggest some fascinating 

biological processes. This intriguing mystery could be solved with additional 

collections and study. 

Tuxophorus collettei Cressey and Cressey 

Little is known about this long-tailed, external copepod of 

cero. 

Name - Named for Dr. Bruce B. Collette, who made many of 

the collections used in the Cressey and Cressey (1980) study. 


cephalosome has moderately spaced lunules, is about as wide 

as long, and about as long as the genital complex. The wingshaped 

plates on the fourth thoracic segment cover the bases of 

the fourth legs and extend posteriorly over part of the genital 

complex. The genital complex is longer than wide and has 

broadly rounded posterior projections. The abdomen is almost 

twice as long as the genital complex. 

Records - Thirteen to 14 females and a male occurred in each 

of 2 cero caught off the U.S. Virgin Islands (USNM 172250-2). 



Length - Female 9.0 mm, male 6.1 mm. 

Detection - This parasite is probably easily lost from the body 

of its host. This may explain why it has been so rarely seen. 

Host Specificity - This copepod is only known from cero. 

Euryphorus brachypterus (Gerstaecker) 

Light to moderate infections of this copepod are commonly found in the 

gills of larger tuna species worldwide. Very heavy infections cause bleeding 

wounds. 

Name - This copepod was previously placed in genus Elytrophora, and 

juveniles placed in Dysgamus, but both are synonyms of Euryphorus. Arnaeus 

thynni Krøyer, D. longifurcus Wilson, D. sagamiensis Shiino, Elytrophora 

atlantica Wilson, E. hemiptera Wilson, E. indica Shiino; various subspecies 

ending in atlantica, brachyptera, or indica; and possibly Caligeria bella Dana 

are synonyms.



cephalosome lacks lunules and is slightly wider than 

long. Two small sets of shields cover the fourth 

thoracic segment and genital complex. The abdomen 

is shorter than the genital complex. Egg strings vary 

in length from much shorter than the body (as shown) 

to 1/3 longer. The body is yellowish brown, the egg 

strings yellow. 

In colder waters of both the northern and southern 

hemispheres, this copepod has the armature on the 

fourth leg with inner setae more strongly developed. 

This variation seems to be caused by temperature 

rather than being an indication of a different species, 

since it occurs on opposite ends of the world. Temperature-related 

size differences have also been noted in 

other fish-parasitic copepods. 

Records - Six to 15 females and 0-6 males occurred 

in blackfin tuna from St. Thomas; on all large tunas 

(albacore, bigeye tuna, blackfin tuna, bluefin tuna, 

yellowfin tuna) throughout the Atlantic; and on many of these and other tunas 

in the Indo-Pacific. It appears to be more abundant in the Atlantic than the 

Indo-Pacific. 


Life History - Males occur on the gills with the females, but in lower numbers 

(40% as often). Juvenile stages of this copepod have been found on jacks and 

other non-scombrid fishes in the Indo-Pacific. 

Ecology - Offshore, pelagic. 

Associations - In very heavy infections, females often had males attached to 

the ventral surface of their genital complexes. 

Location in Host - They are reported from the inside surface of the gill cover, 

on gill filaments, etc. Very heavy infections seem to target the pseudobranchs. 

The exact location in the gill area is unclear since tunas are not examined 

immediately after capture and these copepods may move or be dislodged before 

they can be collected. 

Length - Female 6.1-12.1 mm, egg strings 6.0-15.5 mm; male 5.0-9.0 mm; 

juvenile 3.9-4.8 mm. 

Host Specificity - This parasite is almost genus specific to the larger tunas 

(Thunnus), but has also been reported from a bonito in the Pacific. 

Damage to Host - Very heavy infections of this copepod have been noted in 

bluefin tuna off Europe. The pseudobranch was completely covered with a solid 

"carpet" of overlapping female copepods, open and bleeding wounds were 

produced in the skin, but the hosts appeared to be otherwise healthy. Densely 

packed copepods covering several square centimeters of gill filaments have also 

been reported in tunas. 

199

200 


Euryphorus nordmanni Milne-Edwards 

This copepod occurs in low numbers on dolphin 

around the world. Although widespread, it is poorly 

known. 

Name - Two copepods, Euryphorus coryphaenae 

Krøyer and E. nympha Steenstrup and Lütken, have 

been considered the tropical cousins of this copepod, 

but they are actually synonyms of E. nordmanni. 

"Electrophora coryphaenae" [a misspelling of 

Elytrophora coryphaenae] was described by Pearse 

(1952a) from a single "female" 5.6 mm long without 

egg strings. The copepod he described and figured is 

actually a male of Euryphorus nordmanni. This 

synonymy has possibly been overlooked because 

Pearse drew his copepods inverted (ventral instead of 

dorsal side up) making them more difficult to 

compare. 

Silas and Ummerkutty (1962) incorrectly call "E. 

nordmanni Kirtising, 1937" (=E. nordmanni Milne- 

Edwards, 1840), a synonym of E. nympha Steenstrup 

and Lütken, 1861. 

Diagnostic Characters - It has a caligoid cephalosome 

that is freckled with pigment spots, but lacks lunules. A butterfly-shaped 

plate lies over the fourth thoracic segment. The round genital complex is as 

wide or wider than the cephalosome. The elongate abdomen has large, wingshaped 

expansions. 

Records - One female occurred on 1 of 20 dolphin examined off La Parguera, 

Puerto Rico, and it has been collected numerous times in this host around the 

world. One was found in a cobia from Port Aransas, Texas, and 1 in 1 of 18 

dolphin from Texas, USA (USNM 92673). It also occurred in albacore, bigeye 

tuna, pompano dolphin, yellowfin tuna in the Pacific. 

Geographic Range - Worldwide between 40°S and 40°N. Our collections are 

the first in the Caribbean. 

Life History - Each female often has a male attached to its genital complex 

with the ventral surfaces in contact and body axes aligned. 


Location in Host - Inner surface of operculum, gill chamber or mouth. 


Host Specificity - This parasite prefers dolphin. It is a secondary parasite of 

this host based on the collection records, but may occur more abundantly and 

be lost from the host before it is examined. The single reports from cobia, 

tunas and 1 other fish may be accidental infections or false hosts.


Genus Gloiopotes Steenstrup and Lütken 

These are relatively large, caligoid-shaped copepods distinguished by their 

ornamentation and colorfulness (blue bodies and red egg strings) and the absence 

of lunules. Species in the genus only occur on billfishes (Istiophoridae), 

swordfish (Xiphidae) and wahoo (Scombridae) but not other scombrids. 

These copepods are more host specific than most of the other copepods 

that attach externally to big game fishes. Their presence and abundance has 

been used to suggest relationships among their hosts (see Discussion). Records 

from odd hosts, such as bigeye tuna, were probably based on misidentifications. 

Williams (1978) reported the first record of striped goose barnacle 

attached to a fish-parasitic copepod that was not permanently embedded in its 

host. Causey (1960) reported a similar instance of what was possibly this 

barnacle, Conchoderma sp., on Gloiopotes costatus Wilson [=G. huttoni 

(Tomson)] on sailfish off Mazatlan, Mexico. 

These copepods do not appear to damage the host even when they occur in 

large numbers, although the thousands per sailfish reported by Williams (1967) 

off east Africa, would certainly have injured these hosts. The life history is 

unknown. It may be more complicated than the normal copepod life cycle. 

Simply dispersing the few larvae into the vast ocean would seemingly quickly 

extinct these parasites. They are not known to bite or otherwise harm humans. 

Records of the number of copepods (geographic position caught, species 

and size of host) from fishermen would help us understand the number of hosts 

parasitized and the number of copepods per host. These external copepods are 

so easily lost that our examinations hours after the hosts are caught may have 

little relation to the original infection. 

References - Cressey (1967) revised the genus. 

Yamaguti (1963) incorrectly noted that these 

copepods also occurred on elasmobranchs (sharks and 

rays), but listed none. He also described the first leg 

of these copepods as uniramous (with 1 terminal 

projection) when they are biramous (2 projections). 

Gloiopotes americanus Cressey 

These large and brightly colored copepods are 

found on the skin of sailfish and may be useful as 

biological tags. 

Name - This parasite was confused with Gloiopotes 

ornatus until it was separated and described by 

Cressey (1967). 

Diagnostic Characters - Live copepods have a 

bluish-purple body and red egg strings. It is rather 

similar in appearance to G. ornatus, but differs by 

occurring on different hosts, by having a shorter fifth 

leg (does not extend to or beyond abdomen), and by 

201

202 


other technical characters (see G. ornatus). It differs considerably from G. 

hygomianus by having conspicuous ornamentation and in body shape. The egg 

strings curl back over the posterior of this copepod in a U-shape (We show them 

flattened to avoid obscuring details of the body). Oddly, Cressey (1967) did not 

mention this characteristic. 

Records - We found a female and male on 1 of 2 Atlantic sailfish caught off 

Arecibo, Puerto Rico (USNM); and 1 female on each of 2 Atlantic sailfish off 

Dauphin Island, Alabama, USA. This parasite probably always occurs on sailfish, 

but rinsing, drying and rough handling may knock most of them off before 

fish are examined. It is also known from the Atlantic coast of Florida. 

Geographic Range - Tropical and subtropical Atlantic. Our collection is the 


Ecology - Pelagic and offshore. 

Location in Host - Body. We found it on the gills of 1 sailfish, but all 

previous records were external, and our few specimens may have been washed 

onto the gills when fishermen rinsed the sailfish. 

Length - Female 10.0-11.7; male 8.8-10.0 mm. 

Host Specificity - This parasite only occurs on Atlantic sailfish and is probably 

a characteristic parasite of this host. 

Significance to Sportfishing - The apparently restricted geographic range 

might make it a useful biological tag to distinguish stocks or movements of 

sailfish. 

Gloiopotes hygomianus Steenstrup and Lütken 

This moderate-sized copepod occurs on wahoo 

around the world. It is less ornamented than others in 

the genus, but is still quite colorful. 


bluish-purple body and red egg strings. This copepod 

appears naked compared to the other 2 conspicuously 

ornamented species of Gloiopotes from Atlantic billfishes. 

Records - We found 6-20 females and 1-2 males on 6 

of 15 wahoo from various localities around Puerto 

Rico; and 2-5 females on 4 wahoo off Dauphin Island, 

Alabama, USA. Cressey (1967) also noted this 

copepod from Puerto Rico (USNM). Iversen and 

Yoshida (1957) found this copepod in 52 of 96 

(54.2%) wahoo off the Line Islands in the Pacific. A 

record from albacore in the Pacific is probably an 

error. 


Ecology - Pelagic and offshore. 

Location in Host - Body.



Copepods from Hawaii are smaller than those from other localities worldwide. 

Host Specificity - It only occurs on wahoo and is a characteristic parasite. 

Gloiopotes ornatus Wilson 

These large and brightly colored parasites 

scamper around on the skin of all Atlantic blue 

marlin. 

Name - One of our former graduate students called 

this the "red, white and blue" copepod for all its 

bright colors. It is a spectacularly beautiful animal 

and large enough for sport fishermen to enjoy its 

color. The name "ornatus" describes the elaborate 

ornamentation on this copepod. 


bluish purple body and red egg strings. The rather 

similar Gloiopotes americanus only occurs on Atlantic 

sailfish. Microscopically, adult females of G. 

ornatus differ from G. americanus by having a 

distinct bulbous lateral expansion of the second 

abdominal segment, and more pronounced swollen 

lateral areas of the genital complex. It differs 

considerably from G. hygomianus by having 

conspicuous ornamentation and in body shape. 

Records - We found 25-150 on 40 Atlantic blue marlin; and 2-10 on 3 white 

marlin from various localities around Puerto Rico (USNM). They also 

occurred on these hosts off Havana, Cuba; Brazil; and Woods Hole, 

Massachusetts, USA. Previous records from "swordfish" (USNM 6209) and 

"spearfish" appear to refer to blue marlin and white marlin, respectively. 


Ecology - Strictly an offshore pelagic parasite. 

Location in Host - Body. This copepod is sometimes found in the mouth or 

gill cavities, but this may represent movement after host death, or handling 

contamination. 


Host Specificity - This copepod is only known from blue marlin and white 

marlin. It is family specific to billfishes (Istiophoridae) and a characteristic 

parasite of blue marlin and possibly white marlin. 

Hatschekia amplicapa Pearse 

This small copepod infects the gills of great barracuda. 

Diagnostic Characters - The round cephalosome is attached directly to the 

elongate trunk region. The caudal rami are relatively tiny. 

203

204 


Records - Two to 10 females occurred in 6 of 47 great 

barracuda from various localities around Puerto Rico 

(USNM); and 8 females in a great barracuda from Bimini, 

Bahamas (USNM 88602). 

Geographic Range - West Indies. Our records are the 

first from the Caribbean. 

Life History - The egg strings contained 16 relatively 

large eggs each. 

Ecology - This genus of copepods is usually limited to 

inshore and coral reef fishes, but H. oblongata Wilson 

was once reported from a Pacific crevalle jack, Caranx 

caninus Günther, in the eastern Pacific; and we found a 

Hatschekia sp. once in a little tuna off La Parguera, Puerto 

Rico. The "H. mulli" reported by Causey (1955) on king 

mackerel from Louisiana, USA, was actually 

Pseudocycnoides buccata. 


Length - Females 1.0-1.3 mm, male unknown. 

Host Specificity - This parasite only occurs on great barracuda. We found this 

copepod rarely on French grunt, Haemulon flavolineatum (Desmarest), from La 

Parguera, Puerto Rico (USNM). French grunt is a new host for this copepod, 

but these may have been only accidental infections. 

Pseudocycnus appendiculatus Heller 

This elongate copepod is found on the gills of tunas, 

and a few other big game fishes worldwide. It grows 

larger in colder waters. 

Name - Pseudocycnus spinosus Pearse and P. thynnus 

Brandes are synonyms. 

Diagnostic Characters - It has a distinctively long 

genital complex that is as wide as the cephalosome. The 

caudal rami are long, 1/2 of the length of the genital 

complex. The body is bright red on the dorsal surface 

and yellow-brown on the rest of the body. The egg 

strings are brown. 

Records - We found 1 female and 1 male in 1 of 2 

blackfin tuna, and 2 females in 1 of 40 dolphin from 

various localities around Puerto Rico; 2 females in a 

bluefin tuna, and 2 females in 1 of 2 yellowfin tuna off 

Dauphin Island, Alabama, USA. One to 5 females 

(usually 2) often, and 1-2 males (average 0.3) rarely, 

occur on all large tunas (albacore, bigeye tuna, blackfin tuna, bluefin tuna, 

yellowfin tuna), little tunny and skipjack tuna; and occasionally on Atlantic 

bonito and dolphin from the Atlantic, and some of these hosts and other tunas, 

little tunas and bonitos from the Indo-Pacific.




Length - Female 7.3-24.0 mm, usually 13.0-17.0; male 2.7-5.0 mm. The 

longer specimens of females (18-24 mm) occurred in regions of colder waters. 

The very smallest (7.3 mm) were described from Atlantic bonito. 

Host Specificity - This parasite is almost family specific to scombrids, but 

rarely, and possibly accidentally, occurs in dolphin. Tunas (Thunnini), little 

tunas (Katsuwonini) (little tunny and skipjack tuna in the Atlantic) are the 

preferred hosts, with an infection rate of approximately 10%; but 2% of the 

bonitos (Sardini) examined by Cressey and Cressey (1980) worldwide were 

also infected. This copepod is a secondary parasite of tunas and little tunas. 

Pseudocycnoides buccata (Wilson) 

This parasite occurs in all species and on most specimens of 

Spanish mackerels throughout the Americas. The cumulative loss of 

these valuable sport fishes due to this parasite must be substantial. 

Name - Cybicola elongata Pearse, Hatschekia mulli of Causey, 1953 

and Pseudocycnus elongatus (Pearse) are synonyms. The genus 

Pseudocycnopsis Yamaguti, once used for this parasite, is a synonym 

of Pseudocycnoides Yamaguti. 

Diagnostic Characters - It has a distinctively long genital complex 

that is as wide as the cephalosome. The caudal rami are short, about 

1/10 the length of the genital complex. The first 3 pairs of legs are 

rudimentary. The first pair is located between the bases of maxillipeds 

and not easily seen. The body is red and the egg strings brown. 

Records - We found 1-5 females in 18 of 35 cero and 8 of 14 king 

mackerel from various localities around Puerto Rico (USNM); and 1-2 

in 1 of 2 king mackerel and 1 each in 2 of 4 Spanish mackerel from 

off Dauphin Island, Alabama. Cressey and Cressey (1980) suggested 

that this copepod occurred commonly in light to moderate infections 

on all its hosts (cero, king mackerel, serra Spanish mackerel, Spanish 

mackerel, and Eastern Pacific Spanish mackerels) (USNM 88538). 

Geographic Range - Western Atlantic and Eastern Pacific. 

Life History - Males are seldom seen on the host (6-7% of infected hosts). 

Ecology - This copepod is very similar in appearance to the gill filaments of its 

host, possibly an adaptation to conceal it from cleaner fishes such as remoras. 

Location in Host - Gill filaments, sometimes on pseudobranchs. 

Length - Female 4.0-5.3 mm; male 1.9 mm. Causey (1960) found those in the 

Pacific to be slightly smaller than the ones in the Gulf of Mexico. 

Host Specificity - This parasite is genus specific (Scomberomorus), and is a 

primary parasite of these fishes. 

Damage to Host - Low to moderate infections do little damage to individual 

hosts, but the high prevalence on Spanish mackerels means that minor damage 

occurs throughout the population. Slight growth reduction in hundreds of 

millions of mackerel translates to the loss of millions of kilos of valuable fishes. 

205

206 


Lernanthropus giganteus Krøyer 

This relatively large, obvious copepod occurs on 

jacks around the world in the tropics. It damages the 

gills and heavy infections stunt or kill jacks. 

Name - The specific name obviously refers to its 

large size. Lernanthropus paenulatus Wilson only 

differs from this copepod by having a slightly longer 

shield or skirt and we consider it a synonym. 

Diagnostic Characters - Species in this genus are 

generally cylindrical in shape and have distinctive 

elongate projections from the ventral side of the 

trunk. The ventral side of this species is illustrated so 

that these projections can be seen. The fourth 

segment has an expanded plate on the dorsal side that 

partially covers the projections. The body is beige. 

Records - We found 1 female each in 1 of 3 blue 

runner from Aguadilla, Puerto Rico; 1 of 19 bar jack, 

in 3 of 20 crevalle jack and in 5 of 28 horse-eye jack 

from various localities around Puerto Rico; in a 

yellow jack from St. Lucia; and in 1 of 2 crevalle 

jack from Gulf Shores, Alabama, USA. Bunkley-Williams and Williams 

(1995) found about 15% of crevalle jacks and horse-eye jacks in Puerto Rico 

infected with 1 female copepod each. Both sexes of this copepod occurred in 

blue runner (USNM 42277) and crevalle jack (USNM 42282) from Jamaica. A 

male and a female was reported from a crevalle jack from Texas, USA; and 

both sexes were found on a greater amberjack from Massachusetts and North 

Carolina, USA. It is also reported in other jacks from the tropical Indo-Pacific. 




Host Specificity - This parasite is family specific to jacks. It is a secondary 

parasite of crevalle jack and horse-eye jack. Bar jack, horse-eye jack and 

yellow jack are new hosts. 

Damage to Host - The gill filaments surrounding this copepod have thickened 

surfaces (epithelium). 

Significance to Sportfishing - This is a large and damaging copepod. It 

usually occurs as a single female or as a female-male pair on a host. These low 

levels cause little damage. Heavier infections could stunt or kill jacks. 

Lernaeenicus longiventris Wilson 

This giant, common, and impressive copepod embeds its head in the flesh 

and debilitates dolphin and other big game fishes. The body trails across the 

back of its host like a ribbon. 

Name - It is appropriately named for the unusual shape of the abdomen 

(longus=long, venter=abdomen).


Diagnostic Characters - This is a very large, elongate copepod 

with a T-shaped cephalosome embedded in the flesh of the host. 

The long narrow posterior projection from body is almost as long 

and is similar in appearance to the egg strings. The "neck" is 

longer than the body. The body is white, the cephalosome bright 

pinkish red and the eggs strings are maroon when ripe. 

Records - We found 1 female each on 1 of 19 bar jack, 4 of 20 

crevalle jack, and 1 of 27 horse-eye jack from various localities 

around Puerto Rico; in 3 of 7 horse-eye jack from Joyuda Lagoon, 

Puerto Rico (USNM); in a horse-eye jack from St. Lucia; in 1 of 3 

crevalle jack, and in 1 of 5 dolphin from off Dauphin Island, 

Alabama, USA. It is reported to commonly occur on many dolphin 

off North Carolina, Louisiana and Texas, USA, and in the 

northwest Atlantic (USNM 47800, 47806), and off the west coast 

of Africa; also on greater amberjack, blue runner (USNM 2107, 

47804-5), cobia, crevalle jack (USNM 42346), pompano dolphin 

(USNM 47801), Spanish mackerel (USNM 6192, 47802), off the 

Atlantic and Gulf coasts of USA; and serra Spanish mackerel off 

Brazil. Usually 1 female occurs per host, but we found 9 on a liza, 

Mugil liza Valenciennes. Causey (1953) found 25% of crevalle 

jack off Texas, USA, were infected. 


Life History - Eggs in the egg string which are beginning to hatch 

gradually become a dark maroon color. A newly hatched nauplius 

stage was figured by Wilson (1917). 

Behavior - The female contracts its neck when touched (tactily 

stimulated), pulling 1/2 to 2/3 of the exposed body (genital complex) 

into the host. This might be a defensive reaction to make a smaller 

target for cleaner fishes, or to prevent damage to this long-trailing copepod 

during violent feeding or swimming bouts by the host. 

Associations - The mechanism preventing a second female from infecting a 

host has not been determined. Simple rarity of infective stages does not appear 

to be the answer. If the female is excluding others, this highly interesting 

process should be examined and explained (see Damage to Host). The exposed 

body of this copepod is often covered with hydroids and algae. 

Location in Host - This copepod embeds its head in the host's body usually 

along the dorsal or anal fin near the tail. It was reported once embedded in the 

operculum. 

Length - Females 40.0-50.0 mm, egg strings 10.0-15.0 mm; nauplius stage 

0.25 mm. 

Host Specificity - Crevalle jack, dolphin and pompano dolphin are the 

preferred hosts of this parasite, but it is only a secondary parasite of these hosts. 

It occurs occasionally to rarely on a variety of fishes including many big game 

fishes. Bunkley-Williams and Williams (1995) reported horse-eye jack as a new 

207

208 


host, and 2 other new hosts that were not big game fishes. Bar jack and serra 

Spanish mackerel are additional new hosts. 

Damage to Host - A fibrous membrane forms around the embedded portion 

of the copepod. This large parasite may be detrimental to the host, but the 

extent of injury has not been studied. The normal infection of 1 female per host 

could suggest that fishes cannot survive multiple infections. 

Detection - This long copepod spectacularly streams off the back of its hosts. 

Preparation for Study - The embedded neck and cephalosome must be 

carefully dissected from the host tissues. 

Significance to Sportfishing - This parasite appears to damage many big 

game fishes. Sport fishes infected with this parasite are said to not put up the 

same fight on hook and line as unparasitized fish (Causey 1955). 

Lernaeolophus striatus Wilson 

This large, and long lost, tissue-embedding 

copepod must cause considerable damage to great 

barracuda. 

Name - "Striatus" means fluted or grooved and refers 

to the sides of the cephalosome. 

Diagnostic Characters - This large copepod has 

dendritic projections from the fluted cephalosome. The 

neck and cephalosome are longer than the trunk and 

are embedded in the tissue of the host.. The anterior 

body and horns are a pale wine-red, the trunk is 

white and the appendages pale yellow. A dense mat of 

thick caudal projections is present on the posterior 

end of the body. 

Records - We found 1-2 females in 4 of 44 great 

barracuda from various localities around Puerto Rico 

(USNM). Two females were found in a barracuda 

from Jamaica (USNM 43320). This copepod was 

collected in Jamaica in 1910 (described in 1913) and 

has not been reported again until now (86 years). 

Geographic Range - Caribbean. 

Location in Host - This copepod attaches in the throat just inside the lower 

jaw, with their cephalosomes embedded near the aorta and their bodies trailing 

back along the gill arches. 

Length - Female 27.0-31.0 mm, male unknown. 

Host Specificity - This parasite is only known from great barracuda. Few 

tissue embedding copepods are specific to a single host. 

Damage to Host - This large, tissue embedding copepod must be harmful to 

great barracuda. Infections of more than a few copepods per host could be fatal. 

Preparation for Study - The cephalosome and neck must be carefully teased 

from the host tissues. The caudal projections should be examined for algae and 

other encrusting organisms.


Genus Pennella Oken 

These large elongate parasites attract a lot of attention with their bodies and 

egg strings sticking several feet out of hosts. They were noted in classical 

antiquity, medieval literature, and will probably eventually be recognized in the 

earliest, prehistoric coastal cave paintings of fishes. Aristotle wrote of these 

parasites, which were undoubtedly the first fish-parasitic copepods acknowledged 

by humans. Small ones can be mistaken for scientific fish tags. The name 

Pennella in Latin means "little feather" [penna=feather, -ella=little 

(diminutive)] and refers to the feather-shaped posterior projections. Many 

species in this genus vary extremely in shape and have little host specificity. 

The shape of the cephalosome and horns, and length of neck and trunk, have 

been used to distinguish species, but are unreliable in most cases. Some 

structures change with maturity or vary with the firmness of the surrounding 

host tissue. Older copepods are more elongate. A few specimens of these 

interesting copepods are often saved out of curiosity, but adequate numbers and 

assortments for comparison of these offshore and ephemeral copepods have 

usually eluded taxonomists, resulting in insufficient and repeated descriptions 

(synonyms). The genus became bloated with so many synonyms, poorly defined 

and questionable species that individual Pennella sp. specimens were quite 

difficult to identify to species. Recently, Hogans (1988b) reviewed and 

stabilized this genus and consolidated it into 7 species. 

Females of this genus have a long cylindrical body. The cephalosome 

(armed with 2-3 projections or horns) and neck are embedded through the skin 

or fins of the host into the muscle or other internal organs. The body extends 

from the host and terminates in a dense "beard" or feather-like mat of simple or 

branched lateral projections. Older copepods have larger posterior beards. The 

egg strings are straight and are often as long as or longer than the body. This 

length may be related to the necessity of producing many eggs in the pelagic 

environment where hosts are so sparse and to support a complex two-host life 

cycle. The eggs hatch and release nauplius larvae, a series of larval stages 

develop in the plankton, females settle onto hosts, penetrate the body, and 

develop into adults. Immature females parasitize decoy host squids (Pennella 

varians Steenstrup and Lütken has been found on common cuttlefish, Sepia 

officinalis Linnaeus; elegant cuttlefish, Sepia elegans Blainville; and European 

squid, Loligo vulgaris Lamarck, in the eastern Atlantic). This is probably a 

strategy for finding sparse fish hosts in the open ocean as squid is a prey item 

for big game fishes. It is also an example of a permanent prey-to-predator 

parasite transfer. Males are not parasitic on hosts. Other organisms are found 

attached to the exposed portions of these copepods. Up to 18 striped goose 

barnacles have been found attached. Filamentous algae, hydroids and possibly 

other encrusting organisms grow in the beard at the end of the body and 

sometimes cover it in dense mats. 

All 7 known species may occur in Puerto Rico. Two of the 4 species in big 

game fishes are only known from Puerto Rico (see below). This does not mean 

209

210 


that Puerto Rico is a speciation center for copepods, but just indicates our poor 

state of knowledge. Pennella exocoeti (Holten) occurs on flyingfishes 

(Exocoetus spp.) preyed upon by many big game fishes worldwide and will 

probably be found off Puerto Rico. We collected P. diodontis Oken on a local 

spiny puffer, Diodon holacanthus Linnaeus (USNM, new host and locality). 

The other 2 known species are noted below. Most of these copepods are 

worldwide or at least circumtropical/temperate in distribution. Life histories 

are largely unknown. Many species occur quite sporadically and unpredictably 

on various hosts. These copepods are largely offshore and pelagic. Only P. 

diodontis has inshore, shallow-water hosts. The smallest and most delicate P. 

sagitta (Linnaeus) (2.2 cm long) occurs on sargassumfish, Histrio histrio 

(Linnaeus), or frogfishes. The largest copepod in the world, P. balaenoptera 

Korea and Danielson, occurs on the largest animals on Earth, baleen whales, 

Balaenoptera spp. The body of this copepod is longer than 20 cm and is 

rumored to exceed 1 foot (30.5 cm) with egg strings many feet long. Both of 

these copepod species could occur off Puerto Rico, but may be restricted to 

areas further north because we have not found them on sargassum fish and 

whales we examined. 

The cephalosome of this parasite is often located near vital blood vessels 

or organs where it has an ample source of blood. This placement and 

subsequent damage can be disastrous for small or developing hosts. Host 

reaction can be extreme with enormous cysts occurring around the parasite. 

Many young big game fishes may be killed or debilitated and perish from 

predation because of this parasite. Massive infections that have been rumored, 

but never documented, could kill adult big game fish. The pathological impact 

of these parasites has never been studied, but many authors have suggested that 

they weaken hosts, particularly big game fishes. These parasites are harmless 

to humans. 

The adult female protruding from the side of a host is obvious. Settling 

juveniles, males or developing females must be detected with skin scrapings 

viewed with a microscope. To correctly identify these copepods, the 

cephalosome and its complicated and delicate projections must be dissected out 

of the flesh intact. Simply pulling the copepod out of the host usually breaks 

the copepod at the neck and the cephalosome and neck remain in the flesh of 

the host. Even if the cephalosome is successfully pulled out (5-10% success), 

most of the projections will break and remain in the host. Cutting a large chunk 

of flesh out of the host around the parasite is the best method of saving these 

copepods. Save as many samples as possible. More than 1 species may be 

present on the same host. Many specimens may be needed to identify the 

copepod. Remember copepods that look different, are different sizes, or are 

attached in different positions on the host may represent different species. 

Photographs of unusual groupings of copepods or heavy infections would be 

useful in understanding these parasites.


Pennella filosa (Linnaeus) 

This large, worldwide parasite hangs out of the side of 

many big game fishes, particularly swordfish. 

Name - "Filosus" means thread-like. The only information 

of use in the original description by Linnaeus was the name 

of the host. Pennella costai Richardi, P. crassicornis 

Steenstrup and Lütken, P. germonia Leigh-Sharpe, P. 

germonia fagei Poisson and Razet, P. histiophori 

Thompson, P. orthagorisci Wright, P. pustulosa Baird, and 

P. remorae Murray are synonyms. 

This copepod is morphologically identical to P. 

balaenoptera that occurs on whales. They are essentially 

separated on the basis of their hosts. Morphologically 

identical species are rare but can exist if the biological 

differences maintain isolation. This mystery warrants 

additional study. 

Diagnostic Characters - The female is very large (greater 

than 50 mm long). The large cephalosome is bulbous to 

cylindrical and has 3 short (less than the width of the 

cephalosome) unbranched projections or horns which protrude and extend 

perpendicular to the neck. The flat or truncated portion of the cephalosome is 

completely covered with papillae. The cephalosome and neck are yellow and 

the trunk is dark brown. 

Records - We found 1-50 females on 6 of 40 Atlantic blue marlin, 1 female on 

1 of 3 swordfish and 5 females on 1 of 3 yellowfin tuna from various localities 

around Puerto Rico. It has also been found on albacore, Atlantic sailfish, 

bluefin tuna and white marlin around the Atlantic and the world; on ocean 

sunfish; and on an unidentified dolphin off Australia. 

Hogans, Brattey and Hurlbut (1985) found 182 of 303 swordfish in the 

northwest Atlantic infected with either this copepod or Pennella instructa. A 

maximum of 15 copepods occurred on a single host. A few protruded from 

scar tissue, although whether they caused the damage or were attracted to it was 

not clear. Only 12 copepods were dissected from these hosts and identified, 

thus the numbers of each species, association or competition between them, 

size ranges and location preferences was not determined. The Pennella sp. of 

Cressey and Cressey (1980) in 3 of 57 bluefin tuna and in 1 of 61 wahoo were 

probably P. filosa. 

Geographic Range - Worldwide. Our collections are the first from the Caribbean. 

Life History - Young females have only 2 horns on the cephalosome. The 

third (dorsal) horn develops last and is always smaller. 

Ecology - Ocean sunfish, flyingfishes and pilotfish are probably important in 

maintaining population levels in big game fishes of this pelagic, open ocean 

species. Copepods in ocean sunfish with relatively soft muscle tissue develop 

211

212 


bulbous cephalosomes, while those in relatively tough-muscled tunas develop 

more cylindrical cephalosomes. 

Associations - This copepod is found in tight groups of 8-25 specimens with 

12-50 P. makaira on the skin of Atlantic blue marlins off Puerto Rico. 

Pennella instructa is also found on swordfish, but these 2 copepods have not 

been reported from the same host specimen. This parasite occurs on remoras in 

the Atlantic and Mediterranean, which in turn attach to many of the big game 

hosts of this copepod. These associations may allow new hosts to be infected 

with males or immature females. 

Location in Host - Skin, penetrating into muscle (subcutaneous musculature), 

of the dorsal, lateral and ventral surfaces of the body, but not on the head. 

Length - Female 15.0-20.0 cm, egg strings 12.5-35.0 cm and can be twice the 

body length or more; immature female 9.4 cm; male unknown. 

Host Specificity - The early literature is too inconsistent about copepod and 

host identifications, and modern collections are too limited to determine exact 

host preferences, but swordfish and ocean sunfish may the preferred hosts. 

Atlantic blue marlin is a new host. 

Damage to Host - Sumner et al. (1913) found infections of 30-40 copepods per 

host impaired the vitality of the fish. Hogans, Brattey and Hurlbut (1985) found 

that this copepod weakened the host by damaging the swimming muscles. 

"There is 1 species (Pennella filosa) in the Mediterranean, 7 or 8 inches long, 

which penetrates the flesh of the swordfish, the tunny, and the 

sunfish, and torments them horribly." (Cuvier 1830). 

Pennella instructa Wilson 

This is a very large parasite found in the side of swordfish in 

the western North Atlantic. Sport fishermen could help to 

determine its exact distribution and importance. 

Name - "Instructus" means arranged in definite order and refers 

to the cephalosome papillae. Pennella zeylandica Kirtisinghe is a 

synonym. This species was redescribed by Hogans (1986). 

Diagnostic Characters - The female is very large (greater than 

50 mm long). The bulbous to cylindrical cephalosome has 2 long 

(greater than the width of the cephalosome) unbranched projections 

or horns which protrude and extend posteriorly and parallel to the 

neck. The flat or truncated portion of the cephalosome is 

partially covered with papillae. The cephalosome and neck are 

yellow, the trunk dark brown. 

Records - One to 8 females occurred in swordfish from off 

Massachusetts (USNM 47751) to Maine (USNM 47750), USA, and 

in the northwest Atlantic. Up to 4 copepods commonly occur in a 

single host (see "Records" in Pennella filosa). It is also found in 

black marlin and 2 other species of Pacific billfishes. 

Geographic Range - Worldwide. However, the records are widely spaced 

from the northwest Atlantic, Indian Ocean, and Australia.


Ecology - This species is less common than Pennella filosa. A cylindrical 

cephalosome is found in copepods embedded in hard tissues such as the aorta 

or heart, while a bulbous cephalosome is found in those in soft tissues. 

Associations - Pennella filosa is also found on swordfish, but these 2 copepods 

have not been reported from the same host specimen. This copepod is often 

covered with algae and hydroids but not striped goose barnacles. 

Location in Host - Skin, penetrating through muscle (subcutaneous musculature) 

and attaching in internal organs of the body cavity and major blood vessels 

of the dorsal, lateral and ventral surfaces of the body, but not on the head. 

Length - Female 20.0-25.0 cm, egg strings vary from shorter to longer than the 

body, 10.0-33.0 cm. Two females described by Hogans (1986) were much 

smaller, 14.3 and 15.3 cm long. 

Host Specificity - This copepod is only known from swordfish and may be a 

primary parasite. 

Damage to Host - A thick fibrous cyst up to 5 cm in diameter often forms 

around this copepod. One formed a large cyst in a swordfish ovary. The aorta 

and heart can be injured by this copepod. Hogans, Brattey and Hurlbut (1986) 

found that this copepod weakened the host by damaging the heart. 

Pennella makaira Hogans 

This is an uncommon large copepod that sometimes 

occurs in heavy infections on Atlantic blue marlin near 

Puerto Rico. 

Name - This copepod was described with specimens collected 

off Puerto Rico. 

Diagnostic Characters - The female is the smallest 

reported on billfishes (less than 50 mm long). All other 

members of this genus from billfishes are greater than 50 

mm long. The bulbous cephalosome has 2 long (greater 

than the width of the cephalosome) unbranched projections or 

horns which protrude and extend posteriorly at an oblique 

angle to the neck. The flat or truncated portion of the 

cephalosome is completely covered with papillae. The 

neck is shorter than the trunk. The cephalosome and neck 

are yellow, the trunk dark brown. 

Records - We found 12-50 females on 10 of 40 Atlantic 

blue marlin from various localities around Puerto Rico. 


Associations - This copepod occurs in tight groups of 12- 

50 specimens among 8-25 P. filosa on the skin of Atlantic 

blue marlins off Puerto Rico. The gregariousness of this species could be due 

to preferred sites on the host, attraction to adult individuals, and/or the physics 

of settling onto an actively swimming host. A striped goose barnacle attached 

to a P. makaira on an Atlantic blue marlin off Arecibo, Puerto Rico. 

Location in Host - Skin and muscle. 

213

214 


Length - Females 27.0-29.0 mm; male unknown. 

Host Specificity - This copepod is only known from the Atlantic blue marlin, 

but occurs so seldom that it is only a secondary parasite. 

Significance to Sport Fishing - Its abundance in Puerto Rico and possibly the 

Caribbean, apparent absence in the Gulf of Mexico and other parts 

of the Atlantic, relative long life in the tissue of the host, obvious 

location externally, host specificity, and ease of identification by 

field observation, may make it a highly effective biological tag. 

Pennella sp. 

This is a new species of moderate-sized copepod on dolphin 

which is only known from Puerto Rico. 

Name - We are cooperating with Dr. William Hogans in describing 

and characterizing this new copepod. 

Diagnostic Characters - The female is small (less than 50 mm). 

It is similar to Pennella exocoeti with a cephalosome with 2 pad-like 

processes on each side. 

Records - We found 6-8 females on 13 of 17, and 1 each on 3 of 

13 dolphin off Parguera, Puerto Rico; and 5 females and 1 juvenile 

on a dolphin off Desecheo Island, Puerto Rico. 



Location in Host - Skin or fins embedded in muscle. 

Length - Female 22.5-43.0 mm. 

Host Specificity - This parasite only occurs on dolphin, and is a 

secondary parasite in Puerto Rico. 

Significance to Sport Fishing - See P. makaira. 

Brachiella thynni Cuvier 

This large copepod occurs worldwide on scombrids, especially wahoo. It 

is adapted to hide in the pit behind the pectoral fin base. 

Name - It was named for tunas, but may be found more often on wahoos. 

Diagnostic Characters - The cephalosome and neck are bent or curved 

anteriorly and are longer than the remainder of the body (excluding 

projections). Two attachment processes, approximately 1/2 as long as posterior 

processes, extend from the base of the neck. There are 2 pairs of unequal 

projections from the posterior end of the body; 1 pair (ventral processes) are 

about 2/3 the length of the trunk, the second pair (dorsal processes) are about 

2/3 as long as the first. The body is light brownish yellow. 

Records - We found 1-4 females in each of 19 wahoo, and 1 female on 1 of 40 

dolphin from various localities around Puerto Rico. It has been found in 

albacore, greater amberjack (USNM 39585) bigeye tuna, bluefin tuna, cero, king 

mackerel and yellowfin tuna in the western Atlantic, and these and other 

scombrids around the world. The Brachiella sp. Iversen and Yoshido (1957) 

reported from wahoo in the Pacific was probably this copepod.


Geographic Range - Worldwide. Our collections are 

the first from the Caribbean. 

Life History - The males are rarely found on fishes. 

In females, the right egg string is longer than the left, 

370-860 eggs occur in each egg string. 

Location in Host - The female seems to be adapted in 

body shape and size to the cavity behind the pectoral 

fins (axil) of big game fishes. This site specialization 

is rather unusual. It rarely occurs on other parts of the 

body, mouth or gills. Another species in this genus on 

Pacific Spanish mackerels, Scomberomorus spp., 

attaches in the gills and nasal lamellae. 

Length - Female 12.4-21.7 mm, egg strings 6.2-15.9 

mm; male 1.2-2.3 mm. 

Host Specificity - This copepod is a characteristic 

parasite of wahoo. Wahoo is the preferred host. It also 

occurs on a variety of other scombrids and a few nonscombrids, 

but it is no more than a secondary parasite 

of these hosts. Dolphin is a new host. 

Detection - Surprisingly, this large 

copepod can be missed in a cursory 

external examination. They are often completely hidden in the 

cavity behind the base of the pectoral fin (axil). 

Charopinopsis quaternia (Wilson) 

This characteristic parasite of dolphin occurs in light to 

moderate infections on the gills. 

Name - "Quaternia" means having 4 parts and refers to the 

posterior processes. Brachiella coryphaenae Pearse is a 

synonym. Wilson placed this species in a genus, Charopinus, 

which contains copepods of chimaeras, rays, sharks and skates, 

but Yamaguti (1963) erected a new genus. It has often been 

incorrectly spelled "C. quaternius". 

Diagnostic Characters - It is a tiny to small copepod with a 

cylindrical body. The cephalosome and neck are shorter than 

the trunk. Two elongate projections (posterior processes) extend 

off the posterior end of the trunk. Two attachment processes, 

shorter than the posterior processes, extend from the base of the 

neck, and terminate in bulbs that look like boxing gloves. 

Records - We found 3 females on 1 of 40 dolphin from various 

localities around Puerto Rico; and 1-2 females on 2 of 5 dolphin 

off Dauphin Island, Alabama, USA. Light infections occurred in dolphin from 

the Dry Tortugas, off the Florida Keys, USA (USNM 64009-11) and from the 

Straits of Florida, between Florida and the Bahamas; 11 females on 13 dolphin 

from Port Aransas, Texas, USA (USNM 92663, 92688); and on Spanish 

215

216 


mackerel from Louisiana, USA. It has also been reported from Hawaii in the 

Pacific, and south India in the Indian Ocean. 

Geographic Range - Worldwide (circumtropical/subtropical). But this is 

based on a few, widely spaced, records. Our collections are the first in the 

Caribbean and northern Gulf of Mexico. 

Life History - Burnett-Herkes (1974) suggested that this parasite did not reinfect 

older hosts, if true, this would explain why their numbers are reduced 

and they are finally lost from larger dolphin. What conditions favor younger 

fish becoming infected is not known. 

Ecology - Dolphin smaller than 38 mm are not parasitized (all SL), but young 

dolphin 38.1-39.9 cm are the most heavily parasitized (average 12.0 copepods), 

dolphin 40.0-79.9 cm are parasitized in lower numbers (average 1.7-5.9 

copepods), dolphins 80.0-96.5 cm are more lightly parasitized (average 0.5 

copepod), and older dolphin above 1 meter long do not have this parasite. 

Associations - Burnett-Herkes (1974) found that this copepod had a negative 

relationship with Caligus productus and Euryphorus nordmanni in the gills and 

mouth of 145 dolphin. He suggested that either both of these copepods were 

competing with Charopinopsis quaternia or there was a combination of 

competition and a lack of re-infestation by C. quaternia. 

Location in Host - Gill filaments. Reports from the gill chamber and 

operculum are of copepods dislodged after the death of the host. 

Length - Female 6.2-7.3 mm, egg strings 6.5-8.2 mm; male 0.4 mm. The 

females from the Gulf of Mexico are about 80% as large as those from Hawaii. 

Host Specificity - This is a primary parasite of dolphin, and is almost host 

specific. The single record on Spanish mackerel was probably contamination 

of these copepods from dolphin on sportfishing boats. This copepod must not 

occur very often on Spanish mackerel since it was not found by Cressey and 

Cressey (1980) in their extensive survey of scombrids. The record of 6 females 

and a male on a slender searobin is more difficult to explain. 

Significance to Sportfishing - This distinctive and common copepod might be 

of use as a biological tag, if its distribution is limited. 

Clavellisa scombri (Kutz) 

It occurs on mackerels worldwide, but in light infections on few fishes. 

Name - The name is appropriate for this genus-specific copepod. It was 

inadvertently omitted from the monograph by Yamaguti (1963) although he 

had collected, redescribed and refigured this species (Yamaguti 1939). 

Diagnostic Characters - The cephalosome and neck combined are more than 

twice as long as the trunk. The head has a sclerotized dorsal shield. The trunk 

is more or less circular and has a noticeable anal tubercle on the posterior end. 

No genital process is present. The specimen figured has the microscopic male 

attached on the neck. 

Records - One to 3 females occurred in chub mackerel from Alabama, USA; 

off Campeche, Mexico, tropical west Africa, Peru the Philippines and Japan;


Atlantic mackerel off Europe in the Atlantic and in the 

Mediterranean; and in a Pacific mackerel from Australia 

and Taiwan. 


Ecology - Offshore and pelagic. 



Host Specificity - This copepod is genus specific to 

mackerel (Scomber, all 3 species). It is widespread 

geographically, but occurs in light infections in low 

percentages of the available hosts. It was originally 

described from Atlantic mackerel in Europe, but in an 

extensive survey, examining 97 specimens of this host, 

Cressey and Cressey (1980) failed to find the copepod in 

this host. 

Damage to Host - It is a relatively large copepod 

capable of injuring the gills. Apparently few of these 

parasites occur on a host, thus damage is limited. 

Miscellaneous Copepods 

Genus Alebion - Alebion was one of the sons of Neptune, God of the seas, 

in Roman mythology. These copepods are strictly external parasites of inshore 

sharks. The single females reported, once each, on different big game fishes 

must have been the result of contamination from sharks landed on the same 

deck. The distinctive genus has a caligoid body, a cephalosome about 1/2 the 

length of the body without lunules, segment with fourth legs covered with a 

separate often ornate plate, fourth leg reduced to 1 segment, genital complex 

about 2/3 as wide as cephalosome and usually with posterior projections extending 

along side and beyond the end of the abdomen, abdomen in 2 segments and 

much shorter than genital complex, posterior projections 1/10 to 1/2 as long as 

abdomen and with setae as long as or longer than the projections, egg strings 

long. We found Alebion elegans Capart on great hammerhead, Sphyrna 

mokarran (Ruppell), in Puerto Rico, which is a new locality for the Caribbean. 

Alebion carchariae Krøyer - This copepod was found on a pompano 

dolphin from the Atlantic. It can be distinguished because the segment with the 

fourth legs is about 1/2 as wide as the genital complex and butterfly-shaped, the 

genital complex has posterior projections that extend back beyond the end of the 

abdomen and end in sharp points, the abdomen is shaped like a trident and has 

posterior projections about 1/2 as long as the abdomen and longer than the 

caudal rami. This parasite occurs around the world. The female is 7.6-9.0 mm 

and male 6.0 mm long. 

217

218 


Alebion glaber Wilson - A female (USNM 32840) was found on an 

Atlantic bonito landed off Massachusetts, USA. This copepod can be easily 

distinguished because the segment with the fourth legs is almost as wide as the 

genital complex, the genital complex is almost square and lacks any trace of the 

typical posterior projections, the abdomen is about as long but 1/2 as wide as 

the genital complex, and has a bulbous or round anterior abruptly tapering on 

the posterior 1/2. It is known only from the north Atlantic coast of the USA. 

The female is 10.0-12.0 mm, egg strings 15.0 mm, male 7.0-7.8 mm, chalimus 

on host 2.0 mm, and metanauplius 1.2 mm long. 

Alebion gracilis Wilson - A female (USNM 32727) was found on an 

Atlantic bonito landed off Massachusetts, USA. This copepod is similar to C. 

carchariae but can be distinguished by the lateral projections from the anterior 

end of the abdomen which are shorter than the caudal rami in this copepod, but 

longer in C. carchariae. It is known from the eastern Atlantic coast of the 

USA. Other western north Atlantic and the eastern Pacific records were of 

different species. The female is 9.0-9.6 mm, egg strings 9.0 mm; male 5.3-6.3 

mm; and nauplius 0.3 mm long. This copepod may be more rare than noted as 

it was frequently confused with C. carchariae even by Wilson. 

Anuretes heckelii (Kollar in Krøyer) - This parasite rarely and possibly 

accidentally occurs on big game fishes. Dojiri (1983) redescribed, refigured and 

established types for this species. It was formerly placed in Lepeophtheirus. 

Eirgos anurus Bere is a synonym. It has a caligoid body. The cephalosome is 

more than 1/2 of the total length of the copepod, and lacks lunules. The genital 

complex is ovoid, wider than long, almost as large as the cephalosome, and has 

posterior corners that extend beyond the caudal rami. The abdomen is 

relatively small and much wider than long. The egg strings are about as long as 

the body to longer. It was reported once in crevalle jack from Tuxpan, Mexico; 

and 4 females occurred in a Spanish mackerel from Texas, USA. We found 

females in a spadefish, Chaetodipterus faber (Broussonet), from La Parguera, 

Puerto Rico (USNM) and it has been reported from this host from several 

localities along the U.S. Gulf and Atlantic coasts, and Brazil. Our collection is 

the first in the West Indies. This copepod is known from the western Atlantic, 

and occurs in the gill chamber or mouth of its host. The egg strings contain 22- 

33 eggs each. The female is 2.1-2.9 mm long, egg strings 2.2-3.1 mm, 

immature female 2.4 mm long, male 1.2 mm. This copepod prefers spadefish, 

but has been found (once each) in 4 other hosts from 3 other families of fishes. 

Crevalle jack and Spanish mackerel may only be accidental hosts. 

Brachiella elegans Richiardi - This is either a rare parasite or accidental 

infection in the gills of greater amberjack. It may be a synonym of Brachiella 

thynni, but differs slightly in shape. Both of these copepods have been found 

on greater amberjack. This copepod has a cephalosome and neck that are 

aligned in the same axis as the rest of the body instead of being curved toward


the body and are much shorter than the remainder of the body (excluding the 

projections). Two attachment processes, approximately 1/2 as long as the posterior 

processes, extend from the base of the neck. There are 2 pairs of unequal 

projections from the posterior end of the body, 1 pair (ventral processes) are 

about 2/3 the length of the trunk, the second pair (dorsal processes) are less than 

1/2 as long as the first. The body is dark yellowish brown. Five females and 

1 male occurred in a greater amberjack from Massachusetts, USA (USNM 

39585). It has also found on other fishes in the Mediterranean. The egg strings 

contain 180-210 eggs each. It occurs in the gill chamber of its hosts. The 

female is 10.0-12.6 mm, egg strings 5.0-6.0 mm, and male 1.4 mm long. 

Caligus chelifer Wilson - The preferred hosts of this copepod are menhadens, 

but it has been recorded from a variety of hosts (anchovies, cutlassfish, 

drums, herrings, swordfish). Wilson (1905) described it from swordfish off 

Massachusetts, USA; Wilson (1932) suggested that this was a characteristic 

parasite of the swordfish and that this fish was the "primary" host; and Benz 

(pers. comm.) found 1 on a swordfish from the northwest Atlantic in 1978. The 

rarity of infections on swordfish suggests that these could be acidental infections. 

This female of this copepod was redescribed and the male described by Kabata 

(1972). It has a caligoid body with lunules moderately separated, and an abdomen 

with 2 segments. This copepod is known from the mid-Atlantic coast of 

the USA to Texas in the Gulf of Mexico. A record of this copepod in plankton 

off the west coast of Africa was apparently erroneous. The number of eggs in 

egg strings is 45-50. The female is 4.9-6.5 mm, and egg string 3.2 mm long. 

Caligus confusus Pillai - This copepod is aptly named as it was based on 

the confused specimens of several authors. That was not the end of the confusion, 

Yamaguti (1963) listed a record from dolphin in the eastern Pacific which 

was actually a rainbow runner. Confusion still exists in the Caligus spp. of 

Indo-Pacific jacks. 

Caligus curtus Müller - This copepod was reported on dolphin off Brazil, 

but this parasite is actually confined to the colder regions of the North Atlantic. 

The identity of the copepod in this report cannot be determined. 

Caligus haemulonis Krøyer - Reported on blue runner and cobia from 

Louisiana, USA, by Causey (1953a, 1955). The reports are in doubt since 

Cressey and Nutter (1987) found he misidentified most of these copepods; this 

copepod has not been found on big game fishes before or since; and grunts 

(Haemulon spp.) are, appropriately, the preferred hosts for this copepod with 

other fishes only serving as inadvertent or accidental hosts (Cressey 1991). At 

least one of these copepods has been found on a big game fish. Pearse (1952) 

also described a female copepod from the gills of a blue runner in Texas 

(USNM 92670) as a new species, C. validus, but it is a synonym. It has a 

caligoid body with narrowly separated lunules, cephalosome less than 1/2 of the 

219

220 


body length, roughly circular genital complex which is longer than the abdomen, 

abdomen in 1 segment with short caudal rami which are as wide as long. It is 

only known from the West Indies and Gulf of Mexico. The average length of 

the female is 3.6 mm. 

Caligus patulus Wilson - Two females were found on 2 of 145 dolphin in 

the Straits of Florida (Burnett-Herkes 1974). It was previously known from 

milkfish, Chanos sp., in Panama Bay, Panama, in the eastern Pacific. Dolphin 

may be an accidental host for this copepod. It has a caligoid body with widely 

separated lunules, a genital complex that is wider than long, an abdomen that is 

as wide as long, much shorter than the genital complex and is almost surrounded 

by posterior projections of the genital complex. The female is 6.1 mm long. 

Caligus praetextus Bere - It was reported on crevalle jack from Louisiana, 

USA, by Causey (1953a). However, Cressey and Nutter (1987) found he 

misidentified most of these copepods; and Bere (1936) and Cressey (1991) did 

not find this copepod on crevalle jack. It has a caligoid body with widely 

separated lunules, cephalosome about 1/2 of the body length, roughly 

rectangular genital complex which is longer than the abdomen, abdomen 1segmented 

with inwardly directed caudal rami. Microscopically the sternal furca 

(wish-bone shaped structure in the middle of the underside of the cephalosome) 

is square. It is only known from the northern Gulf of Mexico from the middle 

of the west coast of Florida to Texas. The limited distribution and distinctive 

appearance would make this copepod a useful biological tag, if it parasitized any 

wide-ranging big game fishes. The female is 2.2 mm long. It demonstrates no 

host specificity in abundance or frequency of occurrence on any host species. 

Cecrops latreillii Leach - A female was reported only once from a tuna, 

Thunnus sp., in the Atlantic and the collector suggested that this might have 

been a mistake in labeling or an accidental transference to this host. The body 

is rectangular and more than twice as long as wide with all segments about the 

same width. The cephalosome is almost rectangular (longer than wide) with 

broad posterior lobes, a notch in each side and no lunules. An oval plate (wider 

than long) covers the fourth thoracic segment and overlaps the genital complex. 

Long posterior plates of the genital complex cover the abdomen and look like 

the back of a beetle. Egg strings are irregularly coiled and hidden under dorsal 

plates of genital complex. It occurs worldwide in temperate and cooler waters. 

This copepod breeds from May to October. The ocean sunfish host provides the 

opportunity for this parasite to infect or be transferred to offshore big game 

fishes. It occurs on the gills of the host. Female is 25-30 mm and male 10-17 

mm long. Ocean sunfish is the preferred host, but it occurs on other fishes. 

Lernaeolophus hemiramphi (Krøyer) - A single female of this parasite 

of Caribbean halfbeaks was recorded once from cobia in the Gulf of Mexico. 

This apparently accidental infection could have been caused by a prey-predator


transfer of immature copepods. We examined numerous specimens of halfbeaks 

off Puerto Rico and named a new species of isopod from these hosts, but did not 

find this copepod. It is a large copepod with the anterior portions embedded in 

the tissues and the posterior portion with projections shaped like peacock 

feathers (abdominal process branched terminally). It has a bulbous trunk with 

spiral-shaped egg strings. The female is 16.0 mm long. 

Lernaeolophus sultanus (Nordmann) - One female occurred on a cobia 

from Mississippi, USA; a 13.0 mm female on a swordfish off Ceylon; 1 each 

on a wahoo in the Pacific and on a black jack from an unknown locality. 

Despite these rare records in big game fishes, this copepod almost always occurs 

on inshore, benthic fishes. It is a large tissue-embedded copepod with dendritic 

projections from cephalosome and dense mat of thick, 10 or more branched 

caudal projections along each side of a projection from the posterior end of the 

body. The neck and cephalosome are shorter than the body. We only found 

this copepod on coral reef fishes in Puerto Rico. The copepods we collected 

attached in the roof of the mouth of their hosts and often penetrated through the 

head to caused a large open lesion on the top of the head. The single female in 

a swordfish attached in the body near the anus of the host; and the female in a 

cobia was embedded in the body just posterior to the last dorsal fin ray. These 

are unusual positions for this copepod, and may indicate that the infections 

were accidental. The female is 12.0-15.0 mm long. The caudal projections of 

this copepod are often covered with algae and protozoans, the one from a cobia 

had a striped goose barnacle attached. One we collected was densely covered 

in algae. The projections become more branched in older and larger copepods. 

The head wound and the parts of the copepod that projects into the wound are 

often covered with dense tufts of algae. During an underwater monitoring 

project, we observed a coral reef fish with a large bump on top of its head. 

Three days later this lesion erupted into an open, crater-shaped wound caused 

by this copepod. Whether this copepod always produces a head lesion, and if 

hosts survive this severe injury, is not known. 

Lernanthropus hiatus Pearse - One male 1.7 mm long occurred on the gill 

filaments of an albacore from Bimini, Bahamas (USNM 88560) (Pearse 1951). 

This species was based on a single male specimen and is thus questionable. It 

differs from the known males in the genus by having a wide space between the 

antennae and the mouth tube. Cressey and Cressey (1980) did not find any 

members of this genus of copepods on tunas. 

Lernanthropus micropterygis Richiardi - This copepod was described 

from greater amberjack in the Mediterranean and later found on another jack in 

the Red Sea; and 1 female each on a greater amberjack in the Pacific off Mexico 

and an unidentified jack, Seriola sp., possibly alamaco jack, off Costa Rica. We 

are skeptical of the eastern Pacific identification as this copepod has not been 

found in the Atlantic. As it stands, this parasite is genus specific to 3 species 

221

222 


of jacks (Seriola spp.) and has a peculiarly disjunct geographic distribution. It 

was named for the host, but unfortunately greater amberjack was called 

Micropteryx dumerili in 1885 when it was named. 

Pandarus sinuatus Say - Causey (1953b) possibly found as few as 1 

(number unstated) females on the body of a crevalle jack from Port Aransas, 

Texas. He complained in his paper about sport fishermen "abusing" fishes 

(gaffing, dragging across docks, weighing, hanging, posing and photographing) 

and knocking off or contaminating copepods from other hosts before he was 

permitted to examine them, and also about dislodged copepods clinging to 

improper hosts including human fingers. We believe this characteristic parasite 

of sharks stuck to this jack through such contamination. Causey was impressed 

that these dark brown (almost black) females with long brown egg strings could 

be identified 20 feet away. The average length of the female is 6.5 mm. This 

copepod only parasitizes inshore sharks. Pandarus satyrus Dana was probably 

similarly contaminated once onto a Pacific marlin. Eldridge and Wares (1974) 

found large numbers of caligoid copepods irritating the skin and causing 

redness in eastern Pacific billfishes. Some of these were identified as Pandarus 

sp. Possibly some accidental infections do occur on big game fishes. 

Thysanote longimana Wilson - This shaggy looking copepod is an 

external parasite of larger snappers and probably only accidentally occurred in 

a bar jack. The name means long arms and refers to the second maxillae. The 

cephalothorax is round and attached to a vase-shaped body which lacks 

obvious segments. A pair of attachment arms and 4 dendritic branches extend 

from the neck, the arms joining at their terminal ends. Four more branches 

extend from the trunk just anterior of the posterior end, 2 are simple projections 

and extend posteriorly from between the egg strings. It is yellowish white in 

color. Two females occurred on a bar jack off Carabiñero Beach, Mona Island, 

Puerto Rico (USNM). Our collections appear to be the first reported since this 

parasite was collected more than 85 years ago. It was originally described in a 

misidentified snapper from Jamaica, and we found it in 3 species of snappers 

from Puerto Rico. This copepod is only known from the Caribbean. It occurs 

in inshore, benthic habitats. This parasite attaches on the outside of the throat 

of its hosts. The female is 8.0-9.0 mm, egg strings 4.0 mm; and male 1.3-1.4 

mm long. It is family specific to snappers (Lutjanidae). The 1 record in a jack 

could be accidental. Bar jack is a new host record for this copepod. This 

copepod is easy to see. We speared a bar jack at Mona Island because we 

noticed these copepods on its throat. We have seen many thousands of bar 

jacks before and since this collection and have not noticed this copepod. 

Thysanote ramosa (Richardi) - This copepod was described, as Brachiella 

ramosa, from Mediterranean swordfish. Silas and Ummerkutty (1962) and a 

few uncritical checklists suggest it also occurred off Massachusetts, USA, but 

this appears to be an error.

BRANCHIURA (FISH LICE) 

Fish lice, argulids or branchiurans form a small subclass of crustaceans. 

They can be very harmful to fishes, especially those in hatchery or culture 

facilities. Fish lice can infect the eyes of humans and bite careless handlers of 

live fishes. Approximately 150 species have been described in 5 genera, more 

than 120 in the genus Argulus. They are relatively large parasites varying from 

a few to 20 mm long. 

The body is flattened (strongly depressed) and has a large, expanded head 

(carapace), thorax and an abdomen. The thorax has 4 ill-defined segments and 

the abdomen is completely fused. The appendages of the carapace are 

modified into mouthparts and suckers, and those of the thorax are 4 pairs of 

unmodified legs. Most have large suckers on the under side (dorsal surface) of 

the front of the carapace, something like the caligoid lunules only relatively 

much larger, in a different position and origin. Many have a long and vicious 

sting (stylet) in front of the mouth and between the antennae. The abdomen has 

no appendages, and terminates in a bifurcate tail. 

They mate while free swimming off the host. Eggs are held in the body of 

the female. Females leave the host to deposit eggs in clusters attached to the 

substrate. Argulus spp. hatch as nauplius larvae, but members of the other 

genera hatch as juveniles from eggs in 15-55 days and develop directly into 

adults. Swimming juveniles must find a host in 2-3 days and once attached 

develop into adults in 30-35 days. Sexes are separate. 

They attach on the body, fins, gills and mouth of fishes and sometimes on 

frogs and tadpoles. They are obligate parasites, feeding on blood, but adults are 

capable of changing hosts and spending prolonged periods off any host. They 

may prefer some fishes, but are usually not host specific. Heavy infections can 

kill fishes. A combination of moderate infections, Anilocra acuta, a bacterial 

infection and polluted conditions have caused mortalities in wild inshore fishes 

in the Gulf of Mexico. Fish lice directly transmit viral and bacteria diseases. 

They have introduced microbial diseases into culture facilities and caused 

epizootics. 

They are the only crustacean fish-parasites known to infect humans. 

Others may bite or attack humans, but only Argulus spp. penetrate, survive in, 

and cause diseases in humans. Hargis (1958) reported the first case of human 

argulosis in a child infected by Argulus laticauda Smith while swimming in salt 

water off the Atlantic coast of the USA. We recently interviewed an aquaculture 

specialist from South America who became infected with Argulus sp. during her 

attempt to control losses of cultured tilapia caused by very heavy infections of 

this parasite in fresh water. One of these argulids was splashed into her face 

and lodged between her eye and the orbit. It caused severe irritation and minor 

tissue damage for 24 hours before it was discovered and removed. She was able 

to sleep with this horrid little beast in her eye. All fish lice should be treated 

with caution. 

223

224 


These parasites are more important in fresh, brackish and inshore marine 

waters. None occur in the open ocean realm of big game fishes probably 

because this habitat does not provide substrates for their eggs. 


Subclass Branchiura - fish lice Page 

Order Argulidea 

Family Argulidae 

Argulus bicolor ................................................................................ 224 

Argulus bicolor Bere 

This is a rare and possibly accidental parasite of 

great barracuda in inshore waters. 

Name - The name refers to the predominant green 

and rust colors of the body. 

Diagnostic Characters - The carapace is elongate and 

longer than wide, and 2 suckers occupy almost its 

entire width of the ventral surface (not shown in the 

illustration of the dorsal surface). Lobes of the 

carapace do not extend to the abdomen. The 

posterior corners of the last thoracic segment form 

rounded lobes. The abdomen is relatively large 

(about 1/3 of the total length). 

Records - This louse occurred on a great barracuda 

and another fish from Biscayne Bay, Miami, Florida, 

USA; and on other fishes from the west coast of 

Florida (USNM 69863). 



Length - Female 4.0 mm; male 3.2 mm. 

Host Specificity - Needlefish (Belonidae) appear to be the preferred hosts of 

this parasite. The occurrence on great barracuda may have been accidental. 

CIRRIPEDIA (BARNACLES) 

Barnacles form a subclass in the crustaceans. The common name 

"barnacle" is from the Middle English "bernak" (a goose); and/or the French 

"bernicle." Apparently for the shape of "goose" barnacles. The barnacle goose 

in Europe was reputed by the ancient Greeks to spawn spontaneously from goose 

barnacles. They are famous for incrusting the bottom of your boat and other 

marine structures. Cleaning barnacles from structures, antifouling methods and 

the transport of exotic organisms involve serious economic and environmental 

problems. Barnacles also associate with or parasitize a variety of commercially 

important marine organisms. A great variety of organisms eat these animals,


including "barnacle eaters" (filefishes, Alutera spp.). Barnacles were eaten by 

native Americans, are a delicacy in Europe, and the world's largest barnacle 

supports an important fishery in Chile. 

More than 1000 living species have been described, and most are free 

living. Barnacles vary in size from the minute acrothoracicans, which burrow 

into the calcareous skeletons of corals and sea shells, to parasitic 

rhizocephalans anastomosing throughout the body of large crabs. The 

crustacean body form is greatly modified in all barnacles, but is drastically 

altered in some parasitic species. Most barnacles have a heavy calcarious shell, 

which is unique among crustaceans, composed of several to many parts 

imbedded in a soft mantle which surrounds the animal. Free-living forms are 

either attached by long stalks (goose) or directly to the base (dorsum) of their 

shells (acorn or volcano). Free-living barnacles filter feed by sweeping slender, 

jointed appendages (cirri) through the water. Parasitic forms live on or in 

various marine crabs and other crustaceans, echinoderms (sea stars, etc.) and 

soft corals; and most have lost the shell, appendages and body segmentation of 

free-living barnacles. Some barnacles are specialized associates of particular 

crabs, sea shells, other invertebrates, turtles and whales; while others attach to a 

variety of substrates or organisms. 

The larvae are planktonic. Most barnacles have both female and male 

sexual organs (hermaphroditic), but some groups, particularly parasitic ones, 

have separate sexes. They attach externally, burrow inside skeleton or 

endoparasitically in a variety of hosts. Barnacles vary from free-living through 

various levels of association to permanent parasites. Parasitic barnacles feed on 

the tissues of their host. The shells of barnacles fossilize well and have left a 

good fossil record which shows a comparatively recent radiation (expansion in 

number of species and importance) in the middle of the Carboniferous Period 

(approximately 230 million years ago). Barnacles may have evolved from 

wholly parasitic ancestors as suggested by the many parasitic and hostassociated 

species remaining today and the primitive parasitic ascothoracicans. 

However, closely related subclasses have free-living ancestors, which tends to 

argue against a parasitic origin of barnacles. Fertilization occurs in the mantle 

cavity or at the base of the oviduct; eggs (ova) generally develop to the firststage 

nauplius within the mantle cavity, and are expelled by pumping 

movements of the body. The triangularly-shaped nauplius larva has corners 

tipped with prominent spines. It molts through 6 stages in the plankton, and 

then seeks a host or substrate. The thin, transparent bivalve-shelled, ostracodlike 

cyprid larvae rapidly swims and crawls around the substrate, selects a site 

for attachment, cements itself in place, and metamorphoses into a barnacle. 

One free-living species has been reported associated with big game fishes, and 

another may be confirmed. 

References - Anderson (1993). 

225

226 



Subclass Cirripedia - barnacles Page 

Order Lepadomorpha - goose or pedunculate barnacles 

Family Lepadidae 

Conchoderma auritum ..................................................................... 226 

Conchoderma virgatum ................................................................... 227 

Conchoderma auritum (Linnaeus) - rabbit-ear barnacle 

This large pelagic, rabbit-ear barnacle attaches to 

whales or ship hulls, occasionally sharks, and may be found 

on big game fishes. 

Name - It is called "rabbit-ear" barnacle, or "rabbit-ear 

whale" barnacle, for the projecting "ears" and frequent 

association with whales. We expect that it occurs on big 

game fishes, but has been confused with striped goose 

barnacles. Great variations in the shape of this barnacle 

have inspired the unfortunate description of many 

unjustified species (synonyms). 

Diagnostic Characters - This is a moderate-size to large 

goose barnacle with 2 elongate, ear-shaped projections on 

the anterior end, and usually a base stem (peduncle) that is 

distinct from the body (capitulum). The ears are the only 

consistent characteristic to distinguish this barnacle from 

striped goose barnacles. The ears only occur in adults. 

Immatures often cannot be physically identified. 

Records - This barnacle has been rarely reported from 

sharks and whales in the Gulf of Mexico; whales, sharks and bony fishes in the 

Atlantic, Indo-Pacific and Arctic. It occasionally attaches on ship hulls and 

buoys. 


Ecology - Oceanic, pelagic. 

Location in Host - It attaches to barnacles on the skin of whales, but does not 

attach directly on whale skin. This barnacle can attach to baleen, palate, penis 

and teeth of whales; or any external, exposed hard parts such as bone or shell 

of marine mammals, sea turtles or fishes. 

Length - 10.0-30.0 mm. 

Host Specificity - It is usually found on acorn barnacles, Coronula diadema 

(Linnaeus), attached to whales, but sometimes on striped goose barnacles on 

sea turtles and whales. This barnacle has been reported to only occur on slowmoving 

fishes. This may explain its absence on big game fishes, but its great 

variety of incidental hosts, suggests it may infect almost any fish. It appears to 

prefer living hosts more than inanimate floating objects. 

Preparation for Study - Records on big game fishes should be confirmed by 

preserving specimens of these barnacles in alcohol (151 proof rum, or rubbing 

alcohol) or 10% formalin. If these chemicals are not immediately available,


specimens may be sealed in a plastic bag and held on ice or in a refrigerator for 

no more than 2-4 days before preservation. 

Conchoderma virgatum (Spengler) - striped goose barnacle 

This goose barnacle spectacularly attaches to crustacean 

parasites on fishes, including big game fishes. 

Name - It is called "striped goose barnacle" for its 

markings. The great variations in the shape of this bar- 

nacle have inspired the description of many unwarranted 

species (synonyms). 

Diagnostic Characters - This small to large goose bar- 

nacle lacks the elongate ears of C. auritum. The base stem 

(peduncle) and body (capitulum) are blended together 

without forming a distinct separation. 

Records - We found 1 on 1 of 6 partially embedded copepods, 

Pennella makaira, attached on an Atlantic blue marlin 

off Arecibo, Puerto Rico. It occurred on a partially 

embedded copepod, Lernaeolophus sultanus, on a cobia off 

Mississippi, USA; and on Pennella filosa on swordfish off 

the Atlantic coast of the USA. This barnacle was reported 

on Pennella instructa on Indo-Pacific sailfish, swordfish, and other billfishes 

from the Indian Ocean, but other studies suggest that it does not occur on this 

copepod. It has also been reported from a variety of hosts and substrates 

including floating objects, boat hulls, sea turtles, sea snakes, whales and fishes. 


Ecology - This barnacle is odd because it only grows at shallow, mid-water 

depths. Most other barnacles are quite happy to grow at the surface of the 

ocean. Offshore, big game fishes make excellent platforms for these associates, 

but they require some solid surface unhindered by scales and fish mucus to 

attach. We have found these barnacles attached to parasitic copepods and 

isopods on host fishes, exposed bone from host wounds, and on fish tags. 

This barnacle may attach to offshore sport fishes while they are near shore. 

Leatherback turtles, Dermochelys coriacea (Linnaeus), moving from the Atlantic 

to Caribbean islands to spawn, arrive without any striped goose barnacles, but 

soon become a substrate for this associate. It is also very common on Pennella 

spp. partially embedded in ocean sunfishes and flyingfishes that never venture 

inshore. 

Williams (1978) reported the first record of striped goose barnacle 

attached to a fish-parasitic copepod not permanently embedded in the host and 

Benz (1984) reported 4 more cases. Causey (1960) found a similar instance of 

what was possibly this barnacle (Conchoderma sp.) on Gloiopotes huttoni on the 

Indo-Pacific sailfish off Mazatlan, Mexico. 

Associations - This barnacle attaches to parasitic copepods and isopods on 

fishes. It competes for space with other barnacles on the shells of sea turtles. 

227

228 


Location in Host - It usually attaches on the external surfaces or crustacean 

parasites on hosts, but we found 1 attached to a copepod in the mouth of a fish 

(Williams and Williams 1986). 

Length - 7.2 to 25.0 mm. 

Host Specificity - This barnacle has no host specificity, but occurs on any 

crustacean parasite or exposed hard part on a host. It is sometimes common on 

leatherback turtles; copepods, Pennella spp., partially embedded in offshore 

fishes; and isopods, Nerocila sp., on inshore fishes. Pennella makaira is a 

new copepod host, and Atlantic blue marlin a new fish host. 

Damage to Host - Direct attachment to exposed bone or shell of the host is 

probably benign and may even help to cover or close wounds. These barnacles 

are reputed to increase the water resistance of copepods or isopods by their 

attachments. This force should increase the tissue damage of embedded 

crustacean parasites, but may pull non-permanently attached copepods and 

isopods from their host. 

ISOPODA (ISOPODS) 

Isopods are an order in the crustaceans. The name refers to all legs (pods) 

being approximately similar in size and shape (iso). Isopods kill, stunt and 

damage commercially important fishes. Approximately 9.4% of the chub 

mackerel along the Peruvian coast are parasitized by Meinertia gaudichaudii 

(Milne-Edwards) causing a 15% loss in body weight and costing Peruvian 

fishermen approximately 1.3 billion kilograms of fish annually. This is 

unfortunately not an isolated case. A few fish-parasitic isopods actively swim 

after and bite humans, sometimes alarmingly in mass attacks, but bites are more 

likely to occur when handling infected fishes. Free living isopods are reported 

to clean Saprolegnia spp. (fungus) from fishes. Locally, Anilocra spp. are 

dried and used to make a tea to treat colds. New England fishermen use "salve 

bugs" (Aega spp.) for medicinal purposes. Isopods are eaten by a variety of 

animals. Giant isopods, Bathynomus spp., are fished commercially for human 

food in Japan and Mexico and Hawaiians eat a smaller species. The presence 

of parasitic isopods on marine tropical fishes allegedly indicates that they do 

not contain high amounts of ciguatera (fish poisoning) toxins. This is not 

proven, but highly interesting, particularly since large barracuda and jacks are 

commonly implicated in ciguatera poisoning, and often have attached isopods. 

Approximately 4000 species of isopods have been described, and more than 

450 species are known to associate with fishes. They vary from 0.5-440 mm 

in length. The world's largest species, Bathynomus giganteus Milne-Edwards, 

is found off Puerto Rico and beyond. The head is fused with first thoracic 

segment (cephalothorax), and they have a 7-segmented thorax and 6-segmented 

abdomen (often fused into 2-5). One pair of thoracic appendages is modified 

into mouthparts, and 7 pairs are unmodified. The abdomen has 6 pairs of 

appendages, and ends in a terminal, often shield-shaped segment called the 

pleotelson. Eggs, larval forms and juveniles develop either in a brood pouch


beneath, or in pouches in the abdomen of 

the female. Most isopods possess free 

swimming juveniles that develop into 

adults, but gnathiid juveniles parasitize 

the gills and skin of fishes and are free 

living as adults. Sexes are separate in 

most isopods, while others begin life as 

males and later become females 

(protandrous hermaphrodites). They are 

common in most environments, including 

dry land. They parasitize fishes, crabs, 

shrimp and other isopods. Fishassociated 

isopods vary from accidental 

(cirolanids), temporary or casual 

(corallanids and aegids) to permanent 

(cymothoids) parasites. They attach in a 

variety of locations including the skin, gills, 

inside the mouth, on the fins and some even burrow under the skin to form a 

cyst in the flanks of fish. A broad range of food habits occur. The fishassociated 

forms feed on blood or ooze from wounds. The wounds isopods 

cause may provide entry points for microbial diseases. Isopods can be preserved 

and stored in 70% ethanol (151 proof rum will do) or 40% isopropanol (rubbing 

alcohol). 

Most isopods associated with fishes occur inshore, except those on flyingfishes 

and pelagic needlefishes. The majority of our records from big game 

fishes were accidental infections while these hosts were inshore, or transfers 

from flyingfishes or other prey fishes. Only Cymothoa oestrum on jacks and 

great barracuda and Livoneca redmanii on Spanish mackerels occur commonly 

on big game fishes. Even these isopods first attach when their hosts are inshore. 

Free living isopods are found in the stomachs of big game fishes often 

enough to be listed in some food item studies. Large parasitic cymothoids 

[Glossobius auritus Bovallius, Glossobius impressus (Say), Nerocila excisa 

(Richardson)] have been reported from the stomachs of dolphin in the Pacific, 

and Glossobius impressus has been reported from the stomachs of yellowfin 

tuna in the Caribbean and Pacific. These isopods parasitized flyingfishes that 

were eaten by these big game fishes. At least 2 species of isopods have been 

transferred from prey species to big game fishes (see Discussion). 

Popular references - "Isopods" (Williams and Bunkley-Williams 1997), 

"Marine isopod crustaceans of the Caribbean" (Kensley and Schotte 1989). 


Class Malacostraca Page 

Order Isopoda - isopods 

Family Gnathiidae 

Gnathia spp.* ................................................................................ 230 

229

230 


Family Corallanidae 

Excorollana tricornis ..................................................................... 231 

Family Aegidae 

Rocinela signata ........................................................................... 232 

Family Cymothoidae 

Anilocra acuta ............................................................................... 233 

Cymothoa oestrum ......................................................................... 234 

Livoneca ovalis .............................................................................. 236 

Livoneca redmanii ......................................................................... 237 

Nerocila lanceolata ....................................................................... 238 

_______ 

*Larval form 

Gnathia spp. 

These tiny crustaceans swarm over Caribbean reef 

fishes at night and keep the cleaner fishes and shrimp 

busy picking them off during the day. They are 

rarely reported from big game fishes. 

Name - Benz (1994) called them "sea gnats". The 

juveniles and immature females that occur on fishes are 

extremely difficult to identify to species since most 

species characters are based on the adult, free-living 

male. 

Diagnostic Characters - These minuscule to tiny 

isopods have only 5 pairs of legs (small juveniles of 

other isopods have 6-7 pairs of legs). The head has a 

constricted "neck" instead of being fused into the 

remainder of the body. 

Records - One each occurred in 3 of 45 great 

barracuda and 2 on 1 bar jack from various localities 

around Puerto Rico. It was also reported from a blue runner and a yellow jack 

from Jamaica. 

Geographic Range - Members of this genus occur around the world. 

Life History - Eggs develop into juveniles in cavities in the body of the female 

brood pouch. Juveniles attach, probably at night, to the gill filaments, body or 

fins of fishes and suck blood, their flexible bodies swelling as they fill. They 

eventually drop off the fish and mature into free-living males or females. 

Males develop large heads, some with massive mandibles (jaws), occupy 

benthic habitats (commonly coral reefs) and have harems of many females. 

These isopods were not thought to change hosts or to take more than 1 

blood meal from a host. However, Haemogregarina bigemina (a protozoan 

parasite of fish blood cells) has been shown to complete its life cycle in the body 

of a Gnathia sp., which then transmits the protozoan between fish hosts. For


this to occur, a gnathiid must take a blood meal from a protozoan-infected host, 

drop off long enough for the protozoan to develop, and then contaminate a 

second host by taking another blood meal. Thus, the host relationships of 

gnathiids may be more complex and long lived than once thought. 

Ecology - This is not a parasite of the open ocean. Gnathia spp. are largely 

restricted to inshore or deep sea fishes more closely associated with substrates 

where the adult gnathiids can develop. 

Associations - One Gnathia sp. was found attached to an Excorallana tricornis 

in the mouth of a great barracuda from Puerto Rico. 

Location in Host - Gill filaments and skin. 


Host Specificity - These isopods are assumed to have no host specificity, but 

the biology of most species is unknown. 

Damage to Host - Usually a few isopods do little damage. Heavy infections 

cause tissue damage and kill hosts in confined or culture situations. 

Detection - These small isopods can barely be seen on fishes, and look like tiny 

red spots. They are more easily seen and identified with the use of a dissection 

microscope. 

Significance to Sport Fishing - They appear to be too rare to cause problems 

in big game fishes, but transmission of H. bigmina may injure young big game 


Excorallana tricornis (Hansen) 

These small isopod have horn-like processes 

on the male's head. They casually associate with 

many species of fishes, but will readily "abandon 

ship". 

Name - The name "tricornis" refers to the 3 

horns on the head of the male. Delaney (1984) 

revised this genus and designated E. tricornis as 

the type species. 

Diagnostic Characters - There are 3 large horns 

on the head of the male (more obvious in profile, 

than when viewed from above, as in our figure). 

The body is elongate. There are small spines on 

all legs rather than large hooks. 

Records - We found 2-20 in 6 of 45 great 

barracuda from various localities around Puerto 

Rico. It has been found in blue runner, crevalle jack and great barracuda from 

Jamaica. This isopod is also known from the northern coast of Yucatan 

(Mexico), the Gulf of Mexico, Panama and we recently reported it from 

Colombia (Williams, Bunkley-Williams and Sanner 1994) The eastern Pacific 

"sister species" occurs in fishes including a jack. 

Geographic Range - Caribbean and Gulf of Mexico. 

231

232 


Life History - These isopods probably breed throughout the year, but increased 

numbers of planktonic juveniles occur in the spring and fall. Gravid females 

are seldom captured. They probably do not feed and brood eggs through 

juveniles in seclusion in the substrate in shallow waters. Juveniles leave the 

female and live in the plankton for an unknown period of time, then settle onto 

the substrate and develop into adults. 

Ecology - It is an inshore species that only occasionally infects big game fishes 

when they are near shore. 

Location in Host - Gill chamber, nares or external folds in the throat (gular) 

and gill cover. Occasionally in the mouth, skin or fins. 

Length - Females 4.0-9.6, males 3.9-9.9, juveniles 1.0-3.0 mm. 

Host Specificity - None. 

Damage to Host - Few isopods occur on a host. Blood feeding might be 

significant on small hosts, but no tissue damage has been reported. 

Detection - These isopods are easily seen. They readily detach from the host 

and fall onto the deck, so fishes should be immediately examined after capture. 

Also, look for loose isopods on the deck and under the catch. 

Preparation for Study - Quickly seal gills, heads, etc. in plastic bags. Later, 

the sediment from these bags can be decanted and examined for the presence of 

this isopod. 

Rocinela signata Schioedte and Meinert 

The monogram isopod is the most common and 

ubiquitous medium- to large-sized isopod found in the 

gill chambers of Caribbean fishes. It is known to 

severely bite swimmers and divers, sometimes in 

mass attacks (Garzon-Ferreira 1990). 

Name - Called the "monogram isopod" for the mark 

on the tail (pleotelson). 

Diagnostic Characters - It has an M- or W-shaped 

mark on the tail and is a moderate-sized, flattened 

isopod with an oval body outline. Large hooks occur 

on the first 3 pairs of legs, but only straight segments 

on last 4 pairs. 

Records - We found 1 in 1 of 35 cero, 2 in 1 of 45 

great barracuda, 1 in 1 of 15 king mackerel from 

various localities around Puerto Rico; and 1 in a cero 

from Jamaica. It has also been reported from albacore 

from the Atlantic and in other fishes from Florida, 

USA and Brazil. 

Geographic Range - Tropical western Atlantic. 

Life History - Gravid females do not occur on fishes. They probably consume 

a large blood meal before dropping off a host and do not feed again until the 

offspring emerge as juveniles. Then the female molts and loses the flaps 

forming the brood pouch and feeds again and continues to repeat the


reproductive process. Juveniles have a planktonic stage, and then are found free 

living in or around the substrate or temporarily parasitic on fishes. Isopods of 

all sizes have been collected from fishes. They remain good swimmers 

throughout their lives, except for gravid females. 

Ecology - This is an inshore species. The few isopod reported on offshore 

fishes may have been carried from the shallows to pelagic areas on these hosts. 

They have been collected from depths of 55 m in the Gulf of Mexico, 60-93 m 

off Mexico, and we have seen it on fishes caught in deep water traps off 

Colombia (Williams and Bunkley-Williams 1994). 

Location in Host - Gill chamber, rarely externally. 


Host Specificity - No fish host preference is apparent and since it feeds off 

humans, we suspect that it also attacks sea turtles and mammals, as well. 

Damage to Host - This isopod does not associate for sufficient periods of time 

on the host to cause tissue damage. Usually they occur in too low of numbers 

to cause much injury. 

Detection - The adults are easily seen, but juveniles must be found with a 

dissection microscope. 

Harm to Humans - These isopods are known to bite swimmers and scuba 

divers with mass attacks driving divers out of the water off Colombia (Garzon- 

Ferreira 1990). The bites are painful and can be bloody. One of our aquanauts 

in a saturation NOAA Habitat mission made an emergency stop inside an 

underwater talking bubble to remove a monogram isopod from under his ear. 

When he did so, blood streamed down his neck. This is the only isopod known 

to treat humans as routine prey! 

Anilocra acuta Richardson 

This external rider of gars and other fresh- and brackish 

water fishes, can kill its host. It has been reported from big 

game fishes only once, but in spectacular fashion. 

Name - Acuta (=acute) for its pointed head. 

Diagnostic Characters - The head is distinctly pointed. It 

is a moderate-sized flattened isopod, with the body roughly 

triangular in outline. 

Records - We found this isopod far offshore in the throat of 

a king mackerel off North Carolina. This indicated acciden- 

tal transfer of an adult isopod from a prey to predator host 

and survival of an external isopod in the throat of an new 

host (Williams and Bunkley-Williams 1994). 

Geographic Range - Atlantic through Gulf coast of the 

USA from New York to Texas, and possibly into the Gulf 

coast of Mexico (Williams and Bunkley-Williams in press). 

Ecology - Limited to coastal freshwater and brackish water areas, although it 

survived in full-strength seawater in the king mackerel. Members of the genus 

233

234 


Anilocra are essentially tropical and marine. This is the only exclusively 

temperate member and freshwater species. 

Associations - A combination of this isopod, bacteria, Aeromonas hydrophilacomplex, 

and fish lice, Argulus lepidostei Kellicott, have caused disease and 

mortalities in coastal gars (Lepisosteidae) in the Gulf of Mexico. 

Location in Host - Skin or fins. The occurrence in the throat was highly 

unusual and an example of site transfer. 

Length - Female 24.5-34.2 mm, male 20.4 mm. 

Host Specificity - Usually occurs on gars, but occasionally on other coastal 

fresh and brackish water fishes. The preference for gars appears to imply 

limited host specificity. 

Damage to Host - Although known to cause lethal diseases (see Associations 

above), it more often simply stunts fishes as do all fish-parasitic isopods. 

Cymothoa oestrum (Linnaeus) 

This "jack-choking" isopod stunts wild adults and 

kills aquarium fishes. Caribbean folklore suggests 

that jacks and barracuda with this isopod do not have 

ciguatera fish poison. This fable requires testing. 

Name - They are also called "tongue-biters" for its 

habit of occupying most of the oral cavity of jacks. 

Diagnostic Characters - It is a large, white or 

cream-colored isopod with a rectangular body outline 

and has "shoulder-pad" projections beside the head. 

Records - We found a female-male pair in each of 2 

of 19 bar jack, 1 of 2 blue runner, 7 of 20 crevalle 

jack, 4 of 14 great barracuda, and 9 of 27 horse-eye 

jack from various localities around Puerto Rico; 1 

blue runner each from Bermuda and West Palm 

Beach, Florida, USA; 1 horse-eye jack each from 

Curaçao and St. Lucia; and a crevalle jack from 

Trinidad. It has also been reported in bar jack from 

the Florida Keys, USA, and Carrie Bow Cay Belize; crevalle jack from 

Venezuela; horse-eye jack from the Bahamas and Barbados; and jacks (Caranx 

sp.) from Curaçao and Jamaica. 


Life History - Juveniles are released from the brood pouch, swim in the 

plankton, feed off a variety of small fishes as transfer hosts, finally enter the 

mouth of the final host and mature into adults. The first juvenile to arrive 

becomes the female, and a subsequent juvenile the male. 

Ecology - These parasites range from inshore brackish waters to offshore high 

salinity waters and jacks even carry these isopods into fresh water in Puerto 

Rico (Bunkley-Williams and Williams 1995). They appear to be more common 

on big game fishes living inshore, but are also found in hosts offshore.


Location in Host - The adult female attaches on the top of the tongue, facing 

out. The male is behind and beneath her and lying across the gill rakers. 

Length - Female 23.0-38.0 mm, juveniles 6.5-9.7 mm, first 4 juvenile stages 

6.5, 7.2, 8.1 and 9.0 mm (Williams and Bunkley-Williams 1994). 

Host Specificity - It commonly occurs on crevalle jack and horse-eye jack, and 

occasionally occurs on other jacks and great barracuda. Records from "scombroid 

fishes" appear to have been in error. In marine aquaria it will infect 

almost any fish exposed to juvenile isopods. 

Damage to Host - Little physical damage to the 

host is apparent. There is probably some general 

stunting. Jacks with isopods eat different food 

items, presumably because the isopod takes up 

so much room in the mouth of the host. Large 

numbers of juveniles kill fishes. Juvenile jacks 

can be killed by a few isopods. We have landed 

crevalle jack by hooking them only through the 

isopod in their mouths, demonstrating the strength 

of the isopod's grip on the host. 

Detection - The light-colored female quite 

obviously fills the mouth of the host. 

Harm to Humans - Larger jacks and barracuda 

in Puerto Rico are notorious for carrying harmful 

amounts of ciguatera fish poisoning toxins. The 

sale of these fishes is banned in Puerto Rico. Caribbean folklore suggests that 

fishes with isopods are free from ciguatera. In some fish markets in the Lesser 

Antilles, isopods are illicitly added to fishes to make them sell. This legend has 

not been proven and should not be taken as fact. Research is needed to confirm 

or deny this intriguing idea. 

Significance to Sport Fishing - This parasite may kill a few jacks, so slightly 

fewer are available for sport fishing. Infected jacks are smaller than uninfected 

ones of the same age. Jacks with isopods do not fight as well or as long on 

hook-and-line. Large isopods in the mouth may alarm some sport fishermen, 

but they have no effect on the quality of the fish as food. 

Aquaculture - Thousands of planktonic juveniles of this isopod superinfected 

an adult crevalle jack held in a shallow water trap near Magueyes Island, La 

Parguera, Puerto Rico (Williams and Bunkley-Williams 1994). These isopods 

occur abundantly in the plankton and may cause severe problems for any big 

game fishes cultured in the Caribbean. 

235

236 


Livoneca ovalis (Say) 

This ubiquitous gill-dwelling isopod of the U.S. 

Atlantic coasts is very damaging and important to 

inshore fishes, but relatively rare on big game fishes. 

Name - This isopod is morphologically similar to 

L. redmanii (the next parasite in this book) and 

several authors consider that they are the same. 

Livoneca redmanii is genus specific to Spanish 

mackerels and always occurs in female-male pairs, 

while L. ovalis has no host specificity and occurs 

singly. We believe that these are distinct species. 

"Aegathoa oculata" has been reported from 

Spanish mackerel. This genus was used in the past 

before it was recognized that these isopods were 

juveniles of cymothoid isopods. The genus is now 

considered invalid and is not in use. The isopod was 

probably a juvenile of L. ovalis. 

Diagnostic Characters - This isopod is large, 

cream- or grey-colored, with an oval body shape. It occurs singly (females not 

accompanied by males). 

Records - We found a single immature specimen in a Spanish mackerel from 

Dauphin Island, Alabama, USA. It was also reported once from crevalle jack 

off New York and a juvenile occurred in a Spanish mackerel off Florida, USA. 

Geographic Range - Atlantic and Gulf Coasts of the USA. 

Ecology - Occurs from coastal fresh waters to open ocean, but more commonly 

in inshore marine and brackish water areas. 

Location in Host - Usually in the gill chamber, but occasionally in the mouth. 

It has been reported from the stomachs of several dolphin from off North 

Carolina, as a result of host predation. 


Host Specificity - None. 

Damage to Host - It frequently erodes up to 1/3 of the gill filaments of a host. 

This isopod has been accused of directly transmitting lymphocystis disease, but 

this is unlikely since they permanently associate with a single fish. 

Detection - Easily seen. It sometimes causes an obvious bulge the gill flap and 

often its tail or hind-body is exposed out from under the gill flap of the host. 

Harm to Humans - A court case alleged that this isopod caused injury to 

someone eating an infected Spanish mackerel in August 1994 on the Atlantic 

coast of the USA. We provided testimony in this case and in a similar court 

case in Puerto Rico in 1992 involving an isopod in imported eastern Pacific 

snapper (Isopod Newsletter 1992; 81:1). Fish parasitic isopods in fishes do not 

harm humans that eat these fishes. (If isopods were that dangerous, our isopod 

research would be better funded!)


Livoneca redmanii Leach 

This isopod, found in pairs in the gill-chamber, 

is extremely damaging to mackerels and can kill them, 

causing significant loss of these valuable sport fishes. 

Diagnostic Characters - It is a large, tan, oval- 

shaped isopod that always occurs in female-male pairs 

in the gill chambers of Spanish mackerel. 

Records - We found 2-4 in 26 of 35 cero from 

various localities around Puerto Rico, 2-4 in 2 cero 

from Barbados, 2 in a serra Spanish mackerel from 

Colombia, 2-4 in 2 cero from Jamaica and 2 each in 

3 serra Spanish mackerel from Trinidad. These levels 

may be biased by spearing the hosts. Fishes 

parasitized with this isopod may be more easily 

collected than non-parasitized ones. Records from 

king mackerel and Spanish mackerel are questionable 

and require confirmation. Caribbean records of this isopod on the Spanish 

mackerel probably refer to the serra Spanish mackerel as the 2 fish were 

previously confused and the geographic range of this isopod apparently does 

not overlap that of Spanish mackerel. 

Geographic Range - Caribbean and South American coast to Rio de Janeiro, 

Brazil. It was absent from extensive surveys of Spanish mackerels in Mexico. 

Ecology - Like their Spanish mackerel hosts, they are found around coral reefs 

and in pelagic and offshore habitats. 

Location in Host - The female is located in the ventral portion of the gill 

chamber, facing forward with the dorsal surface against gill flap. The male is 

beneath and partly behind the female. 

Length - Female 19.5-26.0 mm. 

Host Specificity - Only found on cero and serra Spanish mackerel. 

Damage to Host - A female-male pair occupies the gill chamber and destroys 

1/3-2/3 of the gill filaments, thus stunting the growth of the host. Eight of 26 

infections in Puerto Rico and 1 each in Barbados and Jamaica had a pair of adult 

isopods in both gill chambers. These "doubly infected" hosts have very poor 

length-to-body-weight ratios, are generally in poor physical condition, and often 

suffer from other parasite and bacterial diseases. Double infections appear to 

kill the host. A proliferation of gill chamber epithelial tissue (granuloma) was 

found in the identical position of the isopod attachment site of a cero that had 

a pair of isopods in the opposite gill chamber. The productivity of these 

important sport and commercial fishes is drastically reduced by this isopod. 

237

238 


Nerocila lanceolata (Say) 

This dark, external isopod is a "shape shifter" of 

varied body forms that digs large holes in the host flesh. 

Fortunately for big game fishes, it occurs largely on 

inshore fishes. 

Name - The great variety of body forms demonstrated 

by this parasite (see figures) has caused it to be 

repeatedly named as different, unwarranted species. 

Nerocila acuminata Schioedte and Meinert is the most 

popular of these synonyms. 

Diagnostic Characters - It is a large black to black and 

white striped, external isopod. The posterior margin of 

the head is distinctly trilobed. 

Records - Williams, Bunkley-Williams and Rand 

(1994) reported this isopod on a new host, blue runner, 

and new locality in Bermuda. 

Geographic Range - Atlantic and Gulf Coasts of the 

USA and Central America, Bermuda and Cuba. 

Life History - Juvenile or immature isopods occur on 

small fishes, but it is unclear if these are temporary or 

permanent hosts. 

Ecology - This isopod occurs largely on inshore fishes 

but rarely infects big game fishes. 

Associations - It is often infected with external 

hydroids and algae. Striped goose barnacle has been 

reported attached to this isopod and to Nerocila 

californica Schioedte and Meinert. This isopod is 

sometimes found with missing or damaged pleopods, 

pleotelson or legs (pereopods). These injuries are probably due to attacks by 

cleaner fishes. 

Location in Host - Skin and fins. 


Host Specificity - This isopod occurs more commonly on certain inshore, 

bottom (benthic) fishes, but a great variety of fishes are parasitized. Nerocila 

californica occurs on Indo-Pacific sailfish and other billfishes in the eastern 

Pacific, while N. lanceolata does not occur on billfishes in the Atlantic. All 

Nerocila spp. on big game fishes may be a result of prey to predator transfer. 

Damage to Host - This parasite causes considerable tissue damage. External 

lesions often cover more surface area of the host than the isopod. Secondary 

bacterial infections sometimes occur.

PISCES (FISHES) 

Big game fish associates are found in all classes of living fishes (Subphylum 

Pisces). It was spelled "fisch", "fissh" or "fisc" in Middle English, and "fisc" 

or "fish" in Anglo-Saxon. "Fish" is both the singular and plural form for 1 or 

more specimens of a single species of these animals, but the plural "fishes" is 

used for more than 1 fish species. Remoras sometimes harass or bite human 

swimmers and cookiecutter sharks have severely injured fishermen. Fishes are 

our most important source of protein, and have enormous economic value. 

More than 25,000 recently living species of fishes exist, but 4-5 times that 

many names occur. Adult fishes vary in length from the 8 mm central Indian 

Ocean dwarf goby, Trimmaton nannus Winterbottom and Emery, to the 12.6 

meter whale shark. Length of fishes is measured 3 ways: (1) total length (TL) - 

chin to end of tail, (2) fork length (FL) - chin to the middle of the fork of the 

tail, and (3) standard length (SL) - chin to the crease at the base of the tail 

(hypural plate) caused by folding the tail over. TL will be used when available. 

SL is used by scientists to avoid variations caused by worn or damaged fin tips. 

FL replaces SL for big game fishes, sharks etc. whose tails do not fold. See 

Host Summaries for a labeled illustration of lengths and external structures. 

Sailfish may be the fastest fish (clocked at 68 miles-per-hour [122 KPH]); 

and sturgeons, the longest lived (at least 150 years). Fishes live only in water 

(aquatic), respire with permanent gills, have fins, a 2-chambered heart, a skin 

usually covered with scales, an internal skeleton of bone or gristle (cartilaginous) 

and are usually cold blooded (some tunas and offshore sharks are notable 

exceptions to the "cold-blooded" criteria). 

Associates occur in all 3 classes of fishes: (1) Lampreys are primitive, eellike 

and lack jaws, scales, paired fins and bone. Only 1 of the 38 known 

species found in marine and fresh waters in North America and Eurasia 

associates with western Atlantic big game fishes. (2) Sharks have jaws, an 

internal skeleton of cartilage and 5-7 gill slits on each side. Two of the approximately 

2000 known living species associate with our big game fishes. (3) Bony 

fishes have jaws, an internal skeleton of bone and a single gill opening on each 

side. Eight of the 20,000 known living species associate with big game fishes. 

The sexes of fishes are separate, although some species change sexes during 

their lives. Fertilization is external or internal. Development is direct whether 

in the plankton or in the body of the female. Reproduction must be a problem 

for those fish associates isolated on big game fishes. 

Fishes are not permanent (obligate) parasites of big game fishes and can 

move freely from host to host. Lampreys and cookiecutter sharks usually just 

attach to the host for feeding. Lampreys slowly feed on blood and dissolving 

flesh, while cookiecutter sharks are assumed to more quickly twist out plugs of 

muscle and skin. This can be called "micro-predation" or "casual parasitism". 

It is not so different from that found in leeches and aegiid isopods, which are 

traditionally lumped with obligate parasites. Remoras attach, but most are 

239

240 


loosely, and pilotfish do not attach, to big game fishes. They only eat 

abandoned scraps of food, voided fecal material and obligate parasites they pick 

from these hosts. Remoras and pilotfish are traditionally called "commensals" 

of big game fishes. Although some young remoras live in the host gill 

chambers, spend more time with their hosts and may feed directly on host 

materials. Family groups are sometimes reared on a single big game host without 

leaving the fish. These relationships could be more intimate than those of 

the other host associates and could eventually develop into parasitism for at least 

part of their life cycles. Williams and Bunkley-Williams (1994) have suggested 

possible paths that free-living organisms take toward parasitism. In addition, 

these associates may transmit or disseminate parasites and diseases to their large 

hosts although this has not previously received any attention (see Discussion). 

These associates or parasites should be examined for their own parasites 

before they are preserved. To preserve these fishes, the right side of the body 

should be slit into the body cavity, and the whole fish placed in 10% formalin 

that is at least 10 times the volume of the fish. 


Phylum Vertebrata - vertebrates Page 

Subphylum Pisces - fishes 

Class Agnatha - jawless fishes 

Order Petromyzontiformes 

Family Petromyzontidae - lampreys 

Petromyzon marinus - sea lamprey .................................................... 241 

Class Chondrichthyes - cartilaginous fishes 

Order Squaliformes 

Family Squalidae - dogfish sharks 

Isistius brasiliensis - cookiecutter shark .......................................... 242 

Isistius plutodus - largetooth cookiecutter shark ............................. 246 

Class Osteichthyes - bony fishes 

Order Perciformes 

Family Echeneidae - remoras ................................................................. 247 

Echeneis naucrates - inshore remora .............................................. 249 

Echeneis neucratoides - whitefin remora ........................................ 251 

Phtheirichthys lineatus - slender remora ........................................ 252 

Remora brachyptera - spearfish remora ......................................... 253 

Remora osteochir - marlin remora .................................................. 254 

Remora remora - shark remora ....................................................... 255 

Remorina albescens - white remora ................................................ 257 

Family Carangidae - jacks 

Naucrates ductor - pilotfish ............................................................ 258


Petromyzon marinus Linnaeus - sea lamprey 

This lamprey kills or mutilates big game fishes. 

Diagnostic Characters - Sea lampreys are elongate fish with a dorsal fin 

separated into 2 parts by a deep notch. The mouth is a sucking disk lined with 

teeth (horny cusps). The wound this fish produces is circular with the scales 

and skin (epidermis) of the host completely abraded by the teeth of the sucking 

disk, and a hole rasped through the skin in the center of the wound. 

Records - It occurs on Atlantic mackerel and swordfish off Atlantic coast of 

the USA; and on a variety of fishes on both sides of the North Atlantic from 

Labrador (possibly Greenland) to Florida and the northern Gulf of Mexico, and 

from northern Norway through much of the Mediterranean. 

Geographic Range - North Atlantic and Mediterranean. 

Life History - The larvae (ammocoetes) live in the mud of streams 5-13 years, 

before they metamorphose, at a length of 10-20 cm, into adults, and move 

down to the ocean or lakes where they begin to attack fishes. Adults spend 

about 2 years parasitizing a wide variety of fish hosts, then return to freshwater 

streams to breed (anadromous) in April to July. They build nests in streams 

with rapidly moving water and gravel stream beds, and die after spawning. 

They produce 124,000-305,000 eggs, which is the highest for any lamprey 

species. Spawned out lampreys may be parasitized by fungi Saprolegnia spp. 

before they die. 

Ecology - Sea lampreys tolerate a wide range of temperatures and salinities (0- 

35 ppt). They are eaten by other fishes in fresh water and by swordfish and 

striped bass at sea. Sea lampreys appear to be more common in inshore and 

even brackish waters, but are found offshore to a depth of 200 m and 

occasionally down to depths of at least 1100 m. They are not caught on hookand-line, 

and seldom in nets, thus little is known of their habits at sea. 

Parasites and Diseases - Sea lampreys from the ocean have rarely been 

examined for parasites and diseases. A roundworm, Truttaedacnitii stelmioides 

(Vessichelli), occurs in this lamprey from the Mediterranean and North Sea. In 

fresh waters, they have a typical freshwater parasite fauna. 

The liquefying and slurping of host tissue seems to transfer larval parasites 

from the host to the lamprey in fresh waters. The same mechanism probably 

occurs in the ocean. This results in the ironic situation that the final hosts of the 

sea lamprey are the intermediate and intermediary hosts for its parasites. 

Location in Host - Sides. 

Length - 13.5-86.0 cm TL, occasionally up to 120.0 cm. Sea lampreys are the 

largest species of lampreys. 

Host Specificity - Sea lampreys parasitize almost any large fish, but usually 

not whales. They have rarely been reported from a variety of porpoises and 

241

242 


whales. Their Pacific cousin, Pacific lamprey, Lampetra tridentata (Gairdner), 

attacks fishes as well as whales. 

Damage to Host - This parasite rasps a hole in the side of the host's body, and 

sucks blood and body fluids, and slowly dissolves and consumes other tissues. 

More than 1, sometimes 3-4 lampreys, may attack the same host and cause 

severe injury. Weakened hosts with open wounds are more susceptible to other 

parasites and microbial diseases. 

Significance to Sport Fishing - This parasite kills and severely injures 

freshwater sport fishes. The damage to offshore big game fishes must be 

considerable, but is more difficult to determine. Young sea lampreys are used 

as bait for freshwater sport fishes. Sea lampreys are food items for swordfish, 

and in turn sea lampreys attack and feed on swordfish. This is a most circular, 

and peculiar, predator-prey relationship. 

Land-locked populations occur in freshwater lakes. Sea lampreys do terrific 

damage to freshwater sport fishes. After the Welland Canal was opened 

around the barrier of Niagara Falls, this parasite was transferred into most of 

the Great Lakes of North America, and at least 3 species of locally important 

sport and commercial fishes were destroyed. The ecologically and economically 

disastrous transfer of this exotic disease organism into much of the Great Lakes 

is a classic case study of unintended results often incurred during ecological 

modifications. 

Comments - Sea lamprey was considered a delicacy in Europe during the 

Middle Ages, and was a commercial fish in New England until about 1850. The 

flesh is reputed to be delicious, if you can avoid considering how they earn a 

living. They are usually caught when breeding or ascending streams to breed 

(spring run). Sea lampreys are frequently used as preserved specimens in 

biology classes, and live ones can be used in classroom experiments (Cochran 

1989). They may be one of the best studied of all parasites, yet little is known 

of their behavior and habits in the open ocean. 

Isistius brasiliensis (Quoy and Gaimard) - cookiecutter shark 

This small, but violent shark tears fist-sized chunks of flesh out of some of 

our most important offshore game fishes. Wounds and scars from these attacks 

are unfortunately quite common.


Name - It was called "cigar shark" by Robins et al. (1991), but "cookiecutter 

shark" has been used in most publications. 

Diagnostic Characters - It is a relatively small, cigar-shaped shark with a 

darkly pigmented collar behind the head. The caudal fin is high. There are 25- 

31 rows of teeth in the lower jaw. It produces large, cone-shaped wounds, 

sometimes called "crater wounds", on the outside of big game fishes. 

Records - Occasionally 1-3 (usually 1) wounds were seen on albacore, Atlantic 

blue marlin, blackfin tuna, dolphin and yellowfin tuna from various localities 

off Puerto Rico. These wounds are seen on big game fishes off Cuba; are 

common on dolphin off Florida; 1-3 (usually 1) on swordfish from the Gulf of 

Mexico; and albacore, Atlantic blue marlin, bigeye tuna, blackfin tuna, bluefin 

tuna, dolphin, pompano dolphin, wahoo, white marlin and yellowfin tuna 

throughout the oceanic western Atlantic. We saw 1 wound each on albacore, 

bluefin tuna and yellowfin tuna off Okinawa and off the Ryukyu islands east of 

Taiwan. It also occurs on black marlin, Indo-Pacific blue marlin, Indo-Pacific 

sailfish, rainbow runner and other jacks and scombrids from Hawaii; and on 

albacore, dolphin, skipjack tuna and wahoo in the central Pacific. These 

wounds are also found on a variety of marine mammals. 

Geographic Range - Worldwide. Fresh wounds are seen on hosts in warmwater 

areas, while only old wounds and scars are seen in temperate and colder 

areas. This indicates that cookiecutter sharks occur in tropical and subtropical 

areas, and that wounded hosts may move further north or south after attacks. 

Life History - This fish presumably holds eggs in the body and releases 

hatched young (ovoviviparous). Ovaries contain 6-7 large eggs, but embryos 

and litter size is not known. Young are about 14 cm long at birth. Males 

mature at a length of 31-37 cm, females at 38-44 cm. 

Ecology - Besides chunks of host flesh, this shark also eats squids almost as 

large as itself, deep water lightfishes (Gonostomatidae) and crustaceans. It probably 

attacks fishes and mammals basking near the surface. Fast swimming tunas 

and porpoises probably seek out the cookiecutter shark as potential prey, and are 

attacked before they realize their mistake. This has not been observed, but 

circumstantial evidence supports this scenario: (a) cookiecutter shark luminescence 

appears to be a predator attracting device, (b) many wounds are near the 

head, often made from in front of the host, of these large predators, suggesting 

a head-to-head encounter with the cookiecutter shark, (c) wounds only appear 

on some predators after they achieve a body size associated with predation on 

fishes, (d) a cookiecutter shark was found in the stomach of a large Spanish 

mackerel, Scomberomorus sp., caught in the tropical eastern Pacific, although 

they have not been noted in the stomach contents of other big game fishes. 

243

244 

PARASITES OF OFFSHORE BIG GAME FISHES OF PUERTO RICO


This fish is collected from the surface at night, but usually at depths from 

85-3500 m (epipelagic to bathypelagic). Its preferred and maximum depths are 

not known. This shark seems to migrate vertically from great depths to at or 

near the surface at night (vertical migrator, diel cycle). It is more commonly 

found near islands, whether this is caused by using an inshore pupping ground, 

or the attraction of more hosts, is not known. This shark may be able to 

tolerate areas of low oxygen. The relatively large and oily liver and body cavity 

may be a buoyancy adaptation to its deep habitat or its heavy skeleton necessary 

to support violent attacks. The entire belly and particularly the neck is covered 

with luminous organs (photophores) which bathes the shark in a bright, ghostly 

green light at night. This luminescence may lure in large predators (such as the 

cyalume lights used on longlines). Luminescence fades when the fish dies. 

This shark does not appear to be very abundant in most areas, but is so 

common in the eastern Atlantic that it is bottom trawled for fish meal 

production. Studies have reported multiple wounds on every swordfish observed 

in some areas of the eastern Atlantic. The large, offshore game fishes in this 

region must take a beating from these parasites. 

Parasites and Disease - Unknown. 

Location in Host - Body. The wound is often near the head of the host, but 

may be on the flank or any part of the body. 

Length - 14.0-50.0 cm TL; female up to 50.0 cm; male 39.0 cm; wounds 2.0- 

7.0 cm (1.2-5.0 cm wide). 

Host Specificity - This fish has no specificity. It may prefer larger hosts. 

Cookiecutter sharks even bite the rubber sonar domes off nuclear submarines. 

Marine mammals including most dolphins and whales are attacked. It does not 

even spare fellow sharks, such as megamouth shark, Megachasma pelagios 

Taylor, Compagno and Struhsaker. 

Damage to Host - This shark attaches to the body of a host with highly 

specialized sucking lips and a strongly modified pharynx. It can grab hosts with 

the sharp teeth of its upper jaw. This parasite drives the razor-sharp, triangular 

teeth of the lower jaw into the host, and swings its body in a circle, cutting out 

a cone-shaped piece of flesh. 

Compagno (1984) suggests that the mortality rate of offshore game fishes 

might be increased by the attack of this shark. The wounds encourage infection 

by microbial and parasitic diseases. However, the abundance of these scars 

suggest that many hosts survive. 

In an ancient Samoan legend, atu [=skipjack tuna] entering Palauli Bay left 

small round pieces of their flesh near the beach as sacrifices to Chief Tautunu. 

This was evidenced by people catching atu, who found fresh, round wounds on 

the fish sides. 

Harm to Humans - No reports of attacks on free-swimming humans are 

known, possibly because few people swim or dive in oceanic waters at night. 

If you are contemplating a midnight swim off your boat, you should consider 

that this shark seems willing to attack any large animal. As you would expect 

245

246 


in a small fish attacking giant predators, this shark has remarkably quick 

reactions. 

Preparation for Study - This fish should be examined for parasites. The 

stomach and intestine should also be examined for fish flesh and scales which 

may be used to identify its recent hosts. 

Significance to Sport Fishing - Some fishes may die after cookiecutter shark 

attacks, and others may suffer reduced growth. These obvious wounds, or 

scars from the wounds, are distressing in your trophy game fish. 

Commercially, the wounds may reduce the price paid for "perfect" tuna. 

Many whales receive multiple wounds when they migrate into warm 

waters each year. These scars are permanent, but change in color and 

appearance over time. The series of wounds received each year can be 

recognized and used to determine how many migrations have occurred. These 

wounds might be used similarly as biological tags to identify migrating big 

game fishes. 

Comments - These wounds were once thought to be caused by small, internal 

parasites, bacterial diseases or even the isopods sometimes found in these 

wounds. Thus scientific thought about this phenomenon, until recently, was 

little better than the Samoan legend. Solving this mystery was a neat bit of 

oceanography pieced together from the indirect evidence of chunks of flesh 

resembling the wounds and the odd teeth of the cookiecutter shark, and scientists 

twisting holes in pieces of fruit and big game fishes on the deck with dead 

cookiecutter sharks. 

Popular Reference - Klemm (1984). 

Isistius plutodus Garrick and Springer - largetooth cookiecutter shark 

This shark has proportionately the largest teeth of any living shark. 

Fortunately it is rare, as it bites off larger chunks of flesh and bone from big 

game fishes than the cookiecutter shark. 

Diagnostic Characters - This small, cigar-shaped shark lacks a darkly 

pigmented collar behind the head. The tail (caudal fin) is low. There are 19 

rows of teeth in the lower jaw. It gouges out characteristically elongate plugs 

of tissue from its hosts, but it may also produce round wounds which are 

similar to those made by the cookiecutter shark. 

Records - We found 1-2 oval, elongate wounds in 2 of many Atlantic blue 

marlin and in 3 yellowfin tuna caught from various localities off Puerto Rico. 

Wounds have been found on albacore, bluefin tuna and yellowfin tuna in the 

Pacific, but only 2 specimens of largetooth cookiecutter shark have been 

collected (one 100 miles south of Dauphin Island, Alabama, USA; and another 

off Okinawa, Japan).


Geographic Range - Worldwide. Our observations of the wounds of this 

shark are the first from the Caribbean. 

Ecology - This fish was collected from the surface at night over depths from 

814-997 m. Its preferred and maximum depths are not known. This shark 

probably migrates vertically from great depths to at or near the surface at night 

(vertical migrator, diel cycle). The relatively large and oily liver and body 

cavity may be a buoyancy adaptation to its deep habitat or its heavy skeleton 

necessary to support violent attacks. Its smaller fins suggest it is a weaker, less 

active swimmer than the cookiecutter shark, yet it has a larger mouth, more 

powerful jaw, and larger teeth. It has fewer luminous organs (photophores) and 

is therefore not as brightly lit as the cookiecutter shark. The rarity of this shark 

may be due to it possibly occurring at greater depths than the cookiecutter shark. 

This shark also has a more binocular-type vision than the cookiecutter shark. 

This kind of vision, more similar to our own, is well suited to a parasite seeking 

highly mobile hosts. 

Length - at least 42.0 cm TL. 

Host Specificity - Unknown. Atlantic blue marlin is a new host. 

Damage to Host - The wounds caused by this shark are larger than those of 

cookiecutter sharks, and involve bone as well as tissue. They may be more 

harmful. 

Comments - Life History, Location in Host, Parasites and Disease, 

Detection, Harm to Humans, Preparation for Study, and Significance to 

Sport Fishing are the same as for cookiecutter shark above. 

Family Echeneidae (Remoras or Suckerfishes) 

The 8 species of remoras form a family of bony fishes. The Greek 

"echeneis" and Roman "remora" refer to the belief that these fishes could attach 

to ships and hold them back. Various mystical properties have been assigned to 

suckerfishes. Remoras are blamed for the ill fates of Emperor Caligula and 

Mark Antony. Shamans in Madagascar still attach part of the suction disks of 

remoras to the necks of wives to ensure faithfulness to absent husbands. 

Remoras are easily recognized by the sucking disk (a modification of the 

first dorsal fin) on top of their heads. 

This organ consists of numerous pairs of 

cross ridges or laminae. The movable 

disk ridges work much like a Venetian 

blind. When the rear edges are raised, 

a suction for holding on to a host is 

created. Pulling a remora backward only 

increases the suction, while pushing it 

forward releases the hold. The size of the sucking disk in adults can be correlated 

to the activity of their hosts (those on the most active or fastest hosts have 

the largest) and to their body width (the widest or bulkiest have the largest). 

247

248 


Most can swim swiftly, but probably not great distances. Their coloration 

can be quite beautiful, particularly in young, and it is unfortunately, quite 

variable at any age. Identification of young remoras is difficult, and the adults 

of 2 species are very similar in appearance. Many previous host records are 

highly suspect. Development is direct, but some species have a long filament 

on the tail in larval and postlarval stages. Early stages are planktonic. Some 

develop multiple fangs of mysterious function in their jaws. They begin 

attaching to hosts when they reach a length of 4-8 cm, and still have the tail 

filament. Wahoo and a variety of other smaller fishes have been called "trial 

hosts" (intermediary hosts) because only small or young remoras are found on 

them. Larger remoras are more specific. Two species almost always attach to 

billfishes, and 1 each prefer great barracuda, sharks, or manta rays, while whale 

remoras, Remora australis (Bennett), are only found on marine mammals. 

Two species attach to a variety of inshore fishes, sea turtles, mammals and even 

ships and buoys. Some spend considerable time free swimming away from any 

host. Remoras usually attach on the bodies of their hosts, but some can be 

found in the gill cavities or mouths. 

Remoras receive transportation, some degree of protection, and food from 

their hosts. In return, they pick parasites and damaged or diseased tissues from 

the bodies, gills and mouths of their hosts. Host cooperation in this supposed 

"partnership" is less certain. Some hosts appear to try to avoid or shake off 

remoras, and they have been found in the stomach contents of their hosts. 

Much argument but less data occurs in the literature concerning the food 

habits of remoras. They seem to be opportunists taking host ("scraps" or debris 

of food from their large host's meals, ectoparasites, and possibly wastes [feces]) 

and non-host materials (plankton and possibly small fishes from the water 

column). The food of most remoras is pieces of fishes, plankton is less 

important, and parasites occur in token amounts. Besides the standard 

preparations mentioned above, their stomach contents should be quickly 

examined or preserved for parasites (before they are digested) that they may 

have cleaned from their host. The reason that only copepods and isopods have 

been found in the stomachs of remoras may be that the "shells" (exoskeleton) 

of these parasites do not quickly digest or decompose. Stomachs of remoras 

must be immediately removed, opened, and placed in 10% formalin. Only then 

can we find capsalid and other gill worms, tissue flukes and other fragile 

parasites that they may pick off their hosts. 

Remoras are easily seen on the body of hosts, but are less obvious in the 

gill cavity or mouth. Just how important their parasite removal may be to large 

offshore game fishes is unclear. They appear to cause little damage or irritation 

to their large hosts. Remoras attached to lines have been used by fishermen to 

snag large fish and turtles. The most famous report was reputedly by 

Christopher Columbus of suckerfish "living fish hooks" by the Arawak Indians 

in Cuba. The technique is actually worldwide and ancient. 

One fossil remora resembles a pilotfish, suggesting free-swimming 

associated fishes may have evolved into remoras. A Tertiary (1-70 million years


ago) fossil remora had fewer (7-8) and more widely spaced sucker laminae, 

while another specimen had a disk similar to inshore remora. The number of 

laminae may have increased and the spacing decreased during remora 

evolution. Aristotle (384-322 B.C.) observed "dolphin's louse" on dolphin in 

the Mediterranean. The former common names of remoras were established 

before much of their biology and habits were understood, and thus some are 

inappropriate, and they employ a hodgepodge of 4 synonymous terms (diskfish, 

remora, -sucker, suckerfish). 

Echeneis naucrates Linnaeus - inshore remora 

This most common and largest remora is found mostly inshore and on 

almost any kind of host. It often harasses humans. 

Name - The accepted common name of this fish is "sharksucker", and it is also 

called "shark remora". These names are inappropriate as Echeneis naucrates 

is a generalist occurring on a variety of inshore hosts, not just sharks. 

Diagnostic Characters - The body is slender. The sucking disk is relatively 

long, extending backward slightly beyond the middle of the pectoral fin, and 

has 21-28 (usually 23-24) pairs of laminae, with 3-4 rows of spinules along the 

posterior margin. The caudal fin is truncated in the adult, but has a long central 

filament in young 10 cm or smaller. As the remora grows, the filament gradually 

decreases in relative length, until at 18 cm, it is reduced to a short lobe. 

The body is usually striped with brown or black. Normally the belly is white, 

but when it attaches with its body upside down, it reverses its color pattern. 

Records - We found 1-3 (usually 1) on 3 of 47 great barracuda, 2 of 20 

crevalle jack and 1 of 27 horse-eye jack from various localities around Puerto 

Rico. It has also been found on great barracuda from St. Thomas, U.S. Virgin 

Islands, and Florida, USA. We have also seen it on small coral reef fishes, and 

8-12 on 6 of 25 West Indian manatees, Trichechus manatus Linnaeus, from 

various localities around Puerto Rico. This remora attaches to sharks (up to 12 

per host), large bony fishes, rays, whales, boats, floating timbers and other 

objects around the world. 

Geographic Range - Worldwide, except in the eastern Pacific. 

Life History - This remora is often found swimming away from a host. It may 

not have the problem of locating a mate, unlike more host-dependant remoras. 

Ecology - It occurs on both inshore and offshore hosts. An unusual 

development of the respiratory muscles allows this fish to pass a considerable 

amount of water over the gills, thus allowing it to thrive in still waters and on 

hosts with a variety of activity patterns. This remora is not a rapid swimmer. 

It is the most common, inshore remora found in the western Atlantic. Inshore 

249

250 


remoras can easily be caught with hook and line. This remora appears to be 

rather loosely associated with hosts and to change fish hosts frequently. They 

often abandon hosts when hosts are removed from the water. 

Parasites and Disease - Two gillworms occur in the gills of inshore remora, 

Dioncus agassizi from the Indian Ocean and Dioncus remorae from West Indies 

and the New York Aquarium. Dioncus agassizi is also found in cobia, shark 

remora and spearfish remora, and D. remorae occurs in crevalle jack, which is 

a host for inshore remora. Larval tissue flukes, Didymozoides sp., occurred in 

the gill cavity of this remora. One fluke, Parahemiurus merus, occurred in 1 

of 2 inshore remora from Jamaica, and also occurs in jacks that are hosts of this 

remora. Flukes found in the stomach or intestine of this remora include: 

Sterrhurus musculus off Mexico and Florida (also found in crevalle jack and 

great barracuda which are hosts for this remora); Lecithochirium monticellii, 

from Woods Hole, Massachusetts, USA (USNPC 8352), which possibly occurs 

in blue runner and little tunny from the same locality; Echeneidocoelium indicum 

Simha and Pershad from the Indian Ocean; Stephanostomum imparispine 

(Linton) (which also occurs in western Atlantic cobia) and Tubulovesicula 

lindbergi (Layman) from the South China Sea; and Tormopsolus echenei 

Parukihin from the Gulf of Tonkin. Two larval tapeworms, Nybelinia robusta 

in the stomach and Tentacularia coryphaenae encysted postlarvae in the stomach 

and intestine of inshore remora, from Woods Hole, Massachusetts, USA. The 

later species also occurs in a variety of big game fishes. Five roundworms 

occurred in the intestine of this remora, Spirocamallanus olseni Campana and 

Razarihelissoa, from the Sea of Nossi-Bé; Ascarophis sp. of Parukhin, 

Capillaria echenei Parukhin, Raphidascaris sp. of Parukhin, and Spinitectus 

echenei Parukhin from the South China Sea. Three spiny-headed worms, 

Telosentis tenuicornis (Linton), from Massachusetts, USA; Gorgorhynchus 

medius (Linton) from the South Atlantic; Serrasentis sagittifer were found in the 

intestine of inshore remora from the South China Sea. The following copepods 

occur on this host, Tuxophorus caligodes, and on cobia from the Western 

Atlantic; Caligus praetextus from the Gulf of Mexico off Florida; and 

Lepeophtheirus longipes Wilson in the mouth from west Africa. Caligus 

coryphaenae was reported externally on an Echeneis sp.(?) in the eastern 

Pacific. This could have been an inshore remora, but this fish is not supposed 

to occur in the eastern Pacific. This copepod occurs on a variety of big game 

fishes. A fish louse, Argulus varians Bere, was reported on this fish from the 

Gulf of Mexico off Florida, USA. Striped goose barnacles occurred on inshore 

remoras from west Africa. 

Cleaning Behavior - Inshore remoras clean parasites from their host, but 

parasitic copepods and isopods were not a major food item (16% of 87 stomachs 

containing food). Smaller remoras (57-85 mm) occasionally picked parasites, 

medium-sized (86-311 mm) often cleaned, while larger fish (311-630 mm) had 

no parasites in their stomachs. 

It is frequently seen free swimming and may often move from host-to-host. 

Five free-swimming fish had parasitic crustacea in their stomachs, and 1 remora


on a lemon shark, Negaprion brevirostris (Poey), had a Caligus sp. in its 

stomach, a copepod that does not occur on sharks. In 7 other cases, the 

copepods in the stomach of the inshore remora apparently originated from the 

species of host on which it was associated, suggesting that it does remain on 

one host in some cases. We found Caligus lobodes in the stomach of an 

inshore remora from a barracuda, the host of this copepod; and it has previously 

been reported from this association. 

Predation - It occurred in stomachs of Indo-Pacific sailfish and striped marlin 

in the Pacific. 

Associations - Two attached to a white marlin with a white remora; and it 

shares West Indian manatees with whitefin remoras. 

Location in Host - Body, never in the gill cavity or mouth. 

Length - 5.7-100.0 cm (rarely 1.22 m), but usually less than 60.0 cm. Reputed 

observations of 1.3 m inshore remoras have not been confirmed by measurements. 

The largest individuals have been found associated with the largest fish, 

whale sharks. 

Host Specificity - This remora will attach to almost any large to medium-sized 

fish, sea turtle, marine mammal or boat. Oddly, We have seen them attached 

to coral reef fishes that were smaller than the remora, such as sand divers, 

trunkfishes and parrotfishes. Schwartz (1977) found some fishes in 125,000 liter 

tanks could out swim or otherwise avoid attachment of this remora. 

Damage to Host - We have observed many fishes attempting to scrape off or 

knock off inshore remoras. The remora is generally faster than the host, and 

simply dodges the impacts, until the host tires. This would suggest that many 

species of fishes are irritated by the presence of this remora. Schwartz (1977) 

found some species of captive fishes were severely damaged or killed by 

prolonged or repeated disk attachment of inshore remoras, but those most 

injured were not naturally used as hosts in the wild. 

Harm to Humans - Inshore remoras frequently harass bathers and divers. We 

are not certain if most of these "attacks" are attempts to attach or to pick 

imagined parasites off of humans. Few people are sufficiently calm under the 

attention of this remora to learn their intentions. A "crazed" specimen harassed 

human swimmers, including our ichthyology professor, in the "swimming area" 

off the Magueyes Island Marine Laboratories Medusa Dock for days. 

A friend, who was scuba diving in the northern Gulf of Mexico, was 

severely bitten on the breast nipple by an inshore remora. The fish apparently 

mistook this tissue for a dark, juicy copepod or isopod. 

Echeneis neucratoides Zuieuw - whitefin remora 

This less common cousin of inshore remora is poorly understood. These 

2 remoras are very similar in appearance and have often been confused. 

Diagnostic Characters - It is similar to inshore remora, but the body is usually 

striped, stouter, and fins have more white areas (unfortunately, these characters 

are comparative and almost useless, unless fresh specimens of both species are 

available). The sucking disk is long, extending backward slightly beyond the 

251

252 


middle of the pectoral fin, and has 18-23 pairs of laminae. The caudal fin is 

truncated in the adult, and has a long central filament in small young. The body 

is usually striped 

Records - This remora probably occurs on big game fishes inshore, but has 

been confused with inshore remora. We found it on West Indian manatees in 

Puerto Rico, and it has been reported from sharks and other inshore fishes. 

Geographic Range - Western Atlantic. It is the only remora that does not 

occur worldwide. 

Ecology - This remora is an inshore species. It is much less common than 

inshore remoras, but is often mistaken for this species. It can be taken with 

hook and line. 


Food Habits and Parasites - Strasburg (1959) discounted the reports of 

remoras feeding on their hosts' wastes (feces) in Smith (1950). He further suggested 

that materials in the stomach contents of remoras could have been from 

their hosts' scraps or feces, and the 2 sources could not be distinguished. The 

principal item in the stomach contents of a whitefin remora we collected from 

manatees was fecal material from the host. Manatees do not eat fish, thus fish 

scraps are not available for associated remoras. Manatee fecal material is 

largely of plant origin, and can be more certainly identified than feces from 

piscivorous hosts. Our records suggest that fecal material is an important 

component in the diet of remoras. The consumption of fecal material may 

directly transmit alimentary tract parasites between fish hosts and remoras. 

Associations - See inshore remora. 


Length - 6.0-81.0 cm, but usually less than 60.0 cm. 


Phtheirichthys lineatus (Menzies) - slender remora 

This rare remora has only been consistently found in the gills of great 

barracuda. 

Name - The accepted common name of this fish is "slender suckerfish". It is 

also called "slender remora", "striped louse-fish" and "Pega de las Picudas" 

(sucker of barracudas in Spanish). 

Diagnostic Characters - The body is elongate, and the head and sucking disk 

are relatively small. The sucking disk has 9-11 (usually 10) pairs of laminae. 

The caudal fin is rounded in the adult, but has a long central filament in young 

smaller than 9 cm. The body is usually striped.


Records - It occurred often in many great barracuda from Cuba; 1 in a great 

barracuda from Hog Sty Island, Bahamas; and 1 in 1 of many great barracuda 

from Dry Tortugas, Florida, USA. It was also found attached to Pacific sea 

turtles, long-line buoys, bait, or free swimming. 


Life History - Planktonic larva 14 mm or shorter do not have sucking disks. 

The developing disk just touches the back of the head in remoras 21.2 mm 

long. In 32 mm long fish, the disk is 2/3 on the head, and progresses further 

onto the head in larger fish. It is fully formed in 50 mm fish. 


Cleaning Behavior - Parasitic copepods have been found in its stomach. 

Predation - This remora is not immune from predation by its larger hosts. It 

has been found in the stomachs of Atlantic mackerel, in unidentified Pacific 

tuna, and yellowfin tuna. 

Location in Host - Gills, mouth, rarely body. 


Host Specificity - It has been most frequently reported from great barracuda. 

Remora brachyptera (Lowe) - spearfish remora 

It is often found on the body or in the gills of billfishes and swordfishes. 

Its habits are similar to the marlin remora, but it is easily distinguished. 

Name - It is sometimes placed in the genus Remoropsis. 

Diagnostic Characters - The body is short and robust. The sucking disk 

extends posteriorly to slightly beyond the front edge of the base of the pectoral 

fin, and has 14-19 pairs of laminae (15-19 in the Gulf of Mexico, 14-17 in the 

Pacific). The caudal fin is truncated. 

Records - It occurs on Atlantic blue marlin, Atlantic sailfish, longbill spearfish, 

swordfish, white marlin in the Atlantic; black marlin and other billfishes in the 

Pacific; rarely in barracuda and other fishes; and occasionally free swimming. 

Two or more spearfish remoras may occur on a host. 

253

254 



Life History - Larvae and postlarvae have no caudal filament. 

Parasites and Disease - The gillworm, Dioncus agassizi, occurs in the gills 

from the Atlantic coast of the USA. It is also found in cobia, inshore remora 

and shark remora. 

Cleaning Behavior - It does not appear to feed very often on parasitic 

copepods, but only 38 stomachs have been examined (17% with food). 

Immature caligid copepods occurred in the stomach of a spearfish remora from 

an Atlantic sailfish, Gloiopotes watsoni Kirtisinghe from black marlin, and 

Gloiopotes huttoni from striped marlin. 

Predation - This remora has been found in the stomachs of yellowfin tuna and 

unidentified tunas. The stomach contents of this remora contained an 

unidentified remora off Hawaii. 

Location in Host - Body or gill chamber. 


Host Specificity - It is almost always found on billfishes and swordfishes. 

Remora osteochir (Cuvier) - marlin remora 

This remora is often found on the body or in the gills of sailfish and white 

marlin. Its habits are similar to the spearfish remora, but it is easily 

distinguished. 

Name - The accepted common name of this fish is "marlinsucker". This name 

means essentially the same as "marlin remora", and we prefer this 

standardization. Also called "Pega de los Agujas" (sucker of billfishes in 

Spanish). Sometimes placed in the genus Rhombochirus. 

Diagnostic Characters - The body is slender and has a deep longitudinal 

groove along the midline of the lower (ventral) surface. The sucking disk is 

spectacularly large, extending backward well beyond the tip of the pectoral fin, 

and has 17-20 (usually 18) pairs of laminae. The caudal fin is truncated. 

Records - We found 1-2 (usually 2) in 5 of 40 Atlantic blue marlin and 1 in 1 

of 5 Atlantic sailfish from various localities around Puerto Rico. Gudger (1926) 

repeated sportfishermen reports that it occurred in almost every sailfish caught 

off the Florida Keys. Cressey and Lachner (1970) found it on 82 white marlin, 

55 sailfish, but less often on Atlantic blue marlin, swordfish, longbill spearfish, 

and dolphin. It also occurred in great barracuda from the New York Aquarium 

and the Florida Keys; on wahoo, Pacific billfishes, Pacific sharks, ocean sunfish 

from the Pacific; and occasionally free swimming.



Life History - Multiple specimens of this remora occur on many hosts and 

almost always include both sexes. Thus marlin remoras are not dependant on 

the congregation of many hosts to ensure reproduction. Larval and postlarval 

remoras do not have a caudal filament. Small marlin remoras occur on smaller 

fishes, such as wahoo, while small to large ones are found on billfishes. 

Ecology - This small remora seems to be more dependent on the host for 

transport and protection than other species of remoras. It may also feed on 

mucus and materials from the gills of the host. Other remoras are associates of 

fishes, but marlin remoras show some characteristics found in parasites. 

Cressey and Lachner (1970) suggested that these remoras were intimately 

associated with their hosts, probably breed on the host, and form "families" on 

a single host consisting of a gravid female, a ripe male, 1-2 large juveniles and 

several smaller juveniles. This association could be evolving toward parasitism. 

Parasites and Disease - The copepod, Lepeophtheirus crassus (Wilson), 

occurred on the body of this remora from the Gulf of Mexico, on shark remora 

and white remora from the Indian Ocean, and whale remora from the Pacific. 

Cleaning Behavior - Unlike inshore remoras and shark remoras, larger (166- 

230 mm SL) marlin remoras appear to utilize parasites as a source of food more 

often than the smaller (26-125 mm SL) ones. However, this distinction was 

only based on 8 stomachs with copepods. Parasitic copepods Gloiopotes 

americanus were found in the stomachs of marlin remoras on the Atlantic 

sailfish; Gloiopotes ornatus, caligoid, and Pennella sp. from white marlin; and 

Pennella sp. and Caligus sp. from Atlantic blue marlin. These copepods match 

the hosts in which the remoras were found as would be expected in this associate 

that is not thought to often change hosts. 

Predation - This remora has been found in the stomach of a swordfish. 

Location in Host - Gill cavity, sometimes mouth. 

Length - 2.6-38.6 cm, but usually less than 30.0 cm. Those found in Atlantic 

blue marlin are usually larger than those in white marlin, and those on Atlantic 

sailfish are smaller still. Small marlin remoras (24-26 mm SL) have been found 

on small sailfish (24 mm SL) (Cressey and Lachner 1970). 

Host Specificity - This remora is almost always found on sailfish and white 

marlin, but has occasionally been reported from other big game fishes. Cressey 

and Lachner (1970) suggested that marlin remoras only occur on billfishes, and 

records from sharks and other fishes are the result of contaminating other fishes 

with remoras from billfishes in mixed long-line catches on ship decks. 

Remora remora (Linnaeus) - shark remora 

This is the most common remora associating with offshore sharks, and 

occasionally big game fishes. It does not occur inshore. 

Name - The accepted common name of this fish is "remora". This is 

inappropriate and sometimes confusing because the name is also used as a 

general term for any species in the remora family. A better name is "shark 

remora" because it is almost always is found on sharks. 

255

256 


Diagnostic Characters - The body is short and rather robust. The sucking 

disk is comparatively large, extending to the middle of the pectoral fin, and has 

16-20 pairs of laminae. The caudal fin is deeply forked. 

Records - This remora almost always occurs on large sharks; rarely on Atlantic 

sailfish, white marlin, yellowfin tuna, sea turtles, boats, or free swimming in the 

western Atlantic and around the world. Up to 16 remoras may occur on the 

body or in the gills of a host. Whole "families" of various sizes may associate 

on 1 host. 


Life History - Planktonic larva shorter than 8 mm do not have sucking disks. 

The developing disk is behind the head in remoras 9.8 mm long. In 12 mm 

long fish, the disk is 1/2 on the head, and progresses further onto the head in 

larger fish. Larval and postlarval remoras do not have a caudal filament. 

Ecology - It is the most common offshore remora. This associate is limited to 

pelagic hosts, because it requires a swift passage of water over the gills and 

cannot survive in still waters or on inactive hosts. 

Parasites and Disease - The gillworm, Dioncus agassizi, occurs in the gills 

of cobia and shark remora from Woods Hole, Massachusetts, USA; and is also 

found in inshore remora and spearfish remora. The flukes Brachyphallus 

crenatus, Hemiurus montcellii (Linton), and Lecithochirium monticellii occurred 

in the intestine or stomach, and the tissue fluke, Torticaecum fenestratum 

(Linton) (USNPC 8352), encysted in the stomach wall of shark remoras, from 

Woods Hole, Massachusetts, USA. Echeneidocoelium indicum Simha and 

Pershad also occurred in the intestine of this host from the Indian Ocean. The 

larval tapeworms, Bothriocephalus sp., Callitetrarhynchus gracilis, Nybelinia 

bisculcata, Nybelinia robustum and tetraphyllid larvae were found in this remora 

from the western Atlantic; and N. robustum from Europe. The following 

copepods occur on shark remoras: Caligus elongatus from the northeast coast 

of the USA; Pennella filosa on this remora and its hosts worldwide; Perissopus 

oblongus (Wilson) (=Achtheinus dentatus Wilson), a parasite of sharks, with 

which the shark remora associates, that may have only accidentally occurred in 

this remora, from the south Atlantic; Lepeophtheirus crassus from the Indian 

Ocean, and on marlin remora from the Gulf of Mexico, white remora from the 

Indian Ocean, and whale remora from the Pacific. Striped goose barnacles 

occurred on a shark remora in the western Atlantic.


Cleaning Behavior - Smaller (45-165 mm SL) young remoras eat more 

parasitic copepods than larger (166-210 mm SL) old individuals. Parasitic 

copepods from sharks form an important part of their diet, but this has not been 

studied on remoras from big game fishes. Those on sharks feed on copepods 

from the body and gill chambers of their hosts, but usually avoid the fins of 

hosts, possibly due to fin movements. This may be why many copepods are 

isolated on the fins. 

Predation - This remora is sometimes eaten by tunas. 

Location in Host - Adults usually occur on the body, rarely in the gills. 

Smaller remoras are often found in the gill cavity. 


Host Specificity - It usually attaches to large offshore sharks, but can occur 

on a variety of offshore hosts. The relationships of this remora on sharks 

appear to be rather stable and long term, because copepods from their stomach 

match with those found on their host. 

Remorina albescens (Temminck and Schlegel) - white remora 

This small remora is usually found in the gills of manta rays, but can 

rarely be found in the gills of billfishes. 

Name - The accepted common name of this fish is "white suckerfish", but it is 

also called "white remora". 

Diagnostic Characters - The body is rather short, and whitish gray to 

brownish in color. The sucking disk is broad, as wide as 3/4 of the length of 

the disk, and has 12-14, usually 13-14, pairs of laminae. The caudal fin is 

truncated. 

Records - One occurred in a white marlin off the U.S. Atlantic coast (Goldstein 

pers. comm.). It is seldom seen swimming off a host. We have heard rumors 

of this remora being seen on sailfish and longbill spearfish, but these records 

could not be confirmed. 


Life History - Larval and postlarval remoras do not have a caudal filament. 

Females become gravid at a length of 20.6 cm SL. 

Ecology - Rarely seen near shore. 

Parasites and Disease - The copepod, Lepeophtheirus crassus, occurred on the 

body of this remora and shark remora from the Indian Ocean, on marlin remora 

from the Gulf of Mexico, and whale remora from the Pacific. 

257

258 


Associations - The specimen we recorded shared the host with an inshore 

remora that was attached to the head of the fish. This is one of the few records 

of different species of remoras occurring on the same host. 


Location in Host - Gill cavity and mouth, occasionally body. The white 

marlin specimen was in the gill chamber with the tail extending out of the 

chamber. 

Length - 4.7-30.0 cm, but usually less than 30.0 cm. The white remora we 

noted from a white marlin was approximately 30 cm long. 

Host Specificity - This remora prefers manta rays (Family Mobulidae), but is 

also found on sharks and bony fishes. White marlin is a new host. 

Naucrates ductor (Linnaeus) - pilotfish 

Pilotfish do not attach to the fishes they follow, and are the most loosely 

associated fish that we consider. They do pick parasites off their large hosts. 

Name - They are called "pilotfish" because them are so coordinated in copying 

the turns and movements of their larger hosts that they appeared to be guiding 

these fishes. 

Diagnostic Characters - The normal body coloration is 6-7 black bars against 

a light silvery background, but it temporarily turns silvery with blue blotches 

when disturbed. The lobes of the caudal fin have prominent white tips. The 

caudal peduncle has fleshy keels, and dorsal and ventral notches (peduncle 

grooves). 

Records - Pilotfish are always found associated with large sharks, rays, bony 

fishes including offshore big game fishes, sea turtles, ship hulls or driftwood. 


Life History - The larval forms are widespread in oceanic waters between the 

surface and 100 meters depth (epipelagic). Juveniles associate with seaweeds 

and jellyfish. They mature at a length of approximately 23 cm. 

Ecology - They are completely offshore and pelagic organisms. They feed on 

scraps (debris) from their host's meals, small fishes and invertebrates in the


water column, and their host's external parasites. Pilotfish can be caught on 

hook and line. 

Parasites and Disease - The gillworms, Ancyrocotyle bartschi Price, occurred 

on this fish from Puerto Rico; and Ancyrocotyle vallei (Parona and Perugia) 

from the Mediterranean. Neobenedenia melleni, a highly dangerous capsalid 

gillworm, was found on pilotfish from the New York Aquarium, but this was 

probably an accidental infection. The giant stomach fluke, Hirudinella ventricosa, 

occurred in this host from the Canary Islands and is found in many of the 

big game fishes with which pilotfish associate; Stephanostomum naucrotis 

Nagaty, from the Red Sea; and encysted metacercaria from the Mediterranean. 

Two larval tapeworms were found in pilotfish, Lacistorhynchus bulbifer, from 

the western Atlantic, which also occurs in a variety of big game fishes; and 

Nybelinia lingualis from Europe, which also occurs in skipjack tuna and 

swordfish. A spiny-headed worm, Serrasentis lamelliger (Diesing), from the 

Atlantic only occurs in this host. The copepod, Caligus productus, from Puerto 

Rico and the Indian Ocean; and the tissue-embedded copepod, Pennella filosa 

occurred on pilotfish, and both copepods also occur on a variety of big game 


Predation - Mediterranean spearfish, Tetrapturus belone Rafinesque, routinely 

eats pilotfish, and they have been found in the stomachs of striped marlin, Indo- 

Pacific sailfish and yellowfin tuna in the Pacific. Three pilotfish were found in 

the stomach of 1 of 1097 yellowfin tuna in a Pacific food-item study. 

Location in Host - Pilotfish swim ahead, or ahead and beside their hosts. 

Length - up to 70.0 cm, but usually less than 35.0 cm FL. 

Host Specificity - Pilotfish usually accompany large offshore sharks, but 

occasionally associate with other fishes, including big game fishes. 

Damage to Host - They eat scraps (debris) from their host's meals, but these 

materials might have been lost by the host even if the pilotfish was not present. 

They may nip or bite damaged or infected tissues of the host while "cleaning 

off" external parasites, but the tissue removal is probably beneficial to the host. 

Preparation for Study - Besides the standard preparations noted for fishes, 

their stomach contents should be quickly examined for parasites (before they are 

digested), or opened and quickly preserved in 10% formalin. The suggestion 

that they clean parasites from their hosts requires documentation from stomach 

contents. 

259

260 

OTHER DISEASES AND CONDITIONS 

Viruses 

Viral nervous necrosis (VNN) caused by striped jack nervous necrosis virus 

(SJNNV), in Family Nodaviridae, and yellowtail ascites virus (YAV), a birnavirus, 

cause serious losses in seed production of striped jack, Caranx vinctus 

(Jordan and Gilbert), in pen mariculture in Japan (Muroga, Nakai and Nishizawa 

1994). Similar viruses may exist in western Atlantic jacks, but have either not 

been expressed or not yet noted. Fishes held in confined culture conditions are 

observed closely and die-offs due to viral causes are more likely to be detected 

than in fishes in the wild. Viruses may have more opportunities in culture 

conditions for transmission than are possible in the wild. Future efforts at 

culture of big game fishes may create conditions favorable for virus expression 

and discovery. 

Bacteria 

A variety of pathogenic bacteria have killed and injured yellowtail and 

greater amberjack in intensive culture in Japan, including Nocardia seriolae 

Kudo, Hatai and Seino (commonly called N. kampachi Kariya et al.), Pasturella 

pisicida (Bein), Streptococcus sp. and Vibrio anguillarum (Bergman) (Kawahara 

et al. 1986). Descriptions of these bacteria, their known geographic range, 

epizootiology, diagnosis and pathological manifestations, are given by Plumb 

(1994). Dolphin are also being cultured in Hawaii, Japan and possibly in 

Puerto Rico in the future. As western Atlantic big game fishes are held, and 

particularly reared in captivity, primary pathogenic and opportunistic bacteria 

may attack these fishes. The possibility that these bacteria attack wild fishes 

remains unexplored. Bashirullah et al. (1995) reported that 6 of 17 sailfish 

examined for parasites had sloughing, bloody intestinal mucosa and hypothesized 

that it was caused by a bacterial infection. Bacterial isolation or histological 

confirmation was not made. 

Atlantic mackerel are often infected with "fish tuberculosis", Mycobacterium 

spp. Mycobacterium sp. was associated with visceral granulomas in an Atlantic 

mackerel collected off England (RTLA 4634) (Anonymous 1975b), and France 

(RTLA 4689, 4699) (Anonymous 1989), and other big game fishes may be 

affected. Plumb (1994) suggested that "all teleosts should be considered as 

possible hosts" for Mycobacterium spp. Skin granulomas in humans have been 

caused by mycobacteria, especially Mycobacterium marinum Aronson. Human 

infections are usually contracted in freshwater or saltwater swimming pools, 

tropical fish aquaria, or estuaries, but could be acquired from infected fishes. 

Human infections can be extremely resistant to treatment, and have even 

required surgery or amputation.


Multiple nodular or ulcerated lesions occurred in the skin and muscle of 4 

crevalle jacks from north Biscayne Bay off Miami, Florida, USA, (RTLA 2102- 

2105, 3105, 3375, 3403-5, 4133-4, 4148). The lesions were associated with 

inflammation due to infection by unidentified bacteria (Harshbarger 1979). 

Although catch-and-release fishing is unlikely to transmit bacteria from one 

fish to another, the transmission is possible, especially through cross 

contamination by fresh blood or material from active lesions. Cleaning gear 

between catches would be a prudent precaution to prevent possible cross 

infections. 

Tumors (neoplasms) 

Tumors in fishes can be enormous. The largest tumor reported was a 20 

kg lipoma described below. One tumor in a blackfin tuna was 1/9 of its body 

weight. Tumors, more technically called "neoplasms", are masses of new tissue 

which grow independently of their surrounding tissues. Tumors either rarely 

occur in big game fishes, or at least have been rarely reported. Two were found 

in one of the best studied big game fishes, Atlantic mackerel, an iridophoroma 

(neoplastic iridophores in the dermis of the skin) from France (RTLA 4691) 

(Anonymous 1989), and a pineal chondroma (tumor of the pineal organ with 

cartilage formation). Two lipomas (fatty tumors) partly bordered by skeletal 

muscle occurred in a little tunny caught off Miami Beach, Florida, USA (RTLA 

1922) (Harshbarger 1978); and 1 in the body wall of an Atlantic bonito from 

off Miami, Florida, USA, (RTLA 2359) (Harshbarger 1980). In bluefin tuna, 

there are records of a melanoma (RTLA 3919) an osteoma (RTLA 2745), a 

fibroma (RTLA 2744) and 2 lipomas (RTLA 2743 and 1518) (Harshbarger pers. 

comm.) The second lipoma (RTLA 1518) was a 20 kg growth on an 180 kg 

bluefin tuna off Massachusetts. 

An oval, 21x16 cm 

lipoma plus adjacent lobules 

occurred in a southern bluefin 

tuna (RTLA 2456) from 

Australia. A schwannoma 

(invasive malignant peripheral 

nerve sheath tumor) 

protruded from the flank of 

a blackfin tuna off Pensacola, 

Florida, USA (RTLA 

5391); and an osteoma (10 

cm ossified growth) was 

attached to the vertebral 

column of an albacore of 

the Pacific coast of the USA Caudal fin growths on a crevalle jack. 

(RTLA 1384). 

261

262 


Pseudotumors 

Non-neoplastic pseudotumors have also been rarely noted. Three crevalle 

jacks were collected with (1) skeletal muscle granulomas (scar tissue) that 

protruded into the visceral cavity (RTLA 1894), (2) multiple growths in the 

skeletal muscle (RTLA 1900), and (3) growths on the caudal fin (RTLA 1895) 

(see illustration on the previous page) (Harshbarger 1978). Each lesion was 

associated with unidentified protozoans. An abscess occurred in the ovary of 

one dolphin (RTLA 2360), and a nodular growth (muscle atrophy, necrosis and 

chronic inflammation due to traumatic injury) occurred in the muscle of a second 

dolphin (RTLA 2221) (Harshbarger 1980). A mass of granuloma tissue in the 

liver; enteritis of the gut and pyloric ceca with mucosal sloughing; and cecal, 

intestinal and vascular smooth muscle hyperplasia in response to unidentified 

trematodes occurred in an Atlantic sailfish (RTLA 2467) (Harshbarger 1989). 

These 5 non-neoplastic pseudotumors and 2 of the tumors above, came from 

off Miami Beach, Florida, USA, not because the conditions are particularly 

conducive to tumor formation in that region, but through the efforts of one 

interested observer. This is a good example of what can be accomplished by 

any motivated amateur scientist. Additionally, an Atlantic mackerel with 

hemosiderosis (numerous pigment granules in macrophage aggregates of the 

kidney and spleen) was found off Maryland, USA, (RTLA 4421) (Anonymous 

1981). An Atlantic mackerel with nodules in the gill lamellae (branchial 

nodules with epithelial hyperplasia and fibrosis) was found in the northwest 

Atlantic (RTLA 4472) (Anonymous 1989). A granuloma was found in an 

Atlantic mackerel from Europe; and hyperplasia (great increase in the number 

of normal cells arranged normally in a tissue) was reported from a captive chub 

mackerel in Japan. An inflammatory fibrosis was noted in dermis, hypodermis 

and adjacent skeletal muscle in a wild chub mackerel from Japan (RTLA 2038) 

(Harshbarger 1978). 

Anomalies 

Howse, Franks and Welford (1975) found unusual tissue connections 

between parts of cobia hearts (pericardial adhesions) that were probably caused 

by pericarditis. They suggested the abnormal condition occurred frequently 

among cobia in the northern Gulf of Mexico. Mesentery adhesions have also 

been reported in albacore from South Africa. 

Some billfishes may develop without bills or with malformed bills, but the 

reports of these possible anomalies are difficult to distinguish from bills that 

have been simply injured and healed. Reports suggest that fish with partial or 

missing bills appear to adequately feed and survive. A dolphin with a healed 

"broken back" (fracture of centrum of 2 vertebrae with exuberant callus 

formation) was noted off Miami Beach, Florida, USA (RTLA 2167) (Harshbarger 

1979).


Pugheaded (right) and normal (left) cobia. 

Franks (1995) reported the first cobia with a blunt shortened upper jaw and 

forehead, a condition often called "pugheaded", caught in the northern Gulf of 

Mexico in 1991. Feeding, growth and sexual maturity was not altered by this 

condition. One great barracuda with fused vertebrae has been reported on the 

Atlantic coast of the USA, and one bluefin tuna with vertebral anomalies was 

noted in the eastern Atlantic. 

Hermaphroditism 

Three skipjack tuna have been reported with both male and female sexual 

organs occurring in the same fish: one fish from India and 2 from Japan. We 

have received a report of a fourth fish, 51.3 mm and 2.7 km, from the 

Federated States of Micronesia (Itano pers. comm.). 

Stomach ulcers 

Evans and Wares (1972) found gastric ulcers in 73 of 563 striped marlin 

and in 33 of 151 Indo-Pacific sailfish off southern California and Mexico. 

Iversen and Kelley (1974) found gastric ulcers in the stomachs of 10 of 114 

Indo-Pacific blue marlin and 2 of 3 black marlin examined from 1967 to 1969 

at the Hawaiian International Billfish Tournament. Up to 6 ulcers that were 

wider than 10 mm and 50 smaller ulcers occurred in a stomach, ranged in width 

from 3-13 mm, had slightly raised edges, sharply demarcated margins, bases 

covered with brown shaggy material, and had an indurated texture. In tissue 

sections, the lesions could be seen to extend through the wall of the stomach 

obliterating the usual muscular layers. Jolley (1977) occasionally found stomach 

and intestinal ulcers in Atlantic sailfish. Bashirullah et al. (1995) reported small 

"petechiae and/or ulcers" in the empty stomachs and intestines of white marlin, 

Atlantic sailfish and swordfish. We rarely found stomach ulcers in Atlantic blue 

marlin, albacore (RTLA 6329) and in one longbill spearfish from Puerto Rico. 

All are new host records for this condition. Histological examination of the 

longbill spearfish revealed that the submucosa of the stomach was severely 

inflamed. Unidentified tissue fluke fragments and eggs 15-20 µm long were 

present in the outer layer of the muscularis or serosa but there was little 

263

264 


inflammation near the worms, suggesting that they may not have been the 

cause of the gastritis (Grizzle pers. comm.). Iversen and Kelley (1974) 

suggested that the spines of prey fishes caused the wounds, but thought 

nematodes, Maricostula histiophori, might also play some role as they occurred 

in many ulcers. The number and shape of the ulcers supports the spine-wounds 

theory, but the lesions appear to be complicated by bacterial infection. Fresh 

ulcers should be evaluated with standard bacterial isolation procedures. 

Pollution effects 

Most effects occur inshore in highly polluted areas thus only those big game 

fishes entering these regions are involved. "Examination of samples of 

developing embryos of Atlantic mackerel from the New York Bight plankton 

disclosed cytological abnormalities (in the form of disruption of the mitotic 

apparatus) and cytological abnormalities (ranging from stickiness and 

chromosome bridges to complete pulverization). Statistical correlations of high 

prevalences of chromosomal anomalies with degree of environmental 

contamination provided evidence for possible impacts on estuarine/coastal 

populations." (summarized from the literature by Sindermann 1993). As the 

general pollution of the world's oceans progresses, other big game fishes may 

be affected. The example of deteriorating coral reefs may be pertinent to this 

discussion. Coral reefs live in low nutrient environments and can be destroyed 

by what would be considered insignificant pollutants in other environments. 

Coral reefs, unfortunately, form close to shore and therefore are subject to 

human effects. Big game fish also rule a low nutrient realm, which is more 

isolated. Only minor, although global, pollution may cause anomalies in big 

game fishes. The health of big game fishes may prove to be a useful monitor 

of global change. 

Toxins 

Ciguatera fish poisoning 

Ciguatera fish poisoning of humans is caused when fishes contaminated by 

ciguatoxins are eaten. These toxins are found in the flesh but are actually more 

concentrated in the internal organs of affected fishes. Symptoms include nausea, 

headache, reversal of the sensation of hot and cold, tingling of the hands and 

face. Symptoms vary between cases possibly because different combinations of 

a variety of different toxins are involved. This is a seriously debilitating disease 

that is sometimes fatal. Recovery usually takes months and symptoms may 

recur months or years after the initial attack. The severity of the symptoms may 

depend on the levels of toxins in the consumed fish and the prior exposure of 

the victim. No satisfactory treatment is known and there are no tests to identify 

contaminated flesh. Cooking does not break down the toxin. Ciguatera occurs 

in tropical fish worldwide. In Puerto Rico, it is so common in great barracuda 

and many large jacks, including crevalle jacks and horse-eye jacks, that their


sale for human food is forbidden by law. The original sources of the toxins are 

substrate-dwelling dinoflagellates or algae. These toxins appear to accumulate 

up the food chain and particularly concentrate in piscivorous fishes associated 

with coral reefs. Offshore, big game fishes have seldom been implicated in 

ciguatera poisoning of humans, although greater amberjack have poisoned 

people in the Pacific. Ciguatera occurs so commonly in particular food fishes 

in different areas that these resources are not utilized. The disease is underreported 

because medical treatment is seldom sought. Many of our friends and 

colleagues have suffered ciguatera poisoning and research attempting to solve 

this problem has been conducted in our Department of Marine Sciences. 

Scombroid poisoning 

Scombroid poisoning is also called histamine poisoning because it is caused 

by high levels of histamine that are ingested from poorly processed or handled 

fish flesh, particularly in Scombridae and Scomberesocidae. Histamine is 

produced by the bacterial breakdown of histidine which is abundant in the 

muscle of these fishes. Bacteria convert the histidine to histamine in a chemical 

process that can take place even after the bacteria have been killed. Several 

species of bacteria can cause this reaction and are naturally found either on the 

gills or in the intestine of fishes and are transferred to the flesh during 

processing. Human symptoms include rash, flushing of the face, diarrhea, 

sweating and headache. Some victims report a metallic or peppery taste. The 

onset of symptoms is very soon after consuming contaminated fish. These 

symptoms help distinguish this intoxication from ciguatoxin. Prevention is 

accomplished by immediate freezing of the fish and keeping it very cold 

throughout processing. Proper handling during food preparation is also essential 

to prevent the growth of the bacteria on fish flesh. The disease is relatively 

mild and self-limiting and is usually responsive to antihistamine therapy. This 

intoxication occurs worldwide. Several countries keep records of the incidence 

and in the 1980's hundreds of cases were reported in the USA and England. 

Fewer cases are reported in Canada and other more northern regions possibly 

because processed fish stay colder in cold climates. This relatively mild disease 

is thought to occur far more often than it is reported. Fishes in Puerto Rico and 

the western north Atlantic likely to cause this disease are noted in the Host- 

Disease Checklists. 

Burnt Tuna 

Burnt tuna (yake niku=cooked meat in Japanese) was thought to be a 

breakdown of muscle tissue due to the struggle during capture. Actually, it is 

caused by enzyme action after death (calcium-activated proteases and their 

enhancement by high blood catecholamine levels). This condition causes bigeye 

tuna, yellowfin tuna and possibly other species to have flesh that is soft, exuding 

a clear fluid, and slightly sour in taste, instead of red, translucent and firm with 

a delicate flavor. In the Hawaiian sportfish and small-boat long-line fisheries, 

265

266 


half of the tuna trolled and 1/4 taken on long-lines were affected. The condition 

ranges from mild to severe and is found in 5-100% of the flesh of each fish. 

It prevents the use of tuna in raw-fish dishes by sport fishermen and keeps the 

small-boat fishermen from selling tuna in the Japanese market. Burnt fish may 

deteriorate more rapidly, but are otherwise safe to eat. The quality of cooked 

or canned meat is little affected by this condition. 

Mass Mortalities 

A mass mortality killed more than 20,000 tons of mackerel in the Arabian 

Sea during the second week of June 1957 (Jones 1962). This tonnage was 

nearly equal to the world catch of mackerel at that time. The event was never 

studied or explained. 

Strasburg (1959) found a mass mortality of larval frigate tuna south of Lanai 

in the Hawaiian Islands. The cause of death could not be determined, and there 

were no obvious signs of predation, disease or parasitism. It was probably 

caused by the passage of the larvae through an area with marked discontinuities 

in water temperatures. Larvae measured 2.2-8.2 mm TL. The rate of mortality 

increased with size, and larvae about 5 mm were the most affected. Jones 

(1963) suggested that bullet tuna must have also been killed in this mass 

mortality. A similar, sudden mass mortality of shrimps has been observed from 

a submersible. 

Health (1992) suggested that parasites may show developmental stage 

specificity for larval or young fishes and may cause short duration mass 

mortalities in a population. We are not aware of any practical evidence 

supporting this interesting suggestion.

DISCUSSION 

Parasites are a normal part of the biology of their hosts. Usually, parasites 

and hosts have reached an evolutionary point where the host can support the 

parasite without the frequent or premature death of the host. The nature of the 

host-parasite relationship can also tell us much about the host, often including 

its recent and long-term diet and behavior and, occasionally, its evolutionary 

history. For example, parasites tell us that humans were originally tropical 

creatures, because our parasite fauna in this region is more diverse and rich. 

HARM CAUSED BY PARASITES 

The first question everyone asks about parasites is "Can they hurt me?" 

and the second is "Do they hurt the fish?" That is why this information was 

provided for every parasite in the text when it was available, and is discussed 

below. 

Fish Diseases in Humans 

Some big game fish roundworms can infect people [see Nematoda 

(Roundworms)]; fish tuberculosis, ciguatera poisoning and scombroid 

poisoning occasionally injure or kill humans (see Other Diseases and 

Conditions). A few, large tropical big game fishes (jacks and great barracuda) 

should never be eaten, but, in general, properly prepared big game fishes are 

safe, wholesome and delicious. 

Damage to Hosts 

Parasite damage to big game fishes is probably much greater than noted in 

our parasite summaries. Much of the damage occurs in the open ocean where 

it is not recorded. The hosts that can be examined are largely caught by hook 

and line, and this technique selects healthy fishes. Injured or damaged fishes 

are quickly eaten by other fishes or sink out of sight. Scientists were recently 

surprised to learn that the surface waters of the world's oceans are filled with 

viruses. No one had bothered to look before. We know nothing about the 

ecologically important disease processes at work in the open ocean. 

All parasites can damage their host if they are present in sufficient 

numbers. We believe that this situation does not occur often, but we cannot be 

certain. All parasites are like tax collectors, they are always irritatingly present, 

and they always take their cut. Every fish is smaller, takes longer to grow, or is 

sometimes a little slower and gets eaten, because of parasites. This may seem 

minor, but it adds up to trillions of kilos of fishes and potentially billions of 

dollars worth of game and commercial fish value lost each year due to routine 


267

268 


TRANSMISSION OF PARASITES 

Offshore game fishes are usually widely dispersed, as any frustrated 

fishermen can attest! This big sea, few fishes, and little contact between hosts, 

is an enormous problem for tiny parasites and pathological microbes. All 

parasites have complex strategies to get from one host to another, the odds 

against the parasite in the open ocean are even higher than in almost any other 

environment. The tricks big game fish parasites use are incredible. 

Unfortunately, we have discovered very few of them. Some of the more simple 

tricks include releasing offspring when the hosts breed or congregate and 

dropping eggs when the adult host breeds in the inshore nursery area, with the 

eggs timed to hatch when the new generation of host is maturing. 

Parasite Transfer From Associated Fishes 

Pilotfish and remoras pick parasites off their large, big game fish associates, 

but they are also parasitized by some of the same gillworms and copepods as 

their hosts. Thus they also serve as "parasite taxis" carrying parasites between 

hosts, among host species, or from inshore to offshore localities. 

These associates also share many of the internal parasites of their hosts. 

Some of these can be transmitted to big game fishes when they occasionally eat 

their fish associates. These parasites can be easily transmitted to associates that 

consume fecal material of big game fishes. 

These fish associates also greatly benefit the shared parasites by "expanding 

the playing field". One big game fish carrying 5 remoras and surrounded by 4 

pilotfish provides 10 times the potential hosts for a parasite. Just by providing 

many more hosts in the open ocean, and thus more infective units, these 

associates may make parasitism of the sport fish much more likely. 

Prey to Predator Transfer 

Several of the isopod species reported on big game fishes have obviously 

escaped from their normal prey hosts as they were eaten, and reattached to their 

false host predators. The little we know about the life cycles of a few 

gillworms, copepods and possibly flukes suggests that they may occur on inshore 

or prey hosts as immatures, transfer to predators that consume their intermediary 

or decoy host, and mature as adults in the predator. This system could be 

widespread among big game fish parasites. Apparently it is employed by many 

of the tissue flukes. It allows the parasite to use more common, slower, or 

available prey fish and squid hosts as a stepping stone to the appropriate, but 

less available, final host.

DISCUSSION 

International Transfer of Exotic Parasites 

International commerce of big game fishes is largely confined to frozen 

carcasses or cooked and canned flesh. Cooking or adequate freezing kills 

parasites and minimizes the risk of their transmission. However, 2 new trends 

threaten the isolation of big game fish parasites: (1) air shipping of fresh tunas 

and other fishes, (2) aquaculture and subsequent shipping of live brood stock or 

young fish. 

The protozoan, Hexacapsula neothunni Arai and Matsumoto, was 

described from the Tokyo Fish Market, it causes a disease called "jelly-meat" 

that dissolves muscle of albacore, bigeye tuna and yellowfin tuna. This parasite 

has only been found in the Banda Sea (between New Guinea and Borneo) and 

in the Solomon Sea (east of New Guinea). It occurs most frequently in fish 30- 

45 kg and from June to August. One of every 200 fish is affected and jellymeat 

is found in both red and white meat. This is only one among many 

examples of devastatingly dangerous parasites with limited geographic range that 

could easily be introduced with air shipped fresh big game fishes. All that is 

necessary is the careless disposal of contaminated flesh into the environment. 

Something as simple as washing off a cleaning table and allowing the waste 

water to run into the ocean could start this pest. 

Greater amberjack fry were taken from the west coast of the USA, shipped 

to Hawaii, Hong Kong, Okinawa, and the main islands of Japan. A dangerous 

capsalid gillworm, Neobenedenia girellae (Hargis), was introduced with this 

host. It has not only caused disease problems in these cultured fishes, but has 

also injured wild fishes in these localities. Importing exotic amberjacks 

(resulting in the establishment of this exotic parasite) was unnecessary because 

all of these localities have their own local stocks of greater amberjack. 

Two species of blood flukes, Paradeontacylix spp., cause mass mortalities 

of cultured greater amberjack and yellowtail in Japan. These parasites were 

introduced in the 1980s from China and Hong Kong along with greater 

amberjack fry and quickly became a major problem. Our own Atlantic blood 

fluke in this genus from greater amberjack could also be a problem in its 

mariculture. 

Many other dangerous parasites and microbial agents could be just as easily 

be spread. See Bunkley-Williams and Williams (1995) for methods to avoid 

introducing parasites. 

Catch-and-Release Transfer 

Catch-and-release of big game fish is an honorable and ecologically friendly 

concept. However, it may also be a way to spread "the plague" among our 

favorite fishes, if we are not careful. The poor fish has been punctured in the 

mouth with a hook, abraded against the side of the boat, possibly gaffed and 

tangled in lines. If any or all of this equipment is contaminated with disease 

organisms from a previous catch, then it is practically inoculated into these 

269

270 


wounds. Parasites knocked off one fish and laying on the deck are perfectly 

happy to jump on the next host they encounter. 

A caught-and-released fish is about as "stressed out" as it is will ever be 

and still stay alive. Any weakened host is a bonus for a parasite or microbial 

disease. Whatever resistance or immunity the host may have possessed is lost 

or greatly reduced during the recovery period following release. The 

"incubation period" of diseases may be shorter than the recovery period of the 

host. Thus a contaminated fish may contract a fatal disease before it can 

recover from being caught. Most immediate losses of freshwater game fish 

following catch-and-release tournaments are caused by improper handling, but 

most longer-term losses are due to disease complications. The fate of released 

big game fishes has not been explored, but probably is similar to what occurs in 

freshwater game fishes. 

We are not suggesting that catch-and-release program do not work. We 

are merely pointing out that some of these fishes perish. At this point no 

studies have been conducted to determine how many die. What you can do to 

increase the likelihood that your caught-and-released fish will survive is to 

handle it as gently and briefly as possible, and to clean your handling gear and 

deck between catches to prevent any possible cross contamination of diseases 

between fishes. 

USEFULNESS OF PARASITES 

Environmental Indicators 

Parasites of big game fishes may be good indicators of changing 

environmental conditions. Many parasites have an enormous reproductive 

capacity or amazing techniques to find enough hosts to survive. Minor changes 

in the environment may either favor survival or death of parasite reproduction 

and/or disrupt the methods of host finding. These effects would be magnified 

by the system and expressed by either many more parasites per host or many 

fewer (see cero in Host Summaries). Minor environmental changes may cause 

major changes in the abundances of many parasites. Parasite levels may be a 

highly sensitive barometer of changes in the environment of big game fishes. 

Biological Tags 

Parasites have been used as biological tags to distinguish populations of 

fishes including some big game fishes. Those parasites with potential as 

biological tags are discussed in the individual parasite summaries in the text. 

Greater basic knowledge about big game fish parasites and their life cycles may 

provide more tools for fishery biologists to elucidate population movements 

and other population parameters of big game fishes. We are cooperating in an 

international effort to establish and characterize parasite biological tags for big 

game fish.

DISCUSSION 

Parasites as indicators of host relationships 

Cobia and remoras are assumed to be closely related because of similarities 

in shapes and colors of developmental stages and adults. Johnson (1984) 

suggested that cobia were more closely related to dolphins than remoras on the 

basis of their bone structure. The parasites of these species suggest that cobia 

are most similar to remoras, and have little similarities to dolphin. Gillworms 

are the most host specific of any parasites. Thus, the occurrence of Dioncus 

agassizi on cobia and 3 species of remoras seems indicative of a close 

relationship. The adult of Serrasentis sagittifer is host specific to cobia, thus an 

accidental record from inshore remora is interesting. Flukes are not usually 

host specific, thus are less useful in comparisons. However, cobia share their 

western Atlantic flukes with no other big game fishes, but one also occurs in 

inshore remora. Tuxophorus caligodes also occurs on inshore remora, possibly 

other remoras, and jacks. Lernaeenicus longiventris rarely occurs on cobia, but 

it is more typical of dolphin and jacks. Thus, the parasites shared by cobia and 

remoras suggest that they belong in a common family. However, only one 

species of parasite, Dioncus agassizi, would be family specific to a joined 

family, and a high percentage of cobia parasites are unique to this host. The 

isolation suggested by its parasites makes the cobia better suited to remain in its 

own family. On the basis of its parasites, cobia seems to be a remnant of an old 

family that has lost all of its species but one. 

Parasites of members of the fish genus Caranx are all but identical and 

interchangeable. There is some interchange or alternation among a few species 

but the host lists are essentially the same. This may explain some of the 

problems in separating species in the genus Caranx. The parasites suggest that 

none of the host species are very well separated. Some ichthyologists divide this 

genus of fishes into several genera, but the parasites of these fishes argues 

against such action. 

The parasites of jacks in genus Caranx are inshore denizens. The few 

parasites they share with scombrids have little specificity and are only found in 

those scombrids that spend considerable time nearshore. This parasite pattern 

may be due to habitat specificity more than host preference. Only Caligus 

robustus infects both jacks and offshore scombrids. Those with low host 

specificity include Caligus bonito which occurs on inshore and offshore 

scombrids and we found it on horse-eye jack. Lernaeenicus longiventrus prefers 

crevalle jack and dolphins, but is also found on inshore scombrids and cobia. 

It is one of the few parasites sharing Spanish mackerels and jacks. 

Amberjacks have more parasites in common with scombrids than with other 

jacks. Brachiella thynni prefers scombrids, but is also found on greater 

amberjack and we found it on dolphin. Koellikeria bipartita only occurs on 

greater amberjack and large tunas, another species of tissue fluke occurs on 

tunas and Pacific amberjacks, and 3 more occur on tunas, Pacific amberjacks 

and skipjack tuna. The copepod may be relatively non-host specific, but tissue 

271

272 


flukes select specific hosts. Amberjacks seem to have some odd relationship to 

tunas. Amberjacks are obviously not related to scombrids, thus similarities in 

habitat and behavior must be responsible for their shared parasite fauna. 

Dolphin share 7 non-specific parasites with scombrids, jacks and billfishes. 

Otherwise they have no parasites in common with these or any other big game 


Caligus productus rarely, and possibly accidentally, occurs on great 

barracuda. It prefers scombrids, but also occurs on billfishes. This tenuous, 

almost non-existent, overlap with other big game fishes and 2 non-host specific 

protozoans are all that connect great barracuda with fishes other than barracudas. 

Great barracuda parasites suggest that this host is almost completely isolated 

from other fishes. 

Chub and Atlantic mackerel are in the same genus and should be rather 

similar. They have 3 genus-specific parasites in common, but of 90 species of 

parasites found on these 2 hosts, only 14 occur on both. 

The scombrid family has 8 family-specific parasites, while their separate 

tribes have 25 (tunas), 7 (Spanish mackerels), 7 (little tunas), 5 (mackerels), 

and 1 (bonitos) tribe-specific parasites. This suggests that tunas form a strong 

tribe but the other tribes are less cohesive. 

King mackerel occur a bit further offshore than the 3 other western 

Atlantic Spanish mackerels. Their parasites differ a bit from the other 3 which 

have almost identical parasites. King mackerel have a similar, but different 

species of nasal copepod, and lack a second species of nasal copepod that is 

found in all the others. King mackerel have a spiny-headed worm, and a pitdwelling 

copepod that are more typical of offshore hosts. They lack the genusspecific 

isopod and tissue fluke that is shared by the 3 more inshore Spanish 

mackerels. 

The parasites of the western Atlantic Spanish mackerels are otherwise all 

but identical differing only by interchanges of similar species, missing genusspecific 

species that would probably be found with more examinations, or 

slightly different non-host-specific parasites. They share 11 genus-specific 

parasites and 2 almost genus-specific ones, but have only 1 species-specific 

parasite. They have no family-specific parasites, the only group in the 

scombrids to be so isolated. In contrast, the similarly isolated inshore jacks 

(genus Caranx) have 6 family-specific parasites. This suggests that Spanish 

mackerels are the most isolated, and possibly youngest, scombrids. They are 

more separated from each other in nearshore habitats, yet their parasites are 

more uniform than their more wide-ranging relatives, just the opposite of what 

would be predicted. 

The arrangements of parasites in these Spanish mackerels is very similar to 

what is found in the Caranx spp. of the jacks. Possibly the nearshore habitat 

was the last habitat occupied by both jacks and scombrids. The nearshore jacks 

and scombrids would be expected to be the youngest of their families. Each 

time the continents joined and separated this habitat was disrupted, contracted 

or expanded. The more geologically recent ice ages lowered sea level and 

similarly disrupted the nearshore, but not the offshore and oceanic habitats. In

DISCUSSION 

comparison, the offshore and oceanic jacks have enjoyed relatively stable 

habitats. 

The inshore jacks (Caranx spp.) and inshore scombrids (Scomberomorus 

spp.) have shared identical habitats, foods, conditions and histories. If 

environmental conditions determined parasite faunas, then their faunas would be 

identical. Under the same conditions, inshore jacks and scombrids developed 

parallel, but completely different parasite faunas. This is a powerful argument 

for the use of parasite faunas in analyzing host relationships (phylogeny). 

Wahoo is an unusual fish that is difficult to place within the scombrid tribes. 

It shares parasites with most other scombrids. Morphologically, wahoo 

resemble Spanish mackerels, and that is where they are usually placed. They 

share 8 parasites, including one almost host-specific gillworm, with little tunas, 

but only have 3 parasite species in common with Spanish mackerels. 

There are 5 species of copepods in the genus Gloiopotes, 1 is found on 

Atlantic sailfish, 1 on wahoo, 1 on other Atlantic billfishes and 2 species on 

Indo-Pacific billfishes. The cosmopolitan distribution of the wahoo and its 

Gloiopotes copepod contrasts with the other 2 Gloiopotes species that are 

isolated in the Atlantic, and the 2 species restricted to the Indo-Pacific. This 

could indicate that (1) the wahoo association is older, (2) wahoo migrate more 

frequently between the Atlantic and Indo-Pacific, or (3) the various billfish 

species in the Atlantic and Pacific are isolated in each realm and migrate so little 

that separate species of copepods (and host fishes) arose. In Indo-Pacific 

collections identified by Cressey (1967), 16 Indo-Pacific sailfish were parasitized 

by 4 G. huttoni and 12 G. watsoni (1:3 preference); and 22 striped marlin by 

14 G. huttoni and 8 G. watsoni (about a 2:1 preference). The preferred host is 

probably the one on which each parasite originally speciated. The mix of hosts 

and copepods in the Indo-Pacific also suggests that these parasites are older than 

the more host-conservative, and probably more recent, Atlantic species. 

Bonitos have physical features that share characteristics with at least 2 other 

tribes of scombrids. This group of fishes has provided one of the major 

arguments for not splitting up the scombrids, since bonitos are intermediate 

between 2 of the proposed tribes. The parasites of the Atlantic bonito provide 

little definitive evidence for this question. This fish shares its parasites with 

many of the other scombrid tribes. It has no distinct, unique parasite fauna. 

Only one of its parasites could be considered almost tribe specific. Thus there 

is no strong parasitological evidence to join bonitos with any particular scombrid 

tribe, and little to suggest that bonitos form their own cohesive tribe. The best, 

if weak, evidence suggests similarities between bonitos (Sardini) and little tunas 

(Katsuwonini). Atlantic bonito and little tunny only share 1 copepod, 

Ceratacolax euthynni, but it is found nowhere else. One nasal copepod, 

Unicolax collateralis, has been found in most bonitos and little tunas around the 

world. Oddly, it has not been found in the Atlantic bonito. 

Frigate tuna and skipjack tuna are in the little tuna tribe, but have rather few 

parasites in common. Their roundworms are interestingly quite similar. Two 

genera of these worms, Ctenascarophis and Prospinitectus, have only 2 species 

273

274 


each. Frigate tuna harbor C. gastricus and P. mollis; and skipjack tuna the 

"sister" species, C. lesteri and P.exiguus. 

None of the parasites of skipjack tuna are tribe specific to little tunas. The 

other western Atlantic little tunas have 1-5 tribe-specific parasites each. If 

skipjack tuna was placed in the tuna tribe, it would have 9 "tribe-specific" 

parasites. Gillworms often show more specificity than other parasites. 

Skipjack tuna and bluefin tuna are both infected by a gillworm, Hexostoma 

grossum; but skipjack tuna does not share any gillworms with the other little 

tunas. Tissue flukes are host selective since they only develop in specific 

fishes. Four species of tissue flukes occur only in skipjack tuna and bluefin 

tuna (another 5 species occur only in skipjack tuna, and 2-3 other tunas). Nasal 

copepods are almost as specific as gillworms. The 3 other species of western 

Atlantic little tunas are all infected by 2 species, Unicolax collateralis and U. 

mycterobius; but these do not occur in skipjack tuna. The fluke, Syncoelium 

filiferum, has little specificity, but occurs in skipjack tuna and albacore. 

Parasitologically, skipjack tuna is more closely related to tunas (Tribe 

Thunnini), than to little tunas (Tribe Katsuwonini). 

We found 25 parasite species which were tribe specific to tunas. Bigeye 

tuna had 48 species of parasites but almost half (22) were tribe specific. This 

is almost all of the tribe-specific parasites. Yellowfin tuna had the next highest 

with 17 specific, out of 80 total; while the others had many fewer (bluefin 6/70, 

albacore 6/39, blackfin tuna 3/9). The preponderance of shared parasites on 1 

host species might indicate that this fish is evoluntionarily older giving these 

parasites more time to colonize it, and to develop into tribe-specific parasites. 

The description of the host-specific gillworms, Tristoma adcoccineum and 

T. adintegrum, from Pacific swordfish by Yamaguti (1968) is the only strong 

parasite evidence suggesting that Atlantic and Indo-Pacific swordfish might 

represent different species. We suspect that the swordfish are the same, and 

that these Pacific gillworms may be found to be synonyms of the Atlantic, 

Tristoma coccineum and T. integrum. 

The western Atlantic billfishes have relatively few parasites and very few 

host specific ones. This could mean that they are relatively young 

evolutionarily compared to other big game fishes, and especially swordfish.

HOST SUMMARIES AND 

HOST-DISEASE CHECKLISTS 

We present a short summary of basic biological information of each host 

and include a "Name" category where we attempt to list other scientific names 

that have been used for the host when parasites were mentioned. Some of these 

are real synonyms of the fish host (those with authors), but many are mistaken 

names, wrong names or even made-up names that exist no where else. Most of 

the improper names appeared in the fish-parasite literature without authors. 

Some of the improper names are the correct scientific names of existing fishes 

that were only applied erroneously to a parasite host. Older synonyms, or 

synonyms that have never been used to refer to parasites, are not included; thus 

we are not providing complete synonym lists for each host species. We also 

include a short synthesis of the parasites found on each host with comments on 

the phylogenetic relationships among the hosts as indicated by their parasites. 

Parasites may only be an indirect indicator of their host's phylogeny and 

taxonomic relationships, but they are a part of the puzzle and provide additional 

biological evidence of relationships. 

In the disease lists, the parasites, diseases and conditions listed in the left 

column are those normally and 

routinely found in the western 

Checklist Abbreviations 

Atlantic. They are discussed individually 

in the text on the pages 

[acc] - accidental or contamination parasites 

indicated in bold face in the Index. 

[BSea] - Black Sea 

Parasites found on these hosts in 

[EAlt] - Eastern Atlantic 

geographic regions outside of the 

including Europe and Africa 

western Atlantic are noted in the 

[Eur] - Eastern Atlantic around Europe 

right column and are identified 

[fal] - false hosts 

with abbreviations that are noted in 

[Ind] - Indian ocean 

the box at right. Trivial parasites 

[IPac] - Indo-Pacific 

are similarly identified. The "right- 

(Indian and Pacific Oceans) 

column" parasites are included 

[MBS] - Mediterranean and Black Seas 

because some may eventually be 

[Med] - Mediterranean Sea only 

found in our area, and for compari- 

[Pac] - Pacific Ocean 

son with the western Atlantic fauna. 

They are not included in the Index. 

For this book, we chose billfishes, 

bonito, cobia, dolphins, little 

tunas, mackerels, Spanish mackerels, 

swordfish and tunas without 

exception. Only the larger barracuda 

and jacks, that are found offshore, 

were selected. Bluefish is a 

voracious sport fish, but a bit too 

[RSea] - Red Sea 

[SAfr] - South Africa 

[SWAfr] - South Africa and Atlantic off Africa 

[unc] - uncertain status 

(more study is necessary) 

[WAfr] - West Africa (Atlantic off Africa) 

[?] - questionable records 

* - larval parasites 

** - immature adult parasites 

275

276 


small for "big game", oddly does not visit Puerto Rico, and its parasites have 

already been well defined. Salmon (Salmonidae) are definitely big game 

fishes, but occur a bit too far north to be included and their parasites are well 

known. One or 2 sharks (Squaliformes) certainly qualify as big game fish, but 

their parasites are so different, that they would be better considered in a later, 

separate volume on sharks and rays. 

The name "Spanish mackerel" is confusing because it has been used as the 

name of a tribe of mackerels, as a western Atlantic species (Robins et al. 1991), 

and as a name for chub mackerel in the eastern Atlantic. We use the "Spanish 

mackerel" species name in the text because it is well established in the western 

Atlantic (particularly in the USA). The more appropriate common name would 

be "Atlantic Spanish mackerel" as we note in the Checklist. When we use the 

plural "Spanish mackerels" in the text, it refers to the tribe. 

Blue marlin and sailfish of Robins et al. (1991) are separated into Atlantic 

blue marlin and Indo-Pacific blue marlin, and Atlantic sailfish and Indo-Pacific 

sailfish. We use the Atlantic and Indo-Pacific names, but list the parasites of 

the Pacific cohorts as if they were worldwide species. Whether these fishes are 

separate species or not, their parasites are, interestingly, very similar. 

For the purpose of this discussion, we divided the little tunas and tunas into 

separate tribes to conveniently compare their parasites. This division is presented 

for expediency, and is not an attempt to revise their taxonomy (historically, 

proper host taxonomy has been utterly disregarded by fish parasitologists). 

General biological aspects of fishes are discussed under Pisces (Fishes). In 

this section we provide an illustration and a descriptive diagnosis of each host. 

The composite drawing, above, of a mythological fish illustrates the external 

anatomical features discussed in the diagnoses and "Location on Host" sections 

of the parasite descriptions. In a few hosts it was necessary to use characters 

of the gill rakers (finger-like projections on the side of the gill arches opposite 

the gill filaments) and liver to separate physically similar hosts. A morpholo-

HOST SUMMARIES AND HOST-DISEASE CHECKLISTS 

gical feature not easily illustrated is the lateral line which is a system of pressure 

or motion sensory organs on the side of the fish. It is usually obvious with a 

long row of pores through scales. "Muscle" refers to the large skeletal or body 

muscles. The "nose" or nares of both big game fishes and their bony fish 

associates consists of an incurrent pore (nare), a canal and olfactory rosette 

(nasal lamellae) and an excurrent nare. Pseudobranchs are patches of gillfilament-like 

lamellae found in the upper (dorsal) inner surface of the gill cover 

(operculum). "Gills" include the gill filaments, arches and rakers. "Gill 

chamber" includes all inner surfaces except gills and gill cover. "Body" 

includes skin and fins. 

Popular References - Anonymous (1994a), Goldstein (1988), Hammond and 

Cupka (1975), McClane (1974a,b). 

Reference - Fischer (1978). 

Order Perciformes, Family Rachycentridae - cobias 

Rachycentron canadum (Linnaeus) - cobia 

Name - Other common names include ling, lemonfish, black salmon, black 

kingfish, sergeant fish, crab-eater, runner and cabio. 

Diagnostic Characters - The spines of the first dorsal fin are isolated into 

individual finlets. The sides of body are marked with 2 sharply defined, 

narrow, horizontal, silvery bands. 

Geographic Range - Worldwide in tropical and warm-temperate waters. 

Food Habits - It eats crabs, squids and fishes. 

Ecology - Inshore and offshore 

Length - Maximum 200.0 cm, common to 110.0 cm. 

Weight - Maximum 61.5 kg, common 5.0-28.0 kg. 

Parasites - This fish has surprisingly few parasites (31) for such a widespread, 

abundant and relatively well examined host. However, more than 1/2 (17) are 

not reported from the western Atlantic, suggesting that fish from our region 

could be better examined. Further evidence is seen in its copepods, which have 

been well examined and are all reported from the western Atlantic. Most of the 

parasites known from cobia are either host specific (10), or possibly host 

specific (8) suggesting that cobia are not very closely related to any other fishes. 

Digenea (flukes) 

Mabiarama prevesiculata Laruea straightum [Ind] 

277

278 


Stephanostomum imparispine Lecithochirium canadus [Ind] 

Stephanostomum imparispine* Lecithochirium monticellii [?] 

Tormopsolus filiformis Lecithocladium jagannathi [Ind] 

Lepidapedon megalaspi [Pac] 

Paracryptogonimus morosovi [Pac] 

Phyllodistomum parukhini [RSea] 

Plerurus digitatus [Ind] 

Pseudolepidapedon pudens [acc] 

Sclerodistomum rachycentri [Ind] 

Stephanostomum cloacum [Ind] 

Stephanostomum dentatum** [fal] 

Stephanostomum microsomum [Ind] 

Stephanostomum pseudoditrematis[Ind] 

Tormopsolus spatulum [Ind] 

Didymozoidea (tissue flukes) 

Neometanematobothrioides rachycentri [Pac] 

Monogenea (gillworms) 

Dioncus agassizi Dioncus sp. [Pac] 

Cestoda (tapeworms) 

Nybelinia bisulcata* Callitetrarhynchus gracilis* [Med] 

Rhinebothrium flexile* Rhynchobothrium longispine* [unc] 

tetraphyllid* 

Nematoda (roundworms) 

Goezia pelagia 

Iheringascaris inquies 

Acanthocephala (spiny-headed worms) 

Serrasentis sagittifer Serrasentis nadakali [Ind] 

Copepoda (copepods) 

Lernaeenicus longiventris Caligus coryphaenae [Pac?] 

Parapetalus occidentalis Caligus haemulonis [?] 

Tuxophorus caligodes Euryphorus nordmanni [Pac] 

Lepeophtheirus plectropomi [Pac] 

Lernaeolophus hemirhamphi [acc] 

Lernaeolophus sultanus [acc] 

Cirripedia (barnacles) 

Conchoderma virgatum 

Condition 

heart connections (pericardial adhesions) 

Anomalies 

pugheaded


Family Carangidae - jacks 

Alectis ciliaris (Bloch) - African pompano 

Name - It has also been called "Atlantic threadfin", "Cuban jack", "pennantfish", 

Alectis crinitus (Mitchill), Blepharis crinitus, Carangoides ajax Snyder, 

and Hynnis cubensis Poey. 

Diagnostic Characters - The first dorsal fin is lacking in fish 17 cm FL and 

larger. The second dorsal and anal fins have relatively long bases. A row of 

large scutes on the caudal peduncle makes the tail rigid. Tips of the second 

dorsal and anal fins are falcate and long (extremely so in young). 

Geographic Range - Worldwide in tropical and temperate waters. 

Food Habits - It eats fishes and squids. 

Ecology - Inshore to oceanic. 

Length - Maximum possibly 150.0 cm, common to 90.0 cm FL. 

Weight - Maximum 23.0 kg, common to 8.0 kg. 

Parasites - Only 4 parasites have been reported from this host. The flukes and 

tapeworm have no host specificity, and the copepod is almost family specific to 

a variety of Indo-Pacific jacks. This host requires further examination. 

tetraphyllid* 


Lechithochirium sp. [Ind] 




Caligus constrictus [Ind] 

Caranx bartholomaei Cuvier - yellow jack 

Name - It is sometimes placed in genus Carangoides. 

Diagnostic Characters - The first dorsal fin has a relatively short base, and 

the second dorsal and anal fins have relatively long bases. A row of large 

scutes on the caudal peduncle makes the tail rigid. The back of the jaw extends 

to the front margin of the eye. 

279

280 



Food Habits - It eats primarily bottom-dwelling fishes. 


Length - Maximum possibly 100.0 cm (90.0 cm FL), common to 45.0 cm FL. 

One fish from Puerto Rico was 89.5 cm. 


Parasites - With only 15 species of parasites known, this host has obviously 

not been adequately examined. All its parasites are found in other species of 

Caranx. Seven are genus or family specific, 1 only found in jacks and scombrids, 

and 7 generalists with little host specificity (1 prefers jacks). 

Protozoa (protozoans) 

Haemogregarina bigemina 


Alcicornis carangis Genolopa brevicaecum [fal] 

Brachyphallus parvus Podocotyle chloroscombri [fal] 

Bucephalus varicus 

Ectenurus lepidus 

Pseudopecoeloides carangis 

Stephanostomum ditrematis 

Stephanostomum megacephalum 

Tergestia laticollis 


tetraphyllid* 


Caligus chorinemi 

Caligus robustus 

Caligus spinosus 

Lernanthropus giganteus 

Isopoda (isopods) 

Gnathia sp.*


Caranx crysos (Mitchill) - blue runner 

Name - It has also been called Caranx caballus and Paratractus caballus. It 

is replaced by the similar eastern Atlantic blue runner, Caranx fusus Geoffroy, 

in the eastern Atlantic; and green jack, Caranx caballus Günther, in the eastern 

Pacific. Some parasite records for this fish actually refer to these other species. 




to the middle of the eye. The tip of the pectoral fins extend back onto the line 

of tail scutes. 


Food Habits - It eats primarily fishes, but also shrimps, crabs, and other 

invertebrates. 


Length - Maximum possibly 68.0 cm (62.0 cm FL), common to 35.0 cm FL. 


Parasites - One is possibly host specific, 4 genus specific, 3 almost genus 

specific, 4 family specific, 1 only found in jacks and scombrids and 15 

generalists with little host specificity (1 prefers jacks). 




Alcicornis carangis Lecithochirium monticellii [?] 

Brachyphallus parvus 



Parahemiurus merus 





Allopyragraphorus incomparabilis Grubea cochlear [fal] 

Cemocotyle carangis 

281

282 



Callitetrarhynchus gracilis* 

Lacistorhynchus bulbifer* 

Otobothrium crenacolle* 

tetraphyllid* 


Gorgorhynchoides elongatus 

Ostracoda (seed shrimp) 

Vargula parasitica 


Caligus chorinemi Caligus haemulonis [?] 

Caligus elongatus 

Caligus longipedis 

Caligus robustus 

Lernaeenicus longiventris 


Pseudoeucanthus uniseriatus 


Cymothoa oestrum 

Excorollana tricornis 

Gnathia sp.* 

Nerocila lanceolata 

Caranx dentex (Bloch and Schneider) - white trevally 

Name - It is also called "guelly jack". This species is sometimes placed in 

genus Pseudocaranx. Records of this fish in the Indo-Pacific refer to guelly 

jack, Caranx guara (Bonnaterre). 



scutes on the caudal peduncle makes the tail rigid. The first dorsal fin is higher 

than the second dorsal fin. 

Geographic Range - Bermuda, southern Brazil and the eastern Atlantic.


Food Habits - It eats bottom fishes and invertebrates. 

Ecology - Inshore and offshore. 

Length - Maximum 80.0 cm (68.0 cm FL), common to 40.0 cm FL. 


Parasites - With 2 parasite records, this host has obviously not been adequately 

examined. Hysterothylacium aduncum will probably be found in additional 

jacks. 


tetraphyllid* 


Hysterothylacium aduncum 

Caranx hippos (Linnaeus) - crevalle jack 

Name - It is also called "common jack" and "horse crevalle". This fish replaced 

by the similar eastern Atlantic crevalle jack, Caranx sp., in the eastern Atlantic; 

and Pacific crevalle jack, Caranx caninus Günther, in the eastern Pacific. 

Parasite records for "crevalle jack" have been noted around the world, but this 

fish only occurs in the western Atlantic. 




to the back margin of the eye. The chest has only 1 small patch of scales. 

There is a black spot on the pectoral fins. 


Food Habits - It eats mostly fishes, but some shrimps and other invertebrates. 


Length - Maximum 101.0 cm, common to 60.0 cm FL. Old unconfirmed 

records of jacks 150.0 cm could have been this species. 

Weight - Maximum 26.0 kg, common 2.5-6.8 kg. Old, unconfirmed records 

of jacks 32.0 kg could have been this species. 

Parasites - This host has the most parasite species of any of the Caranx (36), 

probably because it has been examined more often. Nine are genus specific, 2 

283

284 


almost genus specific, 5 family specific, 1 only found in jacks and scombrids, 

and 19 generalists with little host specificity (2 prefer jacks). 

Bacteria 

bacterial infection 



Trypanosoma sp. 


Brachyphallus parvus Bucephalopsis gracilescens** [acc] 

Bucephalopsis arcuata 






Stephanostomum sentum 

Sterrhurus musculus 



Allopyragraphorus incomperabilis Dioncus remorae [acc] 


Cemocotyle noveboracensis 

Cemocotylella elongata 

Helixaxine winteri 

Protomicrocotyle mirabilis 



Dasyrhynchus giganteus* 

Eutetrarhynchus lineatus* 

Nybelinia bisulcata* 

tetraphyllid* 


Cucullanus carangis [unc] 


Gorgorhynchoides elongatus 


Caligus chorinemi Caligus praetextus [?] 

Caligus elongatus Anuretes heckelii [acc] 

Caligus longipedis Pandarus sinuatus [acc] 

Caligus robustus Tuxophorus caligodes [?] 

Caligus spinosus 

Holobomolochus crevalleus 

Lepeophtheirus edwardsi 


Lernanthropus giganteus




Livoneca ovalis 

Echeneis naucrates 

granulomas 

nodular lesions 

ciguatera 


Pisces (fishes) 

Neoplasms (tumors) 

Condition 

Caranx latus Agassiz - horse-eye jack 

Name - It is also called "goggle-eye". A similar species, dusky jack, Caranx 

sexfasciatus Quoy and Gaimard, occurs in the Indo-Pacific and is responsible for 

earlier parasite records of horse-eye jack in those regions. 




to the back margin of the eye. The chest is completely covered with scales. 

Geographic Range - Atlantic ocean. 

Food Habits - It eats fishes primarily but some shrimps and other invertebrates, 

including sea butterflies (pteropod mollusks). 


Length - Maximum 80.0 cm, common to 50.0 cm FL. 


Parasites - Six are genus specific, 2 almost genus specific, 3 family specific, 

and 8 generalists with little host specificity (1 prefers jacks). 


Alcicornis carangis Pseudopecoelus elongatus [fal] 

Brachyphallus parvus 


285

286 






Stephanostomum sentum 



Allopyragraphorus incomperabilis 


Cemocotyle noveboracensis 

Helixaxine winteri 



tetraphyllid* 


Caligus bonito 

Lernaeenicus longiventus 






Condition 

ciguatera 

Caranx lugubris Poey - black jack 



scutes on the caudal peduncle makes the tail rigid. The second dorsal and 

pectoral fins have long falcate tips.


Geographic Range - Worldwide in tropical, offshore waters. Around the 

West Indian islands and Bermuda, but well offshore in Florida, the Gulf of 

Mexico and Central and South America in the western Atlantic. 

Food Habits - It eats fishes primarily. 




Parasites - We examined 4 specimens of this host and only found a fluke and 

a larval cestode. This fish may simply harbor few parasites, but it has 

obviously not been examined sufficiently to be certain. 


Lecithochirium sp. Bucephalus carangis [Pac] 

Deretrema carangis [Pac] 

Elytrophallus mexicanus [Pac] 

Lecithochirium priacanthi [Pac] 


tetraphyllid* 


Cucullanus pulcherrimus [unc] 


Caligus longipedis [Pac] 

Lernaeolophus sultanus [acc] 

Caranx ruber (Bloch) - bar jack 

Name - It is sometimes placed in genus Carangoides. 



scutes on the caudal peduncle makes the tail rigid. A dark bar extends along 

the back and down the lower lobe of the caudal fin. 

Geographic Range - Western Atlantic from New Jersey, USA, to Venezuela 

including the West Indies and Bermuda, not in the northern Gulf of Mexico. It 

is the most abundant jack of this genus in the West Indies. 

Food Habits - It eats fishes primarily but some shrimps and other invertebrates. 


287

288 


Length - Maximum 50.0 cm, common to 40.0 cm FL. Larger fish have been 

reported in the Bahamas and Florida Keys, USA, but not confirmed. 


Parasites - Five are genus specific, 2 almost genus specific, 4 family specific, 1 

only found in jacks and scombrids, and 7 generalists with little host specificity 

(1 prefers jacks). 




Alcicornis carangis Lasiotocus truncatus [fal] 

Bucephalus varicus Opecoeloides brachyteleus [fal] 

Cetiotrema carangis Pinguitrema lerneri [fal] 

Ectenurus lepidus Prosorhynchus pacificus [fal] 





Allopyragraphorus incomperabilis 





tetraphyllid* 


Gorgorhynchus elongatus 


Caligus robustus Thysanote longimana [acc] 





Gnathia sp.* 

Elagatis bipinnulatus (Quoy and Gaimard) - rainbow runner 

Name - It is also called "runner", "rainbow yellowtail" or Seriola bipinnulata.



the second dorsal and anal fins have relatively long bases. A finlet with 2 rays 

follows both the second dorsal fin and the anal fin. It is a colorful fish. The 

sides are marked with 2 narrow, light blue or bluish-white stripes, with a 

broader, olive or yellowish stripe in between. 

Geographic Range - Worldwide in tropical and subtropical waters. 

Food Habits - It eats invertebrates and small fishes. 


Length - Maximum 107.0 cm FL (possibly 120.0 cm), common to 80.0 cm FL. 


Parasites - This fish has surprisingly few parasites (12) for such a widespread 

and abundant host. It has probably not been adequately examined, especially in 

the Atlantic with only 3 species of parasites known. 


Ectenurus lepidus Deretrema hawaiiense [Pac] 

Stephanostomum ditrematis Stephanostomum hispidum [Pac] 

Tergestia laticollis [Pac] 


Pseudomicrocotyle elagatis [Pac] 

Pseudomicrocotyle meservei [Ind] 


tetraphyllid* 


Terranova sp. [Pac] 


Caligus confusus [IPac] 

Caligus productus [Ind] 

Shiinoa elagatus [IPac] 


Isistius brasiliensis [Pac] 

Seriola dumerili (Risso) - greater amberjack 

Name - It has also been called S. lalandi(i), S. rhombica Smith, S. 

purpurescens Temminck and Schlegel, S. simplex Ramsey and Ogilby, and S. 

tapeinometapon Bleeker. 

289

290 



the second dorsal and anal fins have relatively long bases. The caudal peduncle 

has distinct grooves on the upper and lower surfaces. There are 11-19 gill 

rakers in fish longer than 20 cm FL. 


Food Habits - It eats fishes and invertebrates. 


Length - Maximum 183.0 cm (150.0 cm FL), common 70.0-110.0 cm FL. 


Parasites - Twenty of the 36 reported occurred in the western Atlantic. Ten 

of these 20 were also found in western Atlantic Caranx spp. Greater amberjack 

had more host specific parasites (5), but fewer genus specific (2) and family 

specific (4) ones, than were found in Caranx spp. Three were possibly host 

specific, 1 almost genus specific, 1 only found in jacks and scombrids, and 20 

generalists with little host specificity (2 prefer jacks). Amberjacks share more 

parasites with scombrids than with other genera of jacks, possibly due to their 

similar habitat and behavior (see Discussion). 

Bacteria 

Nocardia seriolae [Pac] 

Pasturella pisicida [Pac] 

Streptococcus sp. [Pac] 

Vibrio anguillarum [Pac] 




Brachyphallus parvus Paradeontacylix grandispinus [Pac] 

Bucephalus gorgon Paradeontacylix kampachi [Pac] 

Bucephalus varicus Prosorhynchus kahala [Pac] 

Ectenurus lepidus Stephanostomum hispidum [Pac] 

Lecithochirium microstomum Stephanostomum seriolae [Pac] 

Paradeontacylix sanguinicoloides 



Tormopsolus orientalis 


Koellikeria bipartita [EAtl,Med] 

Koellikeria micropterygis [Med] 

Neomtanematobothrioides periorbitalis [Pac] 

Patellokoellikeria seriolae [Pac] 


Allencotyla mcintoshi Aspinatrium kahala [Pac] 

Neobenedenia girellae [Pac] 


Bothriocephalus sp.* Nybelinia puntatissima* [WAfr] 

Dasyrhynchus giganteus*


Pseudogrillotia zerbiae* 

tetraphyllid* 


Hysterothylacium sp.* Hysterothylacium seriolae [Pac?] 




Brachiella thynni Brachiella elegans [acc] 

Lernaeenicus longiventris Caligus lalandei [WAfr] 

Lernanthropus giganteus Colobomatus lichiae [Med] 

Lernanthropus micropterygis [Med,RSea,Pac] 

Nesippus costatus* [Pac,acc] 

Condition 

scombroid poisoning (rare) ciguatera [Pac] 

Seriola fasciata (Bloch) - lesser amberjack 



has distinct grooves on the upper and lower surfaces. There are 23-26 gill 

rakers in fish longer than 20 cm FL. 

Geographic Range - This fish has a peculiarly disjunct distribution in Puerto 

Rico to Antigua, Aruba to Curacao, Cuba, and the Atlantic Gulf coasts of the 

USA and Mexico, and Bermuda. It rarely occurs in the eastern Atlantic. 

Food Habits - It eats squids and fishes. 

Ecology - Inshore to offshore. 

Length - Maximum 67.5 cm FL, common to 39.0 cm FL. 


Parasites - Its 2 parasites are also found in greater amberjack. 


Tormopsolus orientalis 


tetraphyllid* 

291

292 


Seriola rivoliana Cuvier - almaco jack 

Name - It is also called S. bovinoculata Smith, S. colburni Evermann and 

Clark, S. falcata Cuvier, and S. songora Smith. 



has distinct grooves on the upper and lower surfaces. The second dorsal fin is 

more than 3 times the height of the first dorsal fin. 

Geographic Range - Worldwide in tropical, subtropical and sometimes 

temperate waters. 

Food Habits - It eats fishes. 


Length - Maximum 97 cm FL; common to 55-80 cm FL. 

Weight - Maximum 60 kg, common 2.5-3.4 kg. 

Parasites - This fish has obviously not been adequately sampled. Its 3 

parasites are also found in greater amberjack. 


tetraphyllid* 




Lernanthropus micropterygis [Pac] 

Family Coryphaenidae – dolphins 

Coryphaena equiselis Linnaeus - pompano dolphin 

Name - Coryphaena equisetis Linnaeus is often used as a correction of the 

spelling of the original name. 

Diagnostic Characters - This fish has a single dorsal fin that extends from 

behind the eye almost to the caudal fin. The convex anal fin extends from the 

anus to almost the caudal fin. The pectoral fin is 1/2 of the length of the head. 

Geographic Range - Worldwide in tropical and subtropical offshore waters. 

Food Habits - It eats a variety of planktonic and sargassum invertebrates and 

small fishes. 

Ecology - Oceanic.


Length - Maximum 76.0 cm, common to 50.0 cm TL. 


Significance to Sport Fishing - This fish is usually too small to be taken on 

most gear used by big game fishermen. 

Parasites - Almost all of the few parasites known from this host occur on 

dolphin. 


Lecithochirium microstomum Stephanostomum dentatum** [fal] 

Dinurus tornatus 


Callitetrarhynchus gracilis* Nybelinia alloiotica* [WAfr] 

Tentacularia coryphaenae* trypanorhyncha* [EAtl] 

tetraphyllid* 


Hysterothylacium pelagicum Anisakis sp.* [Pac] 

Parascarophis galeata [unc] 


Caligus coryphaenae Alebion carchariae [acc] 

Lernaeenicus longiventris Euryphorus nordmanni [Pac] 

Lernaeenicus hemiramphi [Pac?] 


Isistius brasiliensis 

Coryphaena hippurus Linnaeus - dolphin 

Name - It is also called "common dolphin", "dolphinfish" (to distinguish 

marine mammals), "mahi mahi" (Hawaiian) and "dorado" (Spanish). 

Diagnostic Characters - This fish has a single dorsal fin that extends from 

above the eye almost to the caudal fin. The concave anal fin extends from the 

anus to almost the caudal fin. The pectoral fin is more than 1/2 of the length 

of the head. 

Geographic Range - Worldwide in tropical and warm-temperate waters. 

Food Habits - It eats fishes predominantly, but some crustaceans and squids. 

Ecology - Offshore and oceanic. 

293

294 


Length - Maximum 200.0 cm, common to 100.0 cm. Most fish are 1 year 

old, the largest are 4. 


Significance to Sport Fishing - This is the most commonly caught 

offshore big game fish in the western Atlantic and possibly worldwide. 

Parasites - Thirty of the 44 parasites reported for this host have been found 

in the western Atlantic. Seven are host specific, and 37 generalists with 

little host specificity (3 prefer dolphin). Dolphin share a few parasites with 

scombrids, billfishes and jacks, but appear to be largely isolated from other 

families of big game fishes. Seventeen species of larval tapeworms must 

indicate this fish has a diverse diet of tapeworm intermediate hosts, because 

the average number among big game fishes is less than 5. Some of its hostspecific 

parasites are only found in the Caribbean or parts of the Pacific, 

suggesting that this worldwide host may be separated into populations which 

have little communication. 


Haemogregarina bigemina Kudoa thyrsites [Pac] 

Kudoa sp. [Pac] 


Bathycotyle coryphaenae Dinurus hippuri [Ind] 

Dinurus tornatus Helicometrina nimia [Pac,acc] 

Hirudinella ventricosa 

Stephanostomum coryphaenae 

Tetrochetus coryphaenae 


tissue fluke* [Ind] 


Benedenia hendorffi [Pac] 

Neothroacocotyle coryphaenae [Pac] 

Tristomella laevis [acc] 


Callitetrarhynchus gracilis* Bothriocephalus janikii* [Pac,unc] 

Hepatoxylon trichiuri* Hepatoxylon stenocephala* [EAtl] 

Nybelinia bisulcata* Plerocercoides lonchophorus* [EAlt,unc] 

Otobothrium crenacolle* Pterobothrium acanthotruncatum* [Pac] 

Otobothrium dipsacum* Tetrarhynchus papillosus* [unc]

HOST SUMMARIES AND HOST-DISEASE CHECKLIST 

Pelichnibothrium speciosum* Tetrarhynchus* sp. [Pac,unc] 

Pterobothrium heteracanthum* 

Rhinebothrium flexile* 

Tentacularia coryphaenae* 

tetraphyllid* 


Hysterothylacium pelagicum Anisakis sp. [Pac] 

Ascaris sp.* [Pac,unc] 

Contracaecum sp. [Pac,unc] 

Hysterothylacium marinum [Pac] 


Rhadinorhynchus pristis 

Serrasentis sagittifer* 


Brachiella thynni Caligus balistae [fal] 

Caligus bonito Caligus curtus [?] 

Caligus coryphaenae Caligus patulus [acc?] 

Caligus productus Lernaeenicus hemiramphi [IPac] 

Caligus quadratus Lernaeenicus nodicornis [unc] 

Caligus wilsoni Pennella varians [Eur,unc] 

Charopinopsis quaternia 

Euryphorus nordmanni 


Pennella sp. 

Pseudocycnus appendiculatus 


Anilocra physodes [Med] 

Glossobius impressus [Pac?] 1 


Remora osteochir 

Glossobius auritus [Pac] 1 

Nerocila excisa [Pac] 1 



muscle nodules 

ovarian abscess 

Condition 

broken back head deformities(aquaria) [Pac] 

scombroid poisoning (frequent) 

1Occurred in the stomach of this host, probably parasitized a flyingfish. 

295

296 


Family Sphyraenidae - barracudas 

Sphyraena barracuda (Walbaum) - great barracuda 

Name - It is also called "giant barracuda" and just "barracuda". 

Diagnostic Characters - It is an elongate fish with relatively large, unequal 

vertical teeth in the upper and lower jaws. The 2 dorsal fins are separated by 

a distance greater than the combined lengths of their bases. The pelvic fin 

begins slightly ahead of the first dorsal fin. It usually has several to many inkyblack 

spots of various size on the posterior, lower sides of the body. 

Geographic Range - Tropical and subtropical western Atlantic from 

Massachusetts to southern Brazil. Also eastern Atlantic and western Indo- 

Pacific. 

Food Habits - It eats fishes predominantly, sometimes squids or octopus, 

occasionally shrimps. 

Ecology - Inshore to offshore, but not oceanic. 

Length - Maximum 200.0 cm (rumored to over 300.0 cm), common to 130.0 

cm. We have seen bigger fish than 2 m TL while snorkeling in the waters 

around Puerto Rico, but these fish were measuring us! It is rumored to be 

larger in the Caribbean (Where everything is bigger and better). 


Parasites - This is a well known, popular, worldwide fish that has been well 

examined, but does not have many parasites. Linton (1908) suggested that great 

barracuda in the Dry Tortugas, Florida, USA, and Bermuda had few parasites. 

This has been our observation as well. Eighteen of the 31 parasites reported for 

this host have been found in the western Atlantic. Nine are host specific, 1 

possibly host specific, 5 genus specific, 2 possibly genus specific, and 14 generalists 

with little host specificity. These parasites suggest that this host is almost 

completely isolated from other fishes. The high numbers of host-specific 

parasites (9) indicate that this is a relatively old fish evoluntionarily, and well 

established in its niche. The relatively low number of non-specific parasites (12) 

suggests that it has unique food habits, behavior and habitats. Some of its hostspecific 

parasites are only found in parts of the West Indies or the Pacific, 

indicating that this worldwide host may be separated into populations which have 

little communication. Eight species of tissue fluke occur in 5 other species of 

barracuda in the Mediterranean, Indian Ocean and Pacific. None of these tissue 

flukes has been found on great barracuda even though it occurs around the 

world. More than 600 great barracuda were eviscerated as part of another study


in La Parguera, Puerto Rico, but no tissue fluke lesions were observed in any 

of these fish. The absence of tissue flukes may also suggest that this fish is 

isolated from other species of barracuda. 



Trypanosoma sp. 


Bucephalopsis longicirrus Bucephalopsis attenuata [fal] 

Bucephalopsis longoviferus Bucephalus carangoides [Pac] 

Rhipidocotyle barracudae Bucephalus kaku [Pac] 

Claribulla longula [fal] 

Deretrema sphyraena [Pac] 

Hirudinella ventricosa [WAfr,acc] 

Lecithocladium excisum [fal] 

Neolepedapedon belizense [fal] 

Opegaster hawaiiensis [Pac] 

Plerurus digitatus [Pac] 

Plerurus sphyraenae [Pac] 

Prosorhynchus longicollis [Pac] 

Pseudopecoelus sphyraenae [Pac] 

Rhipidocotyle longleyi [fal] 

Stephanostomum ditrematis [fal] 

Sterrhurus musculus [fal] 

Sterrhurus sp. [Pac] 

Uterovesticulurus sphyraenae [Pac] 


Pseudochauhanea sphyraenae Vallisiopsis sphyraenae [Pac] 


Otobothrium dipsacum* 

Tentacularia sp.* 

tetraphyllid* 


Hysterothylacium marinum [Pac] 

Raphidascaris anchoviellae [unc] 


Caligus isonyx Caligus productus [IPac] 

Caligus lobodes 

Hatschekia amplicapa 

Lernaeolophus striatus 

Branchiura (fish lice) 

Argulus bicolor 


Cymothoa oestrum Alcirona krebsii [acc] 


Gnathia sp.* 

Rocinela signata 

297

298 


Naucrates ductor 

Phtheirichthys lineatus 

Remora brachyptera 


ciguatera 

vertebral 



Condition 

Anomalies 

Family Scombridae, Tribe Scombrini – mackerels 

Scomber japonicus Houttuyn - chub mackerel 

Name - It is sometimes called Pneumatophorus colias (Gmelin), P. grex, P. 

japonicus, and S. colias Gmelin. 

Diagnostic Characters - All scombrids have small finlets behind the dorsal 

and anal fins. The eye has an adipose eye lid. The space between the 2 dorsal 

fins is approximately equal to the length of the base of the first dorsal fin. It 

has dusky blotches on the lower sides and belly. This fish has a gas bladder. 

Geographic Range - Worldwide in temperate waters, including the 

Mediterranean. It is found from Nova Scotia, Canada, to Argentina, but is 

uncommon in the Gulf of Mexico and Caribbean, except in south Florida and 

the off the north central coast of South America. The western and eastern 

Atlantic populations appear to be relatively isolated. Parasites might be useful 

biological tags to measure the amount of contact. 

Food Habits - It eats small pelagic fishes such as anchovies, herrings, and 

silversides; and pelagic invertebrates. 




Parasites - Only 23 of the 75 parasites reported in this worldwide host have 

been found from the western Atlantic, suggesting that it has not been adequately 

examined in our region. Four parasites are host specific, 12 possibly host


specific; 7 genus specific, 1 possibly genus specific; 4 family specific; and 47 

generalists with little host specificity (1 prefers chub mackerel). 


Ceratomyxa inconstans [Pac] 

Goussia pneumatophori [Pac] 

Pseudoalatospora scombri [Pac] 


Coitocaecum extremum Aphallus tubarius [Med] 

Hirudinella ventricosa Aponurus laguncula [acc] 

Lecithocladium excisum Apocreadium misakiense [Pac] 

Neolepidapedon retrusum Cephalolepidapedon saba [Pac] 

Opechona orientalis Dinurus scombri [Pac] 

Tergestia laticollis Ectenurus lepidus [Med] 

Lecithaster gibbosus [Pac] 

Lecithochirium microstomum [Pac] 

Lepocreadium ghanense [WAfr] 

Lepocreadium scombri [Pac] 

Neopechona olssoni [Pac] 

Opechona acanthurus [EAtl] 

Opechona bacillaris [EAtl,Med,Pac] 

Opechona scombri [Pac] 

Tergestia acanthocephala [Pac] 


Didyymozoon sp. Allonematobothrioides baueri [Pac] 

Didymozoon longicolle Allonematobothrioides hirosaba 

[Pac] 

Nematobothrium scombri Allonematobothrioides scombri [Pac] 

Allopseudocolocyntrotrema sp. 

[SAfr] 

Didymocystis wedli [Pac] 

Nematobothrioides gomasabae [Pac] 

Nematobothrioides pneumatophori [SAfr] 

Nematobothrium filliforme [Pac,WAfr] 

Nematobothrium robustum [Pac] 


Grubea cochlear Gastrocotyle japonica [Pac] 

Kuhnia scombercolias Kuhnia sprostonae [Eur,IPac] 

Kuhnia scombri Microncotrematoides inversum [Pac] 

Pseudokuhnia minor [EAtl,IPac] 


Bothriocephalus sp.* Nybelinia sp.* [WAfr] 

Callitetrarhynchus gracilis* Nybelinia surmenicola* [Pac] 

Rhinebothrium flexile* pseudophyllid* [Pac] 

Rhinebothrium sp.* trypanorhynchid* [Pac,WAfr] 


tetraphyllid* 

299

300 



Anisakis sp.* Ascaris scombrorum* [Med,unc] 

Capillaria sp. [Pac] 

Contracaecum sp.* [Pac,WAfr] 

Hysterothylacium aduncum [Pac] 

Hysterothylacium fabri [Pac] 

Hysterothylacium saba [Pac] 

Oncophora melanocephala [WAfr] 

Phocanema sp.* [Pac,WAfr] 

Raphidascaris sp.* [Pac] 


Rhadinorhynchus pristis Bolbosoma sp. [Pac] 

Rhadinorhynchus cadenati [WAfr] 

Rhadinorhynchus japonicus [Pac] 

Rhadinorhynchus lintoni [Eur] 

Rhadinorhynchus seriolae [Pac] 


Caligus mutabilis Advena saba [Pac] 

Caligus pelamydis Lepeophtheirus dissimulatus [Pac] 

Clavellisa scombri Lernaeocera branchialis [Pac?] 

Pumiliopes capitulatus [Pac] 

Sarcotretes inflexus [Eur] 


Meinertia gaudichaudii [Pac] 


Petromyzon marinus 


inflammatory fibrosis [Pac] 

Condition 

scombroid poisoning (frequent) hermaphroditism [Pac] 

hyperplasia (in captivity) [Pac] 

malformed jaws (in captivity) [Pac] 

Scomber scombrus Linnaeus - Atlantic mackerel 

Name - It is often called "mackerel" and "common mackerel".


Diagnostic Characters - All scombrids have small finlets behind the dorsal and 

anal fins. The eye has an adipose eye lid. The space between the 2 dorsal fins 

is greater than the length of the base of either fin. It lacks dusky blotches on 

the lower sides and belly. This fish does not have a gas bladder. 

Geographic Range - Temperate and near-temperate waters of the North 

Atlantic Ocean, including the Mediterranean and Black Sea. In the western 

Atlantic, it is found from Cape Hatteras on the USA Atlantic coast into 

southern Canada. The western and eastern Atlantic populations do not appear 

to be isolated. Parasites might be useful biological tags to measure the amount 

of contact. 

Food Habits - It eats small, planktonic animals (copepods, Calanus sp., other 

invertebrates including squids, fish eggs and fry). 

Ecology - Offshore. This fish swims constantly as it lacks a swim bladder (air 

bladder) for buoyancy. 


Weight - Maximum 1.8 kg, common to 0.5 kg 

Parasites - Forty-six parasites have been reported from this host found 

throughout the Atlantic, however, only 20 have been found in the western 

Atlantic. Two parasites are host specific, 4 possibly host specific, 6 genus 

specific, and 1 family specific; and 33 generalists with little host specificity. 

Bacteria 

acid-fast bacilli [Eur] 

Tuberculosis [Eur] 

Mycobacterium sp. [Eur] 


Goussia clupearum Ceratomyxa parva [Eur,MBS] 

Haematractidium scombri Eimeria sp. [Eur] 

Kudoa histolytica Haemogregarina bigemina [?] 

Fungi (fungus) 

Ichthyophonus hoferi 


Bucephalopsis arcuata* Acanthocolpoides guevarai [WAfr] 

Lecithocladium excisum Brachyphallus crenatus [fal] 

Hemiurus appendiculatus [acc] 

Lecithaster confusus [fal] 

Lecithaster gibbosus [Eur] 

Lecithochirium caudiporum [BSea] 

Opechona bacillaris [Eur,Med] 

Opechona orientalis [Med] 

Opecoeloides vitellosus [?] 

Podocotyle simplex [acc] 

Wardula capitellata [Med] 


Atalastrophion sp. Halvorsenius exilis [Eur] 

Nematobothrium faciale [Eur] 

301

302 


Nematobothrium scombri [Eur,MBS] 

Paranematobothrium triplovitellatum [SAfr?] 


Grubea cochlear Kuhnia sprostonae [Eur,IPac] 

Kuhnia scombri Pseudokuhnia minor [EAtl,Med] 


Bothriocephalus sp.* Echeneibothrium sp.* [Eur] 

Bothriocephalus scorpii* Grillotia angeli [Eur,Med] 

Callitetrarhynchus gracilis* Rhynchobothrium longispine* [unc] 

Grillotia erinaceus* Tetrabothriorhynchus scombri* [unc] 

Lacistorhynchus bulbifer* Tetrarhynchobothrium sp.* [Eur] 


tetraphyllid* 


Hysterothylacium aduncum Anisakis simplex* [Eur} 

ascarid sp.* [Eur] 

Ascaris papilligera* [Med,unc] 

Contracaecum pedum* [Eur,unc] 

Contracaecum scombricum* [unc] 

Cystoopsis scomber [BSea,unc] 

Goezia sp.* [Eur] 

Hysterothylacium aduncum* [Eur] 

Oncophora melanocephala [Eur] 


Rhadinorhynchus pristis Bolbosoma vasculosum [Eur] 


Caligus elongatus Advena paradoxa [Eur,Med] 

Caligus pelamydis Caligus diaphanus [Eur,Med] 

Clavellisa scombri [Eur,Med] 

Lepeophtheirus pectoralis [Eur] 




gill nodules granuloma [Eur] 

hemangioma iridoporoma [Eur] 

hemosiderosis 

pineal chondroma 

Condition 

foreign object embeded intestinal lesion [Eur] 


Anomalies 

embryo-pollution (cytological) 

embryo-pollution (chromosomal)


Tribe Sardini - bonitos 

Sarda sarda (Bloch) - Atlantic bonito 

Name - This fish can only be canned as "bonito", not "tuna" in the USA ["tuna 

(bonito)" in Canada]. It has been called "common bonito" and Pelamys sarda. 


and anal fins. The first dorsal fin is much longer than the second dorsal. It has 

5- 11 dark, slightly oblique stripes running forward and downward on the back 

and upper sides. 

Geographic Range - Tropical and temperate coasts of the Atlantic Ocean, 

including the north central coast of South America, north coast of Yucatan, 

Mexico, northern Gulf of Mexico, southeast coast of South America, Atlantic 

coast of the USA, the Mediterranean and Black Sea. 

Food Habits - It eats small fishes, especially anchovies, herrings, cods, and 

scombrids. 




Parasites - This host has been well examined, but harbors only 27 parasite 

species, 20 of which have been found from the western Atlantic. Four are host 

specific, 1 almost tribe specific, 3 family specific, 1 shares this host and little 

tunny, and 18 generalists with little host specificity (1 prefers this host). These 

parasites do not suggest that bonitos form a cohesive tribe (see Discussion). 


Bucephalopsis arcuata Aponurus tschugunowi [BSea] 

Ectenurus lepidus Lecithochirium caudiporum [BSea] 

Hirudinella ventricosa Opecoelides vitellosus [?] 

Lecithochirium texanum 

Rhipidocotyle capitata 


Atalostrophion sardae 

Nematobothrium pelamydis 


Hexostoma lintoni Cabellerocotyla pelamydis [Med] 

Hexostoma euthynni [acc] 

Hexostoma thynni [Med] 

303

304 



Bothriocephalus sp.* Callitetrarhynchus gracilis* [WAfr] 

Grillotia erinaceus* Tetrarhynchus megabothrium* [unc] 

Lacistorhynchus bulbifer* Tetrarhynchus scomber-pelamys* [unc] 



tetraphyllid* 


Anisakis simplex* Ascaris appendiculata* [Med,unc] 



Caligus bonito Alebion glaber [acc] 

Caligus mutabilis Alebion gracilis [acc] 

Caligus pelamydis 

Ceratacolax euthynni 



Livoneca sp. [WAfr] 


lipoma 

Condition 

scombroid poisoning (rare) 

Tribe Katsuwonidae - little tunas 

Auxis rochei (Risso) - bullet tuna 

Name - Robins et al. (1991) suggested that the name "bullet mackerel" has 

been used incorrectly too long to change, but the "bullet" part is a recent name 

[1967]. This fish cannot be legally canned as "tuna" in the USA. It has also 

been called A. maru Kishinouye, and A. thynnoides Bleeker. 


and anal fins. The first and second dorsal fins are separated by a length 

approximately equal to the base of the first dorsal. There is a complicated


pattern of nearly vertical, dark bars on the posterior back that does not extend 

forward to the posterior end of the first dorsal or pectoral fins. 

Geographic Range - Worldwide in the tropics and subtropics, including the 


Food Habits - It eats small fishes, especially anchovies and herrings; crustaceans, 

especially megalops crab larvae and larval mantis shrimps; and squids. 

Ecology - Offshore and oceanic. It comes inshore around Caribbean islands. 

Length - Maximum 60.0 cm FL, common to 15.0-35.0 cm FL. 


Parasites - Such a widespread host should support more than 17 parasite 

species, and certainly more than the 7 found in the western Atlantic. The 56 

parasites known from the very similar frigate tuna suggest that bullet tuna have 

not been adequately examined. One parasite is genus specific, 1 tribe specific, 

1 almost tribe specific, 1 family specific, 1 shares this host with other little 

tunas and Atlantic bonito, and 1 this host with other little tunas and tunas, and 

11 generalists with little host specificity. 





Didymozoon sp. Didymozoon auxis [Med] 


Callitetrarhynchus gracilis* Tetrarhynchus scomber-rocheri* [unc] 

tetraphyllid* 


Ascaris papilligera* [Med,unc] 



Allorhadinorhynchus sp. [Pac] 

Bolbosoma sp. [Pac] 

Rhadinorhynchus sp. [Pac] 


Unicolax collateralis Caligus asymmetricus [IPac] 

Unicolax mycterobius Caligus coryphaenae [Pac] 

Caligus macarovi [Pac] 

Caligus pelamydis [Pac] 

Caligus productus [Pac] 

Condition 

scombroid poisoning (rare) mass mortality 

Auxis thazard (Lacepède) - frigate tuna 

Name - The name "frigate mackerel" has been used in the English fisheries 

literature since 1884, but this change is necessary, if controversial. This fish 

cannot be canned as "tuna" in the USA. It has also been called A. hira 

Kishinouye, and A. tapeinosoma Bleeker. 

305

306 



and anal fins. The first and second dorsal fins are separated by a length 

approximately equal to the base of the first dorsal. There is a complicated 

pattern of oblique, nearly horizonal, stripes on the posterior back that extends 

forward to the posterior end of the first dorsal and pectoral fins. 

Geographic Range - Possibly worldwide in the tropics and subtropics, but not 

in the Mediterranean. It occurs in abundance off the northern coast of South 

America. This fish was confused with bullet tuna in the western Atlantic until 

recently, thus its distribution has not been determined. 

Food Habits - It eats small fishes, especially anchovies and herrings; 

crustaceans, especially megalops crab larvae and larval mantis shrimps; and 

squids. 



Weight - Maximum 4.5 kg, common to 0.5-1.4 kg. 

Parasites - With 58 parasite species recorded, this host appears to have been 

well examined, except in the western Atlantic where only 9 have been found. 

Six parasites are host specific, 9 possibly host specific, 1 genus specific, 6 tribe 

specific, 2 almost tribe specific, 3 family specific, 1 occurs on this host and 

Spanish mackerels, 1 on bonitos and little tunas, and 2 on little tunas and tunas, 

and 27 generalists with little host specificity. 




Brachyphallus parvus Bucephalopsis gracilescens [Med] 

Lecithocladium excisum** Dinurus euthynni [Pac] 

Tergestia laticollis Dinurus scombri [Pac] 

Hirudinella ventricosa [Pac] 

Lecithochirium texanum [Pac] 

Lecithochirium keokea [Pac] 

Lecithochirium microstomum [Pac] 

Lobatozoum sp. [Pac] 

Phyllodistomum lancea [Pac]


Plerurus carangis [Pac] 

Plerurus kawakawa [Pac] 

Prosorhynchus sp. [Pac] 

Rhipidocotyle capitata [Pac] 

Rhipidocotyle capitata* [Pac] 

Rhipidocotyle pentagonum [Pac] 

Tergestia acanthogobii [Pac] 


Annulocystis auxis [Pac] 

Colocyntotrema auxis [Pac] 

Didymocystis wedli [Ind] 

Didymozoon auxis [Pac] 

Didymosphaera mirabilis [Pac] 

Lobatozoum multisacculatum [Pac] 

Metanematobothrium bivitellatum [Pac] 

Oesophagocystis sp. [Pac] 

Opepherotrema planum [Pac] 

Phacelotrema claviforme [Pac] 

Pseudocolocyntotrema yaito [Pac] 

Sicuotrema auxia [Pac] 


Allopseudaxine macrova [Pac] 

Caballerocotyla manteri [Pac] 

Hexostoma auxisi [Med] 

Hexostoma keokeo [Pac] 


Metapseudaxine ventrosicula [Pac] 

Pseudaxine sp. [Pac] 

Pseudaxine triangula [Pac] 


Callitetrarhynchus gracilis* Tetrarhynchus scomber-rocheri* [unc] 

tetraphyllid* 


Anisakis sp.* [Pac] 

Contracaecum sp.* [Pac] 

Ctenascarophis gastricus [Pac] 

Oncophora melanocephala [Med] 

Prospinitectus mollis [Pac] 


Rhadinorhynchus pristis Filosoma sp. [RSea] 

Neorhadinorhynchus nudus [Pac] 

Rhadinorhynchoides sp. [Pac] 



Unicolax collateralis Caligus asymmetricus [IPac] 

Unicolax mycterobius Caligus biseriodentatus [Ind] 

307

308 


Caligus coryphaenae [Pac] 

Caligus macarovi [Pac] 

Caligus pelamydis [Pac] 

Caligus productus [Pac] 

Condition 

scombroid poisoning (rare) mass mortality [Pac] 

Euthynnus alletteratus (Rafinesque) - little tunny 

Name - It is also called "Atlantic little tunny", "little tuna", Gymnosarda 

alleterata and Thynnus thunnina. 


and anal fins. The first dorsal fin is much longer than the second dorsal. There 

is a complicated pattern of stripes on the posterior back that does not 

extend forward beyond the middle of the first dorsal fin. It usually has a few 

dark spots on the body between the pectoral and pelvic fins. 

Geographic Range - Atlantic Ocean from New England, USA, to Brazil; 

eastern Atlantic and Mediterranean; and sporadically in the Black Sea. All 

coastal areas in the western Atlantic except the Gulf coast of Mexico. 

Food Habits - It eats small pelagic fishes such as anchovies and herrings, fish 

larvae, squids and crustaceans. 




Parasites - This host is abundant and has apparently been rather well 

examined, but only supports 34 species of parasites, 24 from the western 

Atlantic. Three are host specific, 2 possibly host specific, 1 genus specific, 2 

tribe specific, 1 almost tribe specific, 2 family specific, 1 occurs on this host 

and Atlantic bonito, 1 on bonitos and little tunas, and 1 on little tunas and tunas, 

and 20 generalists with little host specificity. This fish has been noted to be the 

most heavily parasitized tuna in the eastern Atlantic and Mediterranean. 

Udonellidea (copepod worms) 

Udonella caligorum



Brachyphallus parvus Lecithochirium acutum [Eur] 

Dinurus scombri Lecithochirium monticellii [?] 

Hirudinella ventricosa Sterrhurus sp. [EAlt] 

Lecithochirium texanum Syncoelium katuwo [WAfr] 




Didymocystis thynni Allopseudocolocyntotrema alioshkini [Eur] 

Oesophagocystis lydiae [Eur] 

tissue fluke** [WAfr] 


Caballerocotyla manteri Hexostoma thunninae [EAtl] 

Hexostoma euthynni Tristomella onchidiocotyle [?] 


Lacistorhynchus bulbifer* Callitetrarhynchus gracilis* [EAlt] 

Otobothrium crenacolle* Otobothrium sp.* [EAlt] 


tetraphyllid* 





Caligus coryphaenae 

Caligus pelamydis 

Caligus productus 

Ceratacolax euthynni 


Unicolax anonymous 

Unicolax collateralis 

Unicolax mycterobius 


Nerocila orbignyi [EAtl] 


lipomas 

Condition 


Katsuwonus pelamis (Linnaeus) - skipjack tuna 

Name - The Japanese soup-stock product, "dried bonito" flakes or katsuobushi, 

is boiled, boned, smoke-dried, molded and finally shaven-flaked skipjack tuna. 

In Japan, what we call "little tunas" are called "bonitos". It has also been called 

"bonito", "ocean bonito","skipjack", "striped tuna", "watermelon tuna"; also 

Euthynnus pelamis (Linnaeus); E. pelamys; Gymnosarda pelamys; K. vagans; 

and Thynnus pelamys. 

309

310 



and anal fins. The first dorsal fin is much longer than the second dorsal. It has 

4-6 conspicuous longitudinal dark bands on the lower sides and belly. 

Geographic Range - Worldwide, cosmopolitan in warm waters, but not in the 

Black Sea. 

Food Habits - It eats fishes, squids, octopus, and crustaceans. 




Commercial Importance - Most "tuna" canned comes from this fish. It is 

also an important fresh fish over much of the globe. 

Parasites - This host appears to have been well examined with 75 parasite 

species recorded, except in the western Atlantic, where only 19 have been 

found. Eight parasites are host specific, 18 possibly host specific, 5 family 

specific, 5 occur on this host and bluefin tuna, 4 on tunas, 2 on bluefin tuna and 

yellowtail, 1 on a Pacific little tuna and a yellowtail, 1 almost genus specific but 

to mackerels (Scomber) not this host, and 3 on little tunas and tunas, and 29 are 

generalists with little host specificity. Its parasites suggest that it belongs in the 

tuna instead of the little tuna tribe (see Discussion). 


Hirudinella ventricosa Dinurus euthynni [Pac] 

Tergestia laticollis Lecithochirium microstomum [Pac] 

Syncoelium filiferum [Pac] 

Syncoelium katuwo [Pac] 


Adenodidymocystis intestinalis [Pac] 

Annulocystis katsuwoni [Pac] 

Coeliodidymocystis abdominalis [Pac] 

Coeliodidymocystis kamegaii [IPac] 

Didymocylindrus filiformis [Pac] 

Didymocylindrus simplex [Pac] 

Didymocystis abdominalis [Pac] 

Didymocystis bilobata [Pac]


Didymocystis ovata [Pac] 

Didymocystis philobranchia [Ind] 

Didymocystis reniformis [Pac] 

Didymocystis rotunditestis [Ind] 

Didymocystis soleiformis [Pac] 

Didymocystis submentalis [Pac] 

Didymocystis thynni [Eur] 

Didymocystis wedli [Pac] 

Didymocystoides intestinomuscularis [Pac] 

Didymocystoides pinnicola [Pac] 

Didymocystoides submentalis [Pac] 

Didymoproblema fusiforme [Pac] 

Didymozoon auxis [Eur?] 

Didymozoon filicolle [Pac] 

Didymozoon longicolle [Pac] 

Didymozoon minus [Pac] 

Koellikeria globosa [Pac] 

Koellikeria orientalis [Pac] 

Koellikeria reniformis [Pac] 

Lagenocystis katsuwoni [Pac] 


Nematobothrium scombri [Pac] 

Neodiplotrema pelamydis [Pac] 

Oesophagocystis dissimilis [Pac] 

Phacelotrema claviforme [Ind] 


Allopseudaxine katsuwonis Allopseudaxine vagans [Pac] 

Caballerocotyla katsuwoni [Pac] 

Hexostoma grossum [Pac] 

Pricea minimae [Ind?] 

Tristomella interrupta [Med] 


Tristomella lintoni [unc] 

Tristomella nozawae [Eur,Pac] 


Nybelinia lingualis* Callitetrarhynchus gracilis* [Pac] 

Tentacularia coryphaenae* Hepatoxylon trichiuri* [Eur] 

tetraphyllid* Pelichnibothrium sp.* [Pac] 

Pseudogrillotia basipuncata* [Pac] 

Rhyncobothrium sp.* [Pac,unc] 

Tentacularia sp.* [Pac] 


Anisakis simplex* Acanthocheilus sp. [Pac] 

Ctenascarophis lesteri Anisakis sp. [Pac] 

Philometra sp. Contracaecum sp. [Pac] 

Prospinitectus exiguus Philometroides sp. [Pac] 

311

312 


spiruroids Terranova sp.* [Pac] 


Rhadinorhynchus pristis Rhadinorhynchus trachuri [Pac?] 

Raorhynchus meyeri [Ind] 

Raorhynchus terebra [Ind] 


Caligus bonito Caligus asymmetricus [Pac] 

Caligus coryphaenae Caligus quadratus [Pac] 

Caligus mutabilis Lepeophtheirus branchialis [Ind,unc] 

Caligus productus Lepeophtheirus salmonis [Pac?] 

Lepeophtheirus bermudensis Unicolax reductus [Pac] 

Lepeophtheirus dissimulatus 



Isistius brasiliensis [Pac] 

Condition 

scombroid poisoning (frequent) hermaphroditism [IPac] 

Anomalies 

lacking body stripes [Pac] 

Tribe Thunnini - tunas 

Thunnus alalunga (Bonnaterre) - albacore 

Name - Only this fish can be called "white meat tuna". Other tunas, except 

bluefin, are called "light tuna". It is sometimes called "longfin tuna", "longfinned 

tunny", Germo alalunga (Bonnaterre), Orcyncnus alalunga, T. germo 

(Lacepede), and Thynnus alalunga. 


and anal fins. The pectoral fins are long, extending beyond the light yellow 

second dorsal and anal fins. The posterior margin of the caudal fin is white, 

and the anal finlets are dark. The ventral surface of the liver is striated. 

Geographic Range - Worldwide in tropical to temperate waters and in all seas, 

including the Mediterranean. This fish is common in the Caribbean, but not in 

the Gulf of Mexico.


Food Habits - It eats a variety of fishes, squids and crustaceans. 




Parasites - With 46 parasite species recorded, this host has been at least 

moderately examined, except in the western Atlantic, where only 13 of these 

parasites have been found. Indicative of their neglect are the 4 gillworms and 

15 tissue flukes not yet found in the western Atlantic. None of these parasites 

are known to be host specific, but 11 are possibly host specific, 8 genus specific 

(which is the same as tribe specific in this case), 4 family specific, 1 occurs on 

tunas and little tunas, 1 on tunas and amberjacks, 3 on tunas and skipjack tuna, 

and 18 generalists with little host specificity. 


Goussia auxidis [Pac] 

Hexacapsula neothunni [Pac] 


Hirudinella ventricosa Syncoelium filiferum [Pac] 


Didymocystis lanceolata [EAtl] 

Didymocystis macrorchis [EAtl] 

Didymocystis philobranchia [Pac] 

Didymocystis philobranchiarca [Ind] 


Didymocystis thynni [Med] 

Didymocystis wedli [Med] 

Didymocystoides alaongae [Pac] 

Didymocystoides buccalis [Pac] 

Didymocystoides opercularis [Pac] 

Didymocystoides superpalati [Ind] 

Didymonaja branchialis [Pac] 

Koellikeria bipartita [EAtl] 

Koellikerioides orientalis [EAtl,Pac] 

Metanematobothrium guernei [EAlt,IPac] 

Nematobothrium latum [EAtl] 

Platocystis alalongae [EAlt,Pac] 

Univitellodidymocystis lingualis [Ind] 


Areotestis sibi [Pac] 

Tristomella nozawae [Pac] 

Capsala thynni [WAfr,unc] 

Hexostoma sibi [Pac] 


Tentacularia coryphaenae* Hepatoxylon trichiuri* [Med,Pac] 

tetraphyllid* Pseudobothrium grimaldii [EAlt,unc] 

Sphyriocephalus tergestinus* [EAlt] 

313

314 


Nematoda (nematodes) 

Hysterothylacium cornutum Anisakis sp.* [Pac] 

Oncophora melanocephala Contracaecum sp.* [Pac] 



Rhadinorhynchus pristis Bolbosoma vasculosum [EAlt] 

Gorgorhynchus sp. [Pac] 


Brachiella thynni Caligus alalongae [EAlt,Ind] 

Caligus coryphaenae Caligus chorinemi [acc] 

Caligus productus Euryphorus nordmanni [Pac] 

Euryphorus brachypterus Lernanthropus hiatus [?] 

Pennella filosa 



Rocinella signata 


Isistius brasiliensis Isistius plutodus [Pac] 



osteoma [Pac] 

Condition 

stomach ulcer mesentary adhesions [SAfr] 


Thunnus albacares (Bonnaterre) - yellowfin tuna 

Name - It is sometimes called "Allison tuna", Neothunnus albacora (Lowe), 

N. macropterus (Temminck and Schlegel), N. albacores, T. argentivittatus 

(Cuvier), T. macropterus, Thynnus albacora, and T. macropterus. 


anal fins. Large fish have a relatively long second dorsal and anal fin, which 

are more than 1/5 of the FL. The pectoral fins are moderately long, reaching


beyond the notch between the dorsal fins. The ventral surface of the liver is not 

striated. 

Geographic Range - Worldwide in the tropics and subtropics. 



Length - Maximum more than 200.0 cm FL, common to 150.0 cm FL. 


Commercial Importance - Most canned "tuna" is skipjack tuna, but some 

comes from this fish. 

Parasites - This host appears to have been well examined and supports 85 

species of parasites, but only 32 have been found in the western Atlantic. 

Three of these parasites are host specific; 8 possibly host specific; 18 genus 

specific (which is the same as tribe specific in this case); 6 family specific; and 

1 each occur on tunas and little tunas, jacks and scombrids, tunas and little 

tunas, and tunas and Atlantic bonito; 1 almost host specific to wahoo; 3 on 

tunas and skipjack tuna, and 42 generalists with little host specificity. Thirteen 

species of larval tapeworms must indicate that this fish has a diverse diet of 

tapeworm intermediate hosts, because the average number among big game 

fishes is less than 5. 


Hexacapsula neothunni [Pac] 


Udonella caligorum 


Brachyphallus parvus Cardicola ahi [Pac] 

Hirudinella ventricosa Macradena sp. [Pac] 

Phyllodistomum thunni [WAfr] 



Atalostropion sardae Angionematoborium cephalodonus [Ind] 

Didymocystis acantyhocybii Dermatodidymocystis vivipara [Pac] 

Didymozoon longicolle Dermatodidymocystis viviparoides [Pac] 

Koellikeria bipartita Didymocystis sp. [Pac] 

Koellikeria globosa Didymocystis irregularis [Pac] 

Koellikeria orientalis Didymocystis orbitalis [Pac] 

Platocystis sp. Didymocystis palati [Pac] 

Didymocystis philobranchia [IPac] 

Didymocystis philobranchiarca [Pac] 


Didymocystis spirocauda [Pac] 

Didymocystis wedli [Med] 

Didymocystoides bifasciatus [Pac] 

Didymocystoides oesophagicola [Pac] 

Didymocystoides superpalati [IPac] 

Koellikeria abdominalis [Pac] 

315

316 


Neophrodidymofrema ahi [Pac] 


Univitellodidymocystis neothunni [Pac] 


Nasicola klawei Areotestis sibi [Pac] 

Caballerocotyla abidjani [WAfr] 

Caballerocotyla biparasitica [Pac] 

Caballerocotyla verrucosa [WAfr] 

Capsala gotoi [Pac] 

Capsala neothunni [Pac] 

Capsala thynni [Eur,unc] 

Hexostoma euthynni [Pac] 


Sibitrema poonui [Pac] 

Tristomella nozawae [Pac] 


Echeneibothrium sp.* Callitetrarhynchus gracilis* [WAfr] 

Grillotia sp.* Dasyrhynchus talismani* [EAtl,Pac] 

Gymnorhynchus gigas* Grillotia erinaceus [?] 

Hepatoxylon trichiuri* Tentacularia coryphaenae* [WAfr] 

Nybelinia sp.* 

Pelichnibothrium speciosum* 

Sphyriocephalus sp.* 


tetraphyllid* 

trypanorhynchid* 


Anisakis simplex* Anisakis sp.* [Pac] 

Hysterothylacium cornutum Ichthyostrongylus thunni [unc] 

Oncophora melanocephala Metanisakis sp. [Pac] 

Monhysterides sp. [Pac] 

Oncophora albacarensis [Eur] 

Philometroides sp. [Pac] 


Rhadinorhynchus pristis Bolbosoma sp. [Pac] 

Bolbosoma vasculosum* [WAfr] 

Neorhadinorhynchus sp. [Pac] 

Rhadinorhynchus trachuri [Pac?] 


Rhadinorhynchus cadenati [WAfr] 


Brachiella thynni Caligus sp. [WAfr] 

Caligus coryphaenae Caligus asymmetricus [IPac] 

Caligus productus Caligus quadratus [Pac] 

Euryphorus brachypterus Caligus robustus [IPac]


Pennella filosa Euryphorus nordmanni [Pac] 

Pseudocycnus appendiculatus Pennella sp. [WAfr,Pac] 


Glossobius impressus 1 



Isistius plutodus 


Remora remora 

Condition 


burnt tuna 

1Occurred in the stomach of this host, probably parasitized a flyingfish. 

Thunnus atlanticus (Lesson) - blackfin tuna 

Name - It is sometimes called "Bermuda tuna" and "black finned albacore". 


and anal fins. The pectoral are fins moderately long, reaching beyond the notch 

between dorsal fins. The right lobe of the liver is longer and the ventral surface 

is not striated. The finlets are dusky. 


Food Habits - It eats predominantly small fishes, but also squids and larval 

crustaceans. 




Parasites - This host has been inadequately examined and presumably supports 

more than 9 species of parasites. None of these parasites are host specific, 3 are 

genus specific (tribe specific in this case), 1 family specific, 5 are generalists 

with little host specificity. 

317

318 





Nasicola klawei 



tetraphyllid* 


Hysterothylacium cornutum 




Euryphorus brachypterus 





schwaunoma 

Condition 


Thunnus obesus (Lowe) - bigeye tuna 

Name - It is sometimes called Parathunnus sibi (Temminck and Schlegel), P. 

mebachi Kishinouye, P. obesus, and Thynnus sibi. 


and anal fins. The pectoral fins are moderately long, extending beyond the 

notch between the dorsal fins, in adults 110 cm FL or larger, and are relatively 

longer in smaller fishes. The finlets are bright yellow and edged with black. 

The ventral surface of the liver is striated. 

Geographic Range - Worldwide in the tropics and subtropics. 

Food Habits - It predominantly eats squids, but also a variety of fishes and 

crustaceans. 

Ecology - Oceanic.




Parasites - This host has been at least moderately examined since 53 parasite 

species have been recorded from it, except in the western Atlantic, where only 

6 species have been found. Two of these parasites are host specific, 11 

possibly host specific, 23 genus specific (tribe specific in this case), 5 family 

specific, 3 occur on tunas and skipjack tuna, and 9 generalists with little host 

specificity. We found 25 parasite species which were tribe specific to tunas. 

This fish harbors almost all of these parasites, possibly indicating that this fish 

species is older than the other tunas (see Discussion). 


Kudoa nova Hexacapsula neothunni [Pac] 


Botulus microporus [WAfr,unc] 

Cardicola ahi [Pac] 


Dermatodidymocystis vivipara [Pac] 

Dermatodidymocystis vivparoides [Pac] 

Didymocystis bifurcata [Pac] 

Didymocystis nasalis [Pac] 

Didymocystis orbitalis [Pac] 

Didymocystis philobranchia [IPac] 

Didymocystis philobranchiarca [IPac] 

Didymocystis poonui [Pac] 


Didymocystoides bifasciatus [Pac] 

Didymocystoides pectoralis [IPac] 

Didymocystoides superpalati [IPac] 


Koellikeria pylorica [Pac] 

Koellikeria retrorbitalis [Pac] 

Koellikeria submaxillaris [Pac] 

Koellikerioides apicalis [Pac] 

Koellikerioides externogastricus [Pac] 

Koellikerioides internogastricus [Pac] 

Koellikerioides intestinalis [Pac] 

Nematobothrium sp. [Ind] 

Neonematobothrioides poonui [Pac] 

Opisthorchinematobothrium parathunni [Pac] 

Orbitonematobothrium perioculare [Pac] 


Univitellodidymocystis neothunni [Pac] 


Areotestis sibi [Pac] 

Caballerocotyla biparasitica [Pac] 

319

320 


Caballerocotyla pseudomagronum [WAfr] 

Caballerocotyla verrucosa [WAfr] 

Capsala gotoi [Pac] 

Hexostoma acutum [Pac] 




Nasicola klawei [Pac,WAfr] 

Neohexostoma robustum [Pac] 

Sibitrema poonui [Pac] 

Tristomella nozawae [Eur,Pac] 

Tristomella onchidiocotyle [WAfr] 


tetraphyllid* Sphyriocephalus sp.* [?] 

Sphyriocephalus dollfusi* [WAfr] 

Dasyrhynchus talismani [EAtl,Pac] 


Philometroides sp. [Pac] 


Brachiella thynni Caligus alalongae [WAfr] 

Caligus coryphaenae Caligus productus [WAfr,IPac] 

Euryphorus brachypterus Euryphorus nordmanni [Pac] 





Condition 

scombroid poisoning (rare) "jelly-meat" [Pac] 

burnt tuna 

Thunnus thynnus (Linnaeus) - (northern) bluefin tuna 

Name - It is also called "horse mackerel", "tunny", Orcynchus thunnus, Scomber 

thunnus, T. orientalis, T. saliens, T. thunnus and Thynnus brachypterus. It


is not a "light tuna" and is not canned, but is highly desired for Japanese rawfish 

dishes. 

We use the name "bluefin tuna" in the text because it is an established 

name (Robins et al. 1991). "Northern bluefin tuna" is technically correct because 

"southern bluefin tuna", Thunnus maccoyii (Castelnau), occurs in the southern 

oceans below 40°S. 


and anal fins. The pectoral fins are relatively short, approximately 80% of the 

head length, and never reach the notch between the dorsal fins. The second 

dorsal fin is reddish-brown. The ventral surface of the liver is striated. The 

first gill arches have 34-43 rakers. 

Geographic Range - Subtropical and temperate areas in the south and north 

Atlantic, north Pacific Ocean, Mediterranean and Black Seas. Known from 

Labrador and Newfoundland down to northeastern Brazil in the Western 

Atlantic. 


Ecology - Oceanic. They do venture inshore in colder waters. Bluefin tuna 

maintain a body temperature up to 18°F higher (22-30°C) than the surrounding 

water (5-30°C), allowing them to function more effectively in cooler waters 

and tripling the power and response of their muscles. They can swim up to 50 

miles/hr (90 KPH), but are not the fastest fish in the sea. 

Length - Maximum over 300.0 cm FL (rumored 420.0 cm), formerly common 

to 200.0 cm FL. 

Weight - Maximum 682.0 kg (rumored to 700.0 kg), formerly common up to 

182.0 kg. 

Aquaculture - Bluefin tuna too small to sell to Japan are "ranched" by holding 

them in net cages or "pounds" in the ocean until they reach marketable size. 

These must experience parasite and disease problems. 

Commercial Importance - Bluefin tuna have become so valuable to the 

Japanese that a large fish sells for more than the annual salary of a fisherman. 

This fish is being hunted to extinction (extirpation) in the northwest Atlantic 

and the international and government agencies appear unwilling to intervene. 

Significance to Sport Fishing - Giant bluefin tuna are the highest prized big 

game sport fish, but unfortunately they may be more highly prized as a food 

fish in Japan. 

Parasites - Only 13 of the 72 known parasites of this fish have been found in 

the western Atlantic. This suggests that this host has not been adequately 

examined in our region. This trend is further substantiated by the fact that its 

7 species of copepods, which have received worldwide study, have all been 

found in the western Atlantic. Nine of the 72 species of parasites are host 

specific, 5 possibly host specific; 6 genus specific (tribe specific in this case); 

6 family specific; 4 occur on tunas, little tunas and amberjacks; 2 on tunas and 

little tunas; 2 on tunas and amberjacks; 3 on skipjack; 1 on little tunny; 4 on 

tunas and skipjack tuna; and 30 generalists with little host specificity. 

321

322 


Bacteria 

Vibrio spp. [Pac] 


Kudoa sp. [Pac] 

Kudoa clupeidae [WAfr] 

Uronema marinum [Pac] 


Hirudinella ventricosa Aponurus lagunculus [Pac] 

Bucephalopsis sibi [Pac] 

Cetiotrema crassum [Pac] 

Lecithaster gibbosus [Med] 

Lecithocladium excisum** [Eur] 

Prosorhynchoides sibi [Pac] 

Rhipidocotyle pentagonum [Pac] 

Rhipidocotyle septpapillata [RSea] 

Sterrhurus imocavus [Med,Pac] 


Koellikeria bipartita Anaplerurus thynnusi [Ind] 

Coeliotrema thynni [Pac] 

Didymocystis thynni [Eur,Med] 

Didymocystis crassa [Pac] 

Didymocystis ovata [Pac] 

Didymocystis reniformis [Pac] 

Didymocystis soleiformis [Pac] 

Didymocystis wedli [Med,Pac] 

Didymocylindrus filiformis [Pac] 

Didymocystoides semiglobularis [Pac] 

Didymoproblema fusiforme [Pac] 

Didymozoon filicolle [Pac] 


Didymozoon pretiosus [Pac] 

Koellikeria globosa [Pac] 

Koellikeria orientalis [Med,Pac] 

Koellikeria reniformis [Pac] 


Nematobothrium sp. [Pac] 

Oesophagocystis sp. [Pac] 


Tristomella onchidiocotyle [?] Caballerocotyla albsmithi [Pac] 

Caballerocotyla gouri [Ind] 

Caballerocotyla magronum [Pac] 

Caballerocotyla paucispinosa [Pac] 

Hexostoma acutum [Pac] 

Hexostoma albsmithi [Pac] 

Hexostoma dissimile [Pac]




Kuhnia thunni [Pac] 

Metapseudaxine ventrosicula [Pac] 

Neohexostoma extensicaudum [Eur] 

Neohexostoma thunninas [Med] 

Tristomella interrupta [Med] 

Tristomella nozawae [Eur] 

Tristomella onchidiocotyle [Med] 


Lacistorhynchus bulbifer* Callitetrarhynchus gracilis* [WAfr] 

Tentacularia coryphaenae* Grillotia sp.* [Ind] 

tetraphyllid* Pelichnibothrium speciosum* [Eur,Pac] 

Tetrarhynchus scomber-thynnus* [unc] 


Hysterothylacium cornutum Anisakis sp.* [Pac] 

Ascaris longestriata* [Med,unc] 

Contracaecum sp.* [Pac] 

Heptachona caudata [Pac] 

Hysterothylacium aduncum [Eur] 




Bolbosoma vasculosum [Pac] 

Neorhadinorhynchus nudus [Pac] 


Brachiella thynni Caligus balistae [fal] 

Caligus bonito Cecrops latreillii [Eur,Med?] 



Euryphorus brachypterus 




Isistius brasiliensis Isistius plutodus [Pac] 



fibroma 

lipomas 

melanoma 

osteoma 

Condition 

scombroid poisoning (rare) severe skin lesions [Pac] 

Uronema-like encephalitis [Pac] 

Anomalies 

vertebral [Eur] 

323

324 


Tribe Scomberomorini - Spanish mackerels 

Acanthocybium solandri (Cuvier) - wahoo 

Name - It is also called "jack-mackerel"; A. petus (Poey) and A. sara. 


and anal fins. The snout is relatively elongate and pointed. The first dorsal is 

much longer than the second dorsal fin. The sides of the body are covered with 

numerous, black vertical bars, which extend below the lateral line. 

Geographic Range - Worldwide in the tropics and subtropics, including the 


Food Habits - It eats moderate-sized to large surface fishes and squids. 

Ecology - Offshore and oceanic surface waters. 

Length - Maximum 210.0 cm FL, common 107.0-140.0 cm. 


Parasites - The 18 parasites reported from this important, worldwide host is an 

amazingly low number. We examined 15 wahoo from Puerto Rico, 4 from the 

northern Gulf of Mexico, and it has been examined elsewhere including an 

exclusive study in the Pacific (Iversen and Yoshida 1957). We are forced to 

conclude that it does not support many parasites. Only 11 of these parasites 

have been found in the western Atlantic. Five are host specific; 1 is tribe 

specific to little tunas, but rarely occurs on wahoo; 1 is family specific 

occurring on tunas, little tunas and wahoo; 11 are generalists with little host 

specificity. Its parasites suggest that it belongs with little tunas and not with 

Spanish mackerels (see Discussion). 


Udonella caligorum 


Hirudinella ventricosa Tetrochetus coryphaenae [acc] 


Didymocystis acanthocybii Nematobothrium spinneri [Pac] 

Univitellodidymocystis miliaris [Pac] 


Caballerocotyla manteri Tristomella nozawae [Pac] 

Neothoracocotyle acanthocybii 


Hepatoxylon trichiuri* 

Tentacularia coryphaenae*


tetraphyllid* 


Brachiella thynni Caligus coryphaenae [Pac] 

Caligus productus Lernaeolophus sultanus [Ind] 

Gloiopotes hygomianus Pennella filosa [?] 

Shiinoa occlusa [Pac] 

Tuxophorus cybii [Ind] 




Remora osteochir** 

Scomberomorus brasiliensis Collette, Russo and Zavalla-Camin 

- serra Spanish mackerel 

Name - This species was described in 1978, before that time it was confused 

with Atlantic Spanish mackerel. 


and anal fins. The sides of the body have many, relatively small, yellow or 

bronze spots. The spots extend from the back onto the belly, and surround the 

pectoral fin. The anterior 2/5 of the first dorsal fin is black, and the highest part 

of the fin is in the middle of the black region. 

Geographic Range - Central and South American coast from Yucatan, Mexico 

to southern Brazil. It is not found in the insular Caribbean. 

Food Habits - It predominantly eats small fishes. 




Parasites - This host has not been adequately or thoroughly examined for 

parasites. Six parasites are genus specific; 1 almost genus specific, rarely 

occurring on other hosts; and 5 generalists with little host specificity. 


Rhipidocotyle baculum 


Didymocystis scomberomori 

325

326 



tetraphyllid* 


Hysterothylacium fortalezae 

Hysterothylacium reliquens* 

Hysterothylacium sp.* 


Caligus mutabilis** 

Holobomolochus divaricatus 


Pseudocycnoides buccata 

Shiinoa inauris 


Livoneca redmanii 

Scomberomorus cavalla (Cuvier) - king mackerel 

Name - It is also called "kingfish". 


and anal fins. The lateral line turns down abruptly under the second dorsal fin. 

The sides of the body are plain and lack markings. 

Geographic Range - Western Atlantic tropics and subtropics. 

Food Habits - It predominantly eats small fishes. 

Ecology - Inshore and offshore, not oceanic. This fish tends to be found 

further offshore than the 3 other species of Spanish mackerels in the western 

Atlantic. 



Parasites - This fish has been well examined for parasites in Puerto Rico and 

many other localities, but it only supports 23 parasite species. One is host 

specific, 6 genus specific, 1 almost genus specific, and 15 generalists with little 

host specificity. The parasites of this host suggests that it differs slightly from 

the other western Atlantic members of the genus (see Discussion). 


Haemogregarina bigemina



Bucephalopsis arcuata Hirudinella ventricosa [acc] 

Lecithochirium sp. Rhipidicotyle capitata [fal] 



Gotocotyla acanthophallus 

Pseudaxine mexicana 

Scomberocotyle scomberomori 

Thoracocotyle crocea 


Nybelinia lamonteae* 




tetraphyllid* 


Hysterothylacium fortalezae Porrocaecum paivai [unc] 




Bolbosoma vasculosum 


Brachiella thynni Caligus bonito [?] 

Caligus mutabilis** Caligus elongatus [?] 


Holobomolochus asperatus 



Anilocra acuta 



Condition 


Scomberomorus maculatus (Mitchill) - (Atlantic) Spanish mackerel 

Name - It is also called "Spanish mackerel" in the USA (see Host Summaries 

introduction), and "spotted Spanish mackerel" (Fischer 1978). 


and anal fins. The sides of the body have few, relatively large, yellow or 

bronze spots. The spots do not extend onto the belly and do not surround the 

pectoral fin. The anterior 1/3 of the first dorsal fin is black. 

Geographic Range - Western Atlantic from Yucatan, Mexico to Maine, USA,. 

Not in Bermuda or the West Indies, except possibly in Cuba and south Florida. 

Previously confused with cero to the north and serra Spanish mackerel in the 

south. 

327

328 


Food Habits - It predominantly eats small fishes, especially anchovies and 

herrings. 

Ecology - In shore and offshore. 



Parasites - This fish has been well examined for parasites from many localities, 

but it is only known to support 24 parasite species. Two are possibly host 

specific, 10 genus specific, and 12 generalists with little host specificity. 


Kudoa crumena 


Bucephalopsis arcuata Bucephalus confusus [acc] 







Scomberocotyle scomberomori 



Callitetrarhynchus gracilis* Dibothriorhynchus speciosum* [unc] 

Lacistorhynchus bulbifer* Rhynchobothrium longispine* [unc] 



tetraphyllid* 



Hysterothylacium fortalezae* 


Hysterothylacium sp.* 

Philometra sp. 


Caligus mutabilis** Caligus bonito [?] 

Holobomolochus divaricatus Caligus elongatus [?] 

Lernaeenicus longiventris Caligus productus [?]


Pseudocycnoides buccata Charopinopsis quaternia [acc] 

Shiinoa inauris Anuretes heckelii [acc] 


Livoneca ovalis 

Condition 


Scomberomorus regalis (Bloch) - cero 


anal fins. The sides of the body have a central line of yellow lines or streaks 

with a row of relatively small, yellow spots above and below. The anterior 1/3 

of the first dorsal fin is black. The pectoral fins are covered with small scales. 

Geographic Range - Western Atlantic from Massachusetts, USA, down the 

Atlantic coast and the West Indies to Brazil. It is not found in the Gulf of 

Mexico or the Central American and Colombian coast. This fish is most 

abundant in the West Indies. 

Food Habits - It predominantly eats small, schooling fishes, but sometimes 

squids and shrimps. 




Parasites - This fish has been well examined for parasites in Puerto Rico and 

from many other localities, but it is only known to support 21 parasite species. 

One is possibly host specific, 9 genus specific, 1 almost genus specific, and 10 

generalists with little host specificity. Heavy to very heavy infections of a tissue 

fluke, D. scomberomori, and heavy infections of an encysted larval roundworm, 

H. reliquens, occurred in this host in a severely contaminated area (Tetra Tech 

1992) in eastern Puerto Rico. This appears to be an example of pollution effects 

increasing the numbers of parasites. 




Bucephalopsis arcuata Myosaccium opisthonema [fal] 




329

330 










tetraphyllid* 

Nematoda (nematode) 




Brachiella thynni 



Holobomolochus divaricatus 


Shiinoa inauris 

Tuxophorus collettei 


Livoneca redmanii 

Condition 


Superfamily Xiphioidea, Family Xiphiidae - swordfishes 

Xiphias gladius Linnaeus - swordfish 

Name - It is sometimes called "broadbill swordfish"; Histiophorus gladius, 

Istiophorus gladius, and X. zeugopteri. 

Diagnostic Characters - All billfish have an elongate bill on the upper jaw. 

The bill of this fish is sword-shaped, wider than thick, and twice as long as the 

head. It has no pelvic fins. The first dorsal fin is relatively large, the second 

dorsal small, and they are widely separated. 

Geographic Range - Worldwide in tropical and temperate waters. 

Food Habits - It predominantly eats small to relatively large fishes, and 

occasionally squids and crustaceans.


Ecology - Inshore to oceanic. It can swim at a speed of 60 miles/hr [108 KPH]. 

Length - Maximum 450.0 cm, formerly common to 220.0 cm. 

Weight - Maximum 536.4 kg, formerly common 50.0-135.0 kg. 

Human Health - Large fish accumulate high concentrations of mercury in their 

flesh. This discovery caused problems with commercial sales until U.S. 

Agencies identified the mercury component harmful to humans and adjusted 

existing regulations. 

Commercial Importance - The high prices paid for swordfish has incited over 

exploitation of this fish. 

Parasites - Fourty-nine species is actually a rather low number of parasite 

species for such an important, worldwide and presumably well studied host. A 

little more than 1/2 of these species have been found from the western Atlantic. 

Of the 49 species of parasites, 13 are host specific, 6 superfamily specific, and 

31 generalists with little host specificity. The high number of host-specific 

parasites (26%) indicate that this fish is evolutionarily older than any of the 

other members of the swordfish-billfish superfamily. This also suggests that it 

is well isolated and distinct from the others and deserves its own family. This 

fish harbors 20 species of larval tapeworms, indicating that it has a rich and 

diverse diet of tapeworm intermediate hosts (the average number among big 

game fishes is less than 5). 


Kudoa musculoliquefaciens [Pac] 


Hirudinella ventricosa Cardicola sp. [Pac] 


Maccallumtrema xiphiados Metadidymozoon branchiale [IPac] 


Neodidymozoon macrostoma [Pac] 

Reniforma multilobularis [Pac] 


Tristoma coccineum Tristoma adcoccineum [Pac] 

Tristoma integrum Tristoma adintegrum [Pac] 


Tristomella pricei [Pac] 


Ceratobothrium xanthocephalum* Bothriocephalus claviger [unc] 

Dasyrhynchus giganteus* Bothriocephalus manubriformis [unc] 

Fistulicola plicatus Floriceps saccatus* [Eur] 

Grillotia erinacea* Gymnorhynchus gigas* [Eur] 

Hepatoxylon trichiuri* Nybelinia sp.* [Ind] 

Molicola horridus* Rhynchobothrium ambiguum* [unc] 

Nybelinia bisulcata* Taenia sp.* [Pac,unc] 

Nybelinia lamonteae* Tetrarhynchus spp.* [unc] 

Nybelinia lingualis* 


331

332 


Otobothrium dipsacum* 

Phyllobothrium delphini* 


tetraphyllid* 


Anisakis simplex* Hysterothylacium aduncum* [Eur] 

Hysterothylacium aduncum Hysterothylacium hanumantharoi [Ind] 

Hysterothylacium corrugatum Hysterothylacium petteri [Pac,unc] 

Maricostula incurva Hysterothylacium reliquens [acc] 

Oncophora melanocephala [fal] 

Paranisakis multipapillus [Ind] 




Caligus coryphaenae Caligus chelifer [acc] 

Caligus elongatus Chondracanthus xiphiae [unc] 

Philichthys xiphiae Gloiopotes huttoni [IPac] 

Pennella filosa Gloiopotes watsoni [Ind] 

Pennella instructa Lernaeolophus sultanus [Ind,acc] 

Pennella sp. [Ind] 

Thysanote ramosa [Med] 


Conchoderma virgatum Conchoderma sp. [Pac] 


Nerocila californica [Pac] 

Nerocila phaiopleura [Ind] 







Condition 

stomach ulcers 

Family Istiophoridae - billfishes 

Istiophorus albicans (Latreille) - Atlantic sailfish 

[Istiophorus platypterus (Shaw and Nodder) - Indo-Pacific sailfish] 

Name - It is also called Histiophorus albicans (Latreille), H. americanus 

Cuvier, H. orientalis, I. americanus (Cuvier), I. greyi, and I. orientalis. The 

Indo-Pacific sailfish is sometimes combined with it to form one worldwide 

species of "sailfish". 


The dorsal fin is enlarged into a broad and wide sail. The pelvic fins are 

approximately twice as long as the pectoral fins.


Geographic Range - Tropical and temperate Atlantic. The Indo-Pacific 

sailfish occurs throughout the tropical and temperate Indian and Pacific 

Oceans. 

Food Habits - It eats a variety of fishes, crustaceans and squids. 

Ecology - Inshore to oceanic. This is the fastest swimming fish. It has been 

recorded swimming 68 miles/hr [122.4 KPH]. 


Weight - Maximum 79.5 kg, common 12.0-20.0 kg. The maximum for the 

Indo-Pacific sailfish is 100.2 kg. 

Parasites - With only 34 parasite species reported from this host, it appears to 

have been under examined, particularly in the western Atlantic where only 12 

are known. Three of the 34 parasites are host specific, 2 possibly host specific, 

1 genus specific, 9 family specific, 5 superfamily specific, and 14 are 

generalists with little host specificity. These parasites provide little definitive 

evidence to divide or combine Atlantic and Indo-Pacific sailfish as species. 

The 1-2 different host-specific parasites on each fish could be due to regional 

parasite distribution or specificity. The differences in tissue flukes may be due 

to under reporting in the Atlantic (see Discussion). 

Bacteria 

bacterial infection 




Hirudinella ventricosa Cardicola grandis [Pac] 

Dinurus scombri [fal] 

Parahemiurus merus [Pac] 


Colocyntotrema sp. Angionematobothrium jugulare [Pac] 

Metadidymozoon branchiale [IPac] 



Neodidymozoon midistoma [Ind] 

Unitubulotestis istiophorusi [Ind] 

333

334 



Tristomella laevis Caballerocotyla marielenae [Pac] 

Caballerocotyla megacotyle [Ind] 

Capsaloides istiophori [Pac] 

Capsaloides sinuatus [Pac] 

Tristomella ovalis [Pac] 

Tristomella pricei [IPac] 


Bothriocephalus manubriformis Callitetrarhynchus gracilis* [Pac] 

Otobothrium dipsacum* Floriceps minacanthus* [Pac] 


tetraphyllid* 


Maricostula histiophori Paracanthocheilus striatus [Ind,unc] 




Gloiopotes americanus Caligus quadratus [Pac] 

Pennella filosa Gloiopotes huttoni [IPac] 

Gloiopotes watsoni [IPac] 

Lepeophtheirus eminens [Pac] 

Lernaeolophus sultanus [Ind] 

Pennella biloboa [Pac,unc] 

Pennella instructa [Pac?] 


Conchoderma virgatum [Pac?] 


Nerocila californica [Pac] 


Naucrates ductor Isistius brasiliensis [Pac] 



Remora remora 

Remorina albescens 


liver granulomas 

Condition 

hyperplasia 

stomach and intestinal tumors 

Makaira indica (Cuvier) - black marlin 

Name - It has also been called Istiompax indicus (Cuvier), I. marlina (Jordan 

and Hill), and M. marlina Jordan and Hill. 

Black marlin are reported in the Atlantic by commercial longline fishermen, 

but their presence has not been confirmed by the scientific community. They


may have inspired great "giant-marlin-that-got-away" stories just off Puerto 

Rico, but have not been examined for parasites in the Atlantic. Please bring us 

one! 


This fish is the only billfish with rigid pectoral fins that cannot be folded against 

the body. The tallest part of the dorsal fin is less than 1/2 the midbody depth. 

Geographic Range - Indian and Pacific Ocean, tropical subtropical and 

sometimes temperate waters, only sporadically in the Atlantic. Rarely reported 

from the Caribbean and west Africa. Some of the giant marlin hooked, but not 

landed, off Puerto Rico may be this fish. 

Food Habits - It eats squids and pelagic fishes. 

Ecology - Inshore to oceanic. This fish is much more abundant in coastal 

waters than in the open sea. 



Commercial Importance - This fish is called "white marlin" in Japan because 

of its white, firm flesh. It commands a high price on the commercial market. 

Parasites - This host probably supports more than the 24 species of parasites 

reported. Too few fish have been examined in the Indo-Pacific, and none in the 

Atlantic. Two of these parasites are host specific, 10 family specific, 5 superfamily 

specific, and 7 generalists with little host specificity. 


Cardicola grandis [Pac] 

Hirudinella ventricosa [Pac] 


Angionematobothrium jugulare [Pac] 

Glomeritrema subcuticola [Pac] 

Makairatrema musculicola [Pac] 

Metadidymozoon branchiale [Pac] 



Torticaecum sp.* [Pac] 


Capsaloides cristatus [Pac] 

Capsaloides isiophori [Pac] 

335

336 


Capsaloides tetrapteri [Pac] 

Tristomella laevis [Ind] 

Tristomella pricei [Pac] 


Bothriocephalus manubriformis [Pac] 

Callitetrarhynchus gracilis* [Pac] 

Floriceps minacanthus* [Pac] 

Otobothrium dipsacum* [Pac] 

Pseudogrillotia zerbiae* [Pac] 

tetraphyllid* [IPac] 


Anisakis sp. [Pac] 

Camallanus sp. [Pac] 

Hysterothylacium pelagicum [Pac,acc] 

Maricostula makairi [Pac] 


Gloiopotes huttoni [IPac] 

Gloiopotes watsoni [Ind] 

Lepeophtheirus eminens [Pac] 

Pennella instructa [?] 


Naucrates ductor Isistius brasiliensis [Pac] 

Remora brachyptera [Pac] 

Condition 

stomach tumors 

Makaira nigricans Lacepède - Atlantic blue marlin 

[Makaira mazara Jordan and Snyder - Indo-Pacific blue marlin] 

Name - It is sometimes called "blue marlin"; also M. ampla (Poey) and M. 

marlina. A "sister species" Indo-Pacific blue marlin is sometimes combined 

with it to form one worldwide "blue marlin". 


The pelvic fins are shorter than the pectoral fins; and the tallest part of the 

dorsal fin is shorter than the midbody depth.


Geographic Range - Atlantic tropical, subtropical and sometimes temperate 

waters. The Indo-Pacific blue marlin is found throughout that region. 

Food Habits - It predominantly eats a variety of small fishes (99%), and some 

squids (1%). They will rarely eat larger fishes. 

Ecology - Offshore to oceanic. It is more abundant in the tropics. 


Weight - Maximum 636.0 kg, common 80.0-182.0 kg. The maximum for 

Indo-Pacific blue marlin is 820.5 kg. 

Parasites - This host has been well examined, but with only 28 species 

worldwide, it is a parasite-poor host, and especially so in the western Atlantic 

where there are only 18. Only 1 of the 28 parasites is host specific, 2 possibly 

host specific, 8 family specific, 4 superfamily specific, and 13 generalists with 

little host specificity. Its single host specific parasite and its low total number 

of parasites, might be due to a relatively short evolutionary history. These 

parasites provide little definitive evidence to divide or combine Atlantic and 

Indo-Pacific blue marlin as species. 




Colocyntotrema sp. Metadidymozoon branhiale [Ind] 

Neodidymozoon macrostoma Unitubulotestis makairi [Ind] 

Wedlia submaxillaris [Ind] 


Tristomella laevis Capsala ovalis [IPac] 

Capsaloides nairagi [Ind] 

Capsaloides sinuatus [Pac] 

Tristomella pricei [IPac] 


Bothriocephalus manubriformis 



Pelichnibothrium speciosum* 

Prosobothrium armigerum* 



tetraphyllid* 


Maricostula sp. Oncophora melanocephala [fal] 

Philometroides sp. 




Gloiopotes ornatus Gloiopotes huttoni [IPac] 

Pennella filosa Gloiopotes watsoni [IPac] 

Pennella makaira Lepeophtheirus eminens [Pac] 

337

338 

Conchoderma virgatum 


Isistius plutodus 




broken bills 


malformed bills 




Condition 

Anomalies 

Tetrapturus albidus Poey - white marlin 

Name - It is sometimes called "spikefish"; also Lamontella albida (Poey), 

Makaira albida (Poey), T. albidus, T. belone, and T. imperator. 


The pelvic fins are shorter than the pectoral fins; and the tallest part of the 

dorsal fin is equal to the midbody depth. The ends of the pectoral and anal fins 

are rounded. 

Geographic Range - Atlantic and Mediterranean tropical, subtropical and 

sometimes temperate waters. 





Parasites - The parasites of the white marlin are very similar in number and 

species to those of Atlantic blue marlin. Only a couple of genus-specific 

gillworms distinguish this host. The parasites of these 2 hosts suggest that they 

are so similar that they should be in the same genus. One of these parasites is 

host specific, 2 genus specific, 4 family specific, 1 superfamily specific, and 9 

generalists with little host specificity. 


Hirudinella ventricosa



Neodidymozoon macrostoma Colocyntotrema sp. [?] 


Capsaloides cornutus Capsaloides perugiai [Med] 

Capsaloides magnaspinosus 

Tristomella laevis 



Gymnorhynchus gigas* 




tetraphyllid* 


Maricostula sp. 


Rhadinorhynchus sp. 



Gloiopotes ornatus 







Remora remora 


Condition 

broken bill 


Tetrapturus pfluegeri Robins and de Sylva - longbill spearfish 


The pelvic fins are longer than the pectoral fins; and the tallest part of the dorsal 

339

340 


fin is slightly greater than the midbody depth. The anus is also well in front of 

the anal fin, instead of adjacent as in other billfishes. 

Geographic Range - Atlantic tropical, subtropical and sometimes temperate 

waters. Mediterranean spearfish replaces it in the Mediterranean. This fish 

overlaps ranges and possibly exchanges parasites with 3 other species of 

spearfishes in the eastern Atlantic. 

Food Habits - It eats primarily pelagic fishes and squids. 

Ecology - Oceanic. 

Length - Maximum 200.0 cm, more often to 165.0 cm. 

Weight - Maximum 45.0 kg, more often to 20.0 kg. 

Significance to Sport Fishing - Rarely caught on hook and line. Taken on 

longlines. 

Parasites - We found 8 species of parasites in a single specimen of this host we 

were able to examine. One fish is hardly an adequate sample. All of these 

parasites are found in white marlin, as would be expected in hosts from the 

same genus. None of these parasites is host specific, 2 genus specific, 2 family 

specific, 1 superfamily specific, and 3 generalists with little host specificity. 


Hirudinella ventricosa Dinurus scombri [fal] 


Neodidymozoon macrostoma 


Capsaloides cornutus 

Capsaloides magnaspinosus 

Tristomella laevis 



tetraphyllid* 








Condition 

stomach ulcer 

Miscellaneous fish 

Sarda orientalis (Temminck and Schlegel) - striped bonito 

Atlantic records of this wide-ranging Indo-Pacific fish were in error (Robins et 

al. 1991).

ACKNOWLEDGMENTS 

ASSISTANCE 

Our research detailed in this book was supported by the Commonwealth of 

Puerto Rico Department of Natural and Environmental Resources (DNER) with 

Wallop-Breaux Sportfish Restoration Funds (Puerto Rico Projects F-28 and F- 

35) and the University of Puerto Rico, Mayaguez Campus (UPRM) in Puerto 

Rico; and the Southeastern Cooperative Fish Disease Project, Department of 

Fisheries and Allied Aquacultures (DFAA), Auburn University, with Wallop- 

Breaux Funds in Alabama. The Caribbean Aquatic Animal Health Project, 

Caribbean Stranding Network and the Caribbean Aquaculture Association also 

provided information and/or support. 

We are deeply grateful to Dr. William G. Dyer, Department of Zoology, 

Southern Illinois University, Carbondale, Illinois, USA, for discussing many 

points of this work with us, preparing and depositing some of the parasites 

collecteded in this study, and encouraging our efforts. 

We thank the members of the organizations listed above and the big game 

fishermen at official tournaments and the many people in the organizations noted 

below who helped collect specimens and cooperated with us in our work, and 

a small army of high school to post-doctoral students who assisted us in 

laboratory and field work related to this study, especially: Mickey Tirado, 

President, Asociación de Pesca Deportiva de Puerto Rico; Arecibo Outboard 

Club, Club Deportivo del Oeste, Club Nautico de Arecibo, Club Nautico 

Boqueron, Club Nautico La Parguera, Club Nautico Mayaguez; Aguadilla, 

Crashboat and La Parguera Fishing Co-ops; Dauphin Island Deep Sea Fishing 

Rodeo; Iris Corujo-Flores, José Berrios, James Timber, Timothy Churchill, Dr. 

Craig G. Lilyestrom, Catherine Aliaume, Alfonso Zerbi, Miguel Figuerola, 

DNER; Dr. Luis R. Almodovar (deceased), Mickey Amador, Aramis Aversa, 

Robin and Andy Bruckner, Dr. Ileana E. Clavijo, Dr. Patrick L. Colin, Dr. 

George D. Dennis, Dr. Michael J. Dowgiallo, Dr. Jorge R. Garcia-Sais, Kathy 

Hall, Dr. Dannie A. Hensley, Nilda Jimenez, Dr. Joseph J. Kimmel, Ed Levine, 

Ivan Lopez, Uchi Mendoza, Dr. Antonio A. Mignucci-Giannoni, Rubby 

Montoya, Dr. Debra P. Moore, Edgardo Ojeda, Edgardo Ortiz, Jorge Rivera, 

Juan Rosado, Marcos Rosado, Victer Rosado, William Rosado, Llena Sang, Dr. 

Rosa M. Steele, Dr. Richard K. Wallace and Dr. Raymond E. Waldner, 

Department of Marine Sciences (DMS), UPRM; Sheila Dunstan, Yolanda Lopez 

and Sylma Martinez de Aymat, DMS Library; Dr. Yolanda Brady (DFAA), 

Lucy and Dolph Bunkley, Adrian Cooper (Jamaica), Patrick Cotter (Bellairs 

Research Institute, Barbados), Sherel and Dallas Durrance, Richard Dyer, 

Antonio Garcia, Marta and Dr. Francisco A. Guzman-Reyes, Froilan Lopez, 

Juan Muñoz (Punta Betin Marine Laboratory, Santa Marta, Colombia), Dr. 

Ronald P. Phelps (DFAA), Dr. Doon Ramsaroop and Dr. Max Sturm (Institute 

of Marine Affairs, Trinidad), Dr. Rand (Bermuda Aquarium), Miguel Rolon 

341

342 


(Caribbean Fishery Management Council) Carol Sanner, Dr. Joseph R. Sullivan 

(Alaska Fish and Game), Wayne Swingle (Gulf of Mexico Fishery Management 

Council), Gerard Van Buurt (Department of Agriculture and Fisheries, Curaçao) 

and Luis Vivoni. 

We received unpublished records of tumors, pseudotumors, conditions and 

parasites from Dr. Harshbarger, copepods from Drs. Hogans and Benz, a 

remora from Dr. Goldstein, gillworms from Dr. Lester, and a hermaphroditic 

skipjack tuna from Dr. David Itano, Hawaii Institute of Marine biology, 

University of Hawaii at Manoa. We received information about ostracodes from 

Dr. McKenzie, fishes from Dr. Dannie A. Hensley (DMS), and bacteria from 

Dr. Esther C. Peters. Christoph Schmitt made translations of parts of German 

articles. 

Our parasite samples were deposited and numbers assigned in the following 

museums: helminths by Dr. J. Ralph Lichtenfels, U.S. National Parasite 

Collection (USNPC), USDA Beltsville, Maryland; crustaceans by Drs. Thomas 

E. Bowman (deceased) and Brian Kensley, Division of Crustacea, Smithsonian 

Institution, U.S. National Museum (USNM); tumors by Dr. John C. 

Harshbarger, Registry of Tumors in Lower Animals (RTLA); and a reference 

collection (for local use) by Dr. Jeff Holmquist, Invertebrate Museum (DMS). 

We thank Drs. E. W. Shell, Grizzle, Rogers and Plumb (DFAU) and Prof. 

Edgardo A. R. Ortiz-Corps, Department of Biology, University of Puerto Rico 

at Humacao for laboratory and office space, library facilities, reference collections 

and accommodations for completing parts of this work. 

ILLUSTRATION SOURCES 

Illustrations of the following species were redrawn or further modified from 

illustrations in the sources listed: 

Protozoans: Trypanosoma sp. (original), Kudoa crumena (Iversen and Van 

Meter 1967), other protozoans (Lom and Dyková 1992). 

Fungi: (Neish and Hughes 1980). 

Copepod worm: (Yamaguti 1963). 

Flukes: Brachyphallus parvus, Lecithochirium sp., Stephanostomum coryphaenae, 

S. sentum (Siddiqi and Cable 1970); Bucephalopsis arcuata, L. 

microstomum, L. texanum, Rhipidocotyle baculum, Tormopsolus orientalis 

(original); B. longicirrus, B. longovifera, Bucephalus varicus, S. 

megacephalum (Manter 1940a,b); B. gorgon (Corkum 1967); Cetiotrema 

carangis (Manter 1947); Coitocaecum extremum (Thatcher 1993); Dinurus 

scombri, R. barracudae (Schell 1985); Hirudinella ventricosa (Gibson and 

Bray 1977); Pseudopecoeloides carangis, Tetrochetus coryphaenae (Yamaguti 

1970); R. capitata (Manter 1944); S. imparispine, Tormopsolus filiformis 

(Sogandares-Bernal and Hutton 1959); other flukes (Yamaguti 1971). 

Tissue flukes: Colocyntotrema auxis, Didymocystis acanthocybii, 

Neodidymozoon macrostoma (Yamaguti 1970); D. wedli (Madhavi 1982); D. 

scomberomori (Overstreet 1969); D. thynni, Didymozoon longicolle (original);

ACKNOWLEDGEMENTS 

Koellikeria bipartita, Nematobothrium pelamydis (Grabda 1991); 

Maccallumtrema xiphiados, N. scombri (Yamaguti 1971); other tissue flukes 

from the original descriptions. 

Gillworms: Allopyragraphorus incomparabilis, Allopseudaxine katsuwonis, 

Cemocotyle carangis, Cemocotylella elongata (Yamaguti 1963); Allencotyla 

mcintoshi (Schell 1970); Capsaloides magnaspinosus, Protomicrocotyle mirabilis, 

Tristoma integrum (original); Dionchus agassizi (Rohde 1978); Helixaxine 

winteri (Caballero and Bravo-Hollis 1965); Pseudochanhanea sphyraenae 

(Yamaguti 1968); other gillworms (Hendrix 1994). 

Tapeworms: Bothriocephalus manubriformis (Yamaguti 1968); Callitetrarhynchus 

gracilis, Pterobothrium heteracanthum (Chandler 1935); Dasyrhynchus 

giganteus, Fistulicola plicatus, Gymnorhynchus gigas, Otobothrium dipsacum 

(Khalil et al. 1994); C. gracilis, Eutetrarhynchus lineatus (Linton 1908, Mac- 

Callum 1921); F. plicatus (Hogans and Hurley 1986); Ceratobothrium xanthocephalum, 

Grillotia erinaceus, Hepatoxylon trichiuri, Lacistorhynchus bulbifer, 

Nybelinia bisulata, P. heteracanthum (Linton 1897b); H. trichiuri (Linton 

1941); L. bulbifer, (Schmidt 1986); N. lamonteae (Nigrelli 1938); N. lingualis 

(Schmidt 1986); O. crenacolle, Rhinebothrium flexile (Linton 1905); Pelichnibothrium 

speciosum (Wardle and McLeod 1952); Pseudogrillotia zerbiae 

(Palm 1995). 

Roundworms: Aniskis simplex, Oncophora melanocephala (original); 

Hysterothylacium aduncum (Berland 1961); H. cornutum, Maricostula 

histiophori (Bruce and Cannon 1989); H. fortalezae, H. reliquens, Iheringascaris 

inquies, M. incurva (Deardorff and Overstreet (1981); other roundworms from 

the original descriptions. 

Ostracod: (Wilson 1913). 

Copepods: life cycle stages (Wilson 1905); Brachiella thynni, Clavellisa 

scombri, Gloiopotes americanus, G. ornatus, Hatschekia amplicapa, Pennella 

sp., Tuxophorus colletti (original); Caligus balistae, C. chorinemi, C. 

coryphaenae, C. isonyx, C. longipedis, C. wilsoni (Cressey 1991); C. bonito, 

C. mutabilis, C. pelamydis, C. productus; Ceratacolax euthynni, Euryphorus 

brachyptera, Holobomolochus asperatus, H. divaricatus, Pseudocycnus appendiculatus, 

Pseudocycnoides buccata, Shiinoa inauris, Unicolax collateralis 

(Cressey and Cressey 1980); C. quadratus, C. spinosus (Shinno 1959, 1960); 

H. crevalleus (Cressey 1981); Pennella makaira (Hogans 1988a); Philichthys 

xiphiae (Kabata 1979); U. anonymous, U. mycterobius (Vervoort 1965); other 

copepods (Yamaguti 1963). 

Fish louse: (Yamaguti 1963). 

Isopods: Rocinella signata (Menzies and Glynn 1968); other isopods 

(Kensley and Schotte 1989). 

Fish associates: Isistius brasiliensis and I. plutodus (Compagno 1984); 

pilotfish (Fischer 1978). 

Fish hosts: (Fischer 1978). 

Other diseases: Pseudotumor drawn from a specimen (RTLA 1895) 

donated to the RTLA by Walter Kandashoff. 

343

344 

BIBLIOGRAPHY 

These citations include literature cited and/or used in this book and 

additional references which we believe may be useful in the study of these 

parasites. Due to space limitations, references concerning parasites from 

outside the western Atlantic are not cited. Popular articles are marked with 

an asterisk (*). 

Amin, O.M. 1985. Classification. Pages 27-72 In Crompton, D.W.T. and B. B. 

Nickol (Eds.) Biology of the Acanthocephala. Cambridge University 

Press, 519 p. 

Anderson, D. 1993. Barnacles, structure, function, development, and evolu- 

tion. Chapman & Hall, New York, USA, 376 p. 

Anderson, R.C., A.G. Chabaud, S. Willmott. 1974-83. CIH keys to the 

nematode parasites of vertebrates. Commonwealth Agricultural Bureaux, 

Farnham Royal, Bucks, England, Numbers 1-10. 

Anonymous. 1976, 1981, 1989. Registry of marine pathology, volumes 1-3, 

NMFS Laboratory, NOAA, Oxford, Maryland, USA. 

_____ 1980. Proceedings of the 1979 sea lamprey International Symposium. 

Canadian Journal of Fisheries and Aquatic Sciences 37:1585-2214. 

*_____ 1992. Fishes of Puerto Rico [in Spanish and English]. 2nd Edition, 

Sea Grant Program, University of Puerto Rico, Mayaguez, Puerto Rico, 

UPRSG-E-43:38 p. 

*_____ 1994a. World Record Game Fishes. The International Game Fish 

Association, Pompano Beach, Florida, 352 p. 

_____ 1994b. Evidence for mixing based on parasites. Appendix E and 

Pages 139-145 In: An assessment of Atlantic bluefin tuna. National 

Academy Press, Washington, DC, USA, 148 p. 

Arandas-Rego, A. and C.P. Santos. 1983. Helminths of mackerel, Scomber 

japonicus, from Rio de Janeiro. Memórias do Instituto Oswaldo Cruz 

78:443-448. 

Barse, A.M. 1988. A contribution to the biology of white marlin, Tetrapturus 

albidus, off the Delaware and Maryland coast (USA) with special emphasis 

on feeding ecology and helminth parasites. MS Thesis, University of 

Maryland, 92 p. 

_____ and C.H. Hocutt. 1990. White marlin parasites: Potential indicators of 

stock separations, seasonal migrations, and feeding habits. Pages 41-49 

In: R.H. Stroud (Ed.) Proceedings of the Second International Billfish 

Symposium, Part 2. 

Bashirullah, A.K.M., N. Aguado, M. Alvarez, I.A. Marcano R. and J.J. 

Alio. 1995. Preliminary analysis of the parasites of billfishes captured in 

Venezuela [in Spanish]. Proceedings of the Gulf and Caribbean Fisheries 

Institute (in press).

BIBLIOGRAPHY 

_____ and J.C. Rodriguez. 1992. Spatial distribution and interrelationship of 

four monogenoidea of jack mackerel, Caranx hippos (Carangidae) in the 

north-east of Venezuela. Acta Cientifica Venezolana 43:125-128. 

Bane, G.W. 1969. Parasites of the yellowfin tuna, Thunnus albacares, in the 

Atlantic Ocean (Pisces: Scombridae). Wasmann Journal of Biology 27: 

163-175. 

Benz, G.W. 1984. Association of the pedunculate barnacle, Conchoderma 

virgatum (Spengler, 1790), with pandarid copepods (Siphonostomatoida: 

Pandaridae). Canadian Journal of Zoology 62:741-742. 

*_____ 1994a. Parasitic crustaceans: Back to basics. Pages 76-83 In: Regional 

Conference Proceedings, American Zoo and Aquarium Association, 

Wheeling, West Virginia, USA, 412 p. 

_____ 1994b. Host index to Yamaguti's Parasitic Copepoda and Branchiura of 

Fishes 1963. Journal of Aquatic Animal Health 6:162-175. 

Bere, R. 1936. Parasitic copepods from gulf of Mexico fish. American Midland 

Naturalist. 17:577-625. 

Berland, B. 1961. Nematodes from some Norwegian marine fishes. Sarsia 2: 1- 

50. 

Bliss, D.E. (Ed.). 1982-85. The biology of Crustacea. Volumes 1-10, Academic 

Press, New York, USA. 

Bravo-Hollis, M. 1953. Gill monogenea of Mexican coastal fishes [in Spanish]. 

Congreso Cientifico Mexicano Memoria 7:139-146. 

Brown, R.J. 1970. Pathology of pompano with whirling disease and Spanish 

mackerel with enteric cestodiasis. Proceedings of the World Mariculture 

Society 1:132-136. 

Bruce, N.L., R.D. Adlard and L.R.G. Cannon. 1994. Synoptic checklist of 

ascaridoid parasites (Nematoda) from fish hosts. Invertebrate Taxonomy 

8:583-674. 

_____ and L.R.G. Cannon. 1989. Hysterothylacium, Iheringascaris and 

Maricostula new genus, nematodes (Ascaridoidea) from Australian pelagic 

marine fishes. Journal of Natural History 23:1397-1441. 

Bunkley-Williams, L., W.G. Dyer and E.H. Williams, Jr. 1996. Some 

aspidogastrid and digenean trematodes of Puerto Rican marine fishes. 

Journal of Aquatic Animal Health 8:87-92. 

_____ and E.H. Williams, Jr. 1994. Diseases caused by Trichodina spheroidesi 

and Cryptocaryon irritans (Ciliophora) ciliates in wild coral reef fishes. 

_____ 6:360-361. 

_____ and _____ 1995. Parasites of Puerto Rican freshwater sport fish [in 

Spanish]. Department of Natural and Environmental Resources, San Juan, 

and Department of Marine Sciences, University of Puerto Rico, Mayaguez, 

Puerto Rico, 190 p. 

Burnett-Herkes, J. 1974. Parasites of the gills and buccal cavity of the dolphin, 

Coryphaena hippurus, from the straits of Florida. Transactions of the 

American Fisheries Society 103:101-106. 

345

346 


Caballero y Caballero, E., and M. Bravo-Hollis. 1965. Fish Monogenea from 

the Mexican Gulf and Caribbean coasts. I [in Spanish]. Bulletin of Marine 

Science of the Gulf and Caribbean 15:535-547. 

_____ and _____ 1967. Fish Monogenea (Van Beneden, 1858) Carns, 1863, 

from the Mexican Gulf and Caribbean coasts. III. Anales del Instituto de 

Biologia de Universidad Nacional Autonoma de Mexico 38:27-34. 

Cable, R.M. and J. Linderoth. 1963. Taxonomy of some acanthocephala from 

marine fish with reference to species from Curaçao, N.A., and Jamaica, 

W.I. Journal of Parasitology 49:706-716. 

_____ and B.A. Mafarachisi. 1970. Acanthocephala of the genus Gorgorhynchoides 

parasitic in marine fishes. H.D. Srivastava Commemorative Volume, 

p. 255-261. 

Causey, D. 1953a. Parasitic copepoda from Grand Isle, Louisiana. Louisiana 

State University Marine Laboratory Occasional Papers 7:18 p. 

_____ 1953b. Parasitic copepoda of Texas coastal fishes. Publications of the 

Institute of Marine Sciences 3:7-16. 

_____ 1955. Parasitic copepoda from Gulf of Mexico fish. Louisiana State 

University Marine Laboratory Occasional Papers 9:19 p. 

_____ 1960. Parasitic Copepoda from Mexican coastal fishes. Bulletin of 

Marine Sciences of the Gulf and Caribbean 10:323-337. 

Chandler, A.C. 1935. Parasites of fishes in Galveston Bay. Proceedings of the 

U.S. National Museum 83:123-157. 

_____ 1937. A new trematode Hirudinella beebi from the stomach of a 

Bermuda fish Acanthocybium petus. Transactions of the American 

Microscopical Society 56:348-354. 

_____ 1941. Two new trematodes from the bonito, Sarda sarda, in the Gulf of 

Mexico. Journal of Parasitology 27:183-184. 

_____ 1942. Some cestodes from Florida sharks. Proceedings of the U.S. 

National Museum 92(3135):25-31. 

*Cochran, P.A. 1989. Maintaining parasitic lampreys in closed laboratory 

systems. American Biology Teacher 51:115-119. 

Compagno, L.J.V. 1984. Hexanchiformes to Lamniformes. FAO Fisheries 

Synopsis No. 125, Volume 4. Sharks of the world, Part 1:249 p. 

Corkum, K.C. 1967. Bucephalidae (Trematoda) in fishes of the northern Gulf 

of Mexico: Bucephalus Baer, 1827. Transactions of the American Microscopical 

Society 86:44-49. 

_____ 1968. Bucephalidae (Trematoda) in fishes of the Northern Gulf of 

Mexico: Bucephaloides Hopkins, 1954 and Ripidocotyle Diesing, 1858. 

_____ 87:342-349. 

Crane, J. 1936. Notes on the biology and ecology of giant tuna, Thunnus 

thynnus Linnaeus, observed at Portland, Maine. Zoologica 21:207-212. 

Cressey, R.F. 1967. Genus Gloiopotes and a new species with notes on host 

specificity and intraspecific variation (Copepoda: Caligoida). Proceedings 

of the United States National Museum 122:1-21.

BIBLIOGRAPHY 

_____ 1975. A new family of parasitic copepods (Cyclopoida: Shiinoidae). 

Crustacea 28:211-219. 

_____ 1981,1983,1991. Parasitic copepods from the Gulf of Mexico and 

Caribbean Sea, I. Holobomolochus and Neobomolochus. II: Bomolochidae. 

III: Caligus. Smithsonian Contributions to Zoology 339:1-24; 389: 1- 

35; 497:1-53. 

_____ and B.B. Collette. 1970. Copepods and needlefishes: a study in hostparasite 

relationships. Fishery Bulletin 68:347-432. 

_____, B.B. Collette and J.L. Russo. 1983. Copepods and scombrid fishes: A 

study in host-parasite relationships. _____ 81:227-265. 

_____ and H.B. Cressey. 1980. Parasitic copepods of mackerel- and tuna-like 

fishes (Scombridae) of the world. Smithsonian Contributions to Zoology 

311:1-186. 

_____ and E.A. Lachner. 1970. The parasitic copepod diet and life history of 

diskfishes. Copeia 1970:310-318. 

_____ and P. Nutter. 1987. Reidentification of David Causey's Caligus 

collections (Crustacea: Copepoda). Proceedings of the Biological Society 

of Washington 100:600-602. 

Crites, J.L., R.M. Overstreet and M. Maung. 1993. Ctenascarophis lesteri n. sp. 

and Prospinitectus exiguus n. sp. (Nematoda: Cystidicolidae) from the 

skipjack tuna, Katsuwonus pelamis. Journal of Parasitology 79:847-859. 

Cuvier, G. 1830. The Animal Kingdom according to their organization in order 

to serve as a basis for the natural history of the animals and the 

introduction to their comparative anatomy [in French]. Nouvelle Edition, 

Volume 3, Déterville, Paris, 504 p. 

Davies, A.J. 1982. Further studies on Haemogregarina bigemina Lavern & 

Mesnil, the marine fish Blennius pholis L., and the Isopod Gnathia 

maxillaris Montagu. Journal of Protozoology 29:576-583. 

Dawes, B. 1968. Trematoda. Cambridge University Press, London, 644 p. 

Deardorff, T.L. and R.M. Overstreet. 1980. Taxonomy and biology of North 

American species of Goezia (Nematoda: Anisakidae) from fishes, 

including three new species. Proceedings of the Helminthological Society 


_____ and _____ 1981a. Review of Hysterothylacium and Iheringascaris (both 

previously = Thynnascaris) (Nematoda: Anisakidae) from the northern 

Gulf of Mexico. Proceedings of the Biological Society of Washington 

93:1035-1079. 

_____ and _____ 1981b. Larval Hysterothylacium (Thynnascaris)(Nematoda: 

Anisakida) from fishes and Invertebrates in the Gulf of Mexico. Proceedings 

of the Helminthological Society of Washington 48:113-126. 

_____ and _____ 1982. Hysterothylacium pelagicum sp. n. and H. cornutum 

(Nematoda: Anisakida) from marine fishes. _____ 49:246-251. 

_____ and _____ 1986. Piscine adult nematode invading an open lesion in a 

human hand. American Journal of Tropical Medicine and Hygiene 35: 

827-830. 

347

348 


_____ and _____ 1991. Seafood-transmitted zoonoses in the United States [of 

America]: The fishes, the dishes, and the worms. Pages 211-265 In: D. R. 

Ward and C. R. Hackney (Eds.). Microbiology of marine food products. 

Van Nostrand Reinhold, New York, USA, 450 p. 

Delamare-Deboutteville, C. and L. Nunes-Ruivo. 1958. Copepods parasitic on 

Mediterranean fishes [in French]. Vie Milieu 9:215-235. 

Delaney, P.M. 1984. Isopods of the genus Excorallana Stebbing, 1904 from 

the Gulf of California, Mexico (Crustacea, Isopoda, Corallanidae). Bulletin 

of Marine Science 34:1-20. 

Dojiri, M. 1983. Revision of the genera of the Caligidae (Siphonostomatoida), 

copepods predominantly parasitic on marine fishes. PhD Dissertation, 

Boston University, 721 p. 

Dyer, W.G., E.H. Williams, Jr., and L.B. Williams. 1985. Digenetic trematodes 

of marine fishes of the western and southwestern coasts of Puerto Rico. 

Proceedings of the Helminthological Society of Washington 52:85-94. 

_____, _____ and _____ 1986. Some trematodes of marine fishes of 

southwestern and northwestern Puerto Rico. Transactions of the Illinois 

Academy of Sciences 79:141-143. 

_____, _____ and L.B. Williams. 1988. Digenetic trematodes of marine fishes 

of Okinawa, Japan. Journal of Parasitology 74:638-645. 

_____, _____ and _____ 1992a. Neobenedenia pargueraensis n. sp. (Monogenea: 

Capsalidae) from the red hind, Epinephelus guttatus, in Puerto 

Rico. _____ 78:330-333. 

_____, _____ and _____ 1992b. Tristomella laevis (Verrill, 1875) Guiart, 1938 

(Monogenea: Capsalidae) on white and blue marlins from the 

southwestern coast of Puerto Rico and Desecheo Island. Transactions of 

the Illinois State Academy of Science 85:183-185. 

_____, _____ and _____ 1992c. Homalometron dowgialloi sp. n. 

(Homalometridae) from Haemulon flavolineatum and additional records 

of digenetic trematodes of marine fishes in the West Indies. Journal of the 

Helminthological Society of Washington 59:182-189. 

_____, _____ and _____ 1997. Helminths of the dolphinfish (Coryphaena 

hippurus) in Puerto Rico. _____ (in press). 

Eggleston, D.B. and E.A. Bochenek. 1990. Stomach contents and parasite 

infestation of school bluefin tuna Thunnus thynnus collected from the 

middle Atlantic Bight, Virginia. Fishery Bulletin 88:389-395. 

Eldridge, M.B. and P.G. Wares. 1974. Some biological observations of 

billfishes taken in the eastern Pacific Ocean, 1967-1970. NOAA Technical 

Report NMFS SSRF-675(2):89-101. 

Fernandes, B.M.M. 1970. Occurrence of Dinurus tornatus (Rudolphi, 1819) 

Loss, 1907 in Brasil [in Portuguese]. Atas da Sociedade de Biologia do 

Rio de Janeiro 14:91-92.

BIBLIOGRAPHY 

_____, A. Kohn and R. Magalhães-Pinto. 1985. Aspidogastrid and digenetic 

trematode parasites of marine fishes of the coast of Rio de Janeiro State, 

Brazil. Revista Brasileira de Biologia 45:109-116. 

Finlayson, J.E. 1982. The alleged alternation of sexual phases in Kuhnia 

scombri, a monogenean of Scomber scombrus. Parasitology 84:303-311. 

Fischer, W. (Ed.). 1978. FAO Species Identification Sheets for Fisheries 

Purposes, Western Central Atlantic (Fishing Area 31). Volumes 1-7, FAO 

United Nations, Rome, Italy. 

Fischthal, J.H. 1977. Some digenetic trematodes of marine fishes from the 

barrier reef and lagoon of Belize. Zoologica Scripta 6:81-88. 

Franks, J.S. 1995. A pugheaded cobia (Rachycentron canadum) from the northcentral 

Gulf of Mexico. Gulf Research Reports 9:143-145. 

Garzon-Ferreira, J. 1990. An isopod, Rocinela signata (Crustacea: Isopoda: 

Aegidae), that attacks humans. Bulletin of Marine Science 46:813-815. 

Gibson, D.I. and R.A. Bray. 1977. The Azygiidae, Hirudinellidae, Ptychogonimidae, 

Sclerodistomidae and Syncoeliidae (Digenea) of fishes from the 

northeast Atlantic. Bulletin of the British Museum (Natural History) 

Zoology Series 32:167-245. 

*Goldstein, R.J. 1988. Offshore fishing from Virginia to Texas. John F. Blair, 

Winston-Salem, North Carolina, USA, 248 p. 

*_____ 1987. Billfish parasites. Sport Fishing (Oct/Nov):58-64. 

Grabda, J. 1991. Marine fish parasitology. (English translation, 1981 Polish 

publication) VCH Verlagsgesellschaft, Weinheim, Germany. 306 p. 

Guitart-Manday, D. 1964. Fishery biology of the swordfish, Xiphias gladius 

Linnaeus (Teleostomi: Xiphiidae), in Cuban waters [in Spanish]. Poeyana, 

Ser. B(1):37 p. 

*Hammond, D.L. and D.M. Cupka. 1975. A sportsman's field guide to the 

billfishes, mackerels, little tunas and tunas of South Carolina. South 

Carolina Marine Resources Department, Educational Report 3:32 p. 

Harding, J.P. 1966. Myodocopan ostracods from the gills and nostrils of fishes. 

Pages 369-374 In: H. Barnes (Ed.) Contemporary studies in marine 

science, George Allen and Unwin Ltd., London, England. 

Hargis, W.J., Jr. 1955,1957. Monogenetic trematodes of Gulf of Mexico fishes. 

V. The superfamily Capsaloidea. XIII. The family Gastrocotylidae Price, 

1943. Transactions of the American Microscopical Society 74:203-225; 

76:1-12. 

_____ 1956a. Monogenetic trematodes of Gulf of Mexico fishes. XI. The 

family Microcotylidae Taschenberg, 1879. Proceedings of the Helminthological 

Society of Washington 23:153-162. 

_____ 1956b. Monogenetic trematodes of Gulf of Mexico fishes. XII. The 

family Gastrocotylidae Price, 1943. Bulletin of Marine Sciences of the 

Gulf and Caribbean 6:28-43. 

_____ 1958. The fish parasite Argulus laticauda as a fortuitous human epizoon. 

Journal of Parasitology 44:45. 

349

350 


Harshbarger, J.C. 1965-1981 (each year). Activities report of the Registry of 

Tumors in Lower Animals, National museum of Natural History, 

Smithsonian Institution, Washington, District of Columbia, USA. 

Hasegawa, H., E.H. Williams, Jr. and L. Bunkley-Williams. 1991. Nematode 

parasites from marine fishes of Okinawa, Japan. Journal of the 


Hauck, K. 1977. Occurrence and survival of the larval nematode Anisakis sp. in 

the flesh of fresh, frozen, brined, and smoked pacific herring, Clupea 

harengus Pallasi. Journal of Parasitology 63:515-519. 

Health, M.R. 1992. Field investigations of the early life stages of marine fish. 

Advances in Marine Biology 28:1-174. 

Hendrix, S.S. 1994. Platyhelminthes: Monogenea. NOAA Technical Report 

NMFS 121:107 p. 

*Hilderbrand, K.S., Jr. 1984. Parasites in marine fishes: Questions and answers 

for seafood retailers. Oregon State University Sea Grant 79:2 p. 

Hogans, W.E. 1985. Occurrence of Caligus coryphaenae (Copepoda: 

Caligidae) on the Atlantic bluefin tuna (Thunnus thynnus L.) from Prince 

Edward Island, Canada. Crustaceana 49:313-314. 

_____ 1986. Redescription of Pennnella instructa Wilson, 1917 (Copepoda: 

Pennellidae) from the swordfish (Xiphias gladius L.). Canadian Journal of 

Zoology 64:727-730. 

_____ 1987. Morphological Variation in Pennella balaenoptera and P. filosa 

(Copepoda: Pennellidae) with a Review of the Genus Pennella Oken, 1816 

Parasitic on Cetacea. Bulletin of Marine Science 40:442-453. 

_____ 1988a. Pennella makaira, new species (Copepoda: Pennellidae) from 

the Atlantic Blue Marlin, Makaira nigricans, in the Caribbean Sea. 

Proceedings of the Biological Society of Washington 101:15-19. 

_____ 1988b. Redescription of Pennella sagitta (Copepoda: Pennellidae) from 

Histrio histrio (Pisces) in the north-west Atlantic Ocean with a provisional 

review of the genus Pennella. Journal of Zoology (London) 216: 379-390. 

_____ 1995. Parasitic Copepoda in the collection of the Atlantic Reference 

Centre, St. Andrews, New Brunswick, Canada. Canadian Technical Report 

of Fisheries and Aquatic Sciences No. 2028:6 p. 

_____ and J. Brattey. 1982. Parasites of the gills and gastrointestinal tracts of 

swordfish (Xiphias gladius) from the northwest Atlantic Ocean, with an 

assessment of their use as biological tags. Report to Canada Department 

of Fisheries and Oceans No. 07SC.FP706-1-CO33:36 p. 

_____, _____ and T.R. Hurlbutt. 1986. Pennella filosa and Pennnella instructa 

(Copepoda: Pennellidae) on swordfish (Xiphias gladius L.) from the 

northwest Atlantic. Journal of Parasitology 71:111-112. 

_____, _____, L.S. Uhazy and P.C.F. Hurley. 1983. Helminth parasites of 

swordfish (Xiphias gladius L.) from the northwest Atlantic Ocean. _____ 

69:1178-1179.

BIBLIOGRAPHY 

_____ and P.C.F. Hurley. 1986. Variations in the Morphology of Fistulicola 

plicatus Rudolphi (1802) (Cestoda: Pseudophyllidea) from the Swordfish, 

Xiphias gladius L., in the northwest Atlantic Ocean. Fishery Bulletin 84: 

754-757. 

Holliday, M. 1978. Food of Atlantic bluefin tuna, Thunnus thynnus (L.), from 

the coastal waters of North Carolina to Massachusetts. MS Thesis, Long 

Island University, New York, USA, 27 p. 

Howse, H.D., J.S. Franks and R.F. Welford. 1975. Pericardial adhesions in 

cobia Rachycentron canadum (Linnaeus). Gulf Research Reports 5:61-62. 

Iles, C. 1971. Fistulicola plicatus (Cestoda) and Tristoma spp. (Trematoda) on 

swordfish from the Northwest Atlantic. Journal of the Fisheries Research 

Board of Canada 28:31-34. 

Iversen, E.S. and N.N. Van Meter. 1967. A new Myxosporidian (Sporozoa) 

infecting the spanish mackerel. Bulletin of Marine Sciences 17:268-273. 

_____ and H.O. Yoshida. 1957. Notes on the biology of the wahoo in the Line 

Islands. Pacific Science 11:370-379. 

Iversen, R.T.B. and R.R. Kelley. 1974. Occurrence, morphology, and parasitism 

of gastric ulcers in blue marlin, Makaira nigricans, and black 

marlin, Makaira indica, from Hawaii. NOAA Technical Report NMFS 

SSRF-675(2):149-153. 

Jahn, T.L., E.C. Bovee and F.F. Jahn. 1979. How to Know the Protozoa. 2nd 

Edition, William C. Brown Company, Dubuque, Iowa, 279 p. 

Johnson, G.D. 1984. Percoidei: development and relationships. Pages 464-498 

In: H.G. Moser et al. (Eds.) Ontogeny and systematics of fishes. American 

Society of Ichthyology and Herpetology, Special Publication 1. 

*Johnson, S.K. 1977. Worm parasites of the edible flesh of Texas coastal 

fishes. Texas Agricultural Extension Service, FDDL-M1:6 p. 

*_____ 1977. Fish worms harmless to humans. Texas Trawler Jan.-Feb. 

1977:3-4. 

Johnstone, J. 1912. Tetrarhynchus erinaceus van Beneden. I. Structure of larva 

and adult worm. Parasitology 4:364-415. 

Jolley, J.W., Jr. 1977. The biology and fishery of Atlantic sailfish, Istiophorus 

platypterus, from southeast Florida. Florida Marine Resources Publication 

No. 28:31 p. 

Jones, E.C. 1971. Isistius brasiliensis, a squaloid shark, the probable cause of 

crater wounds on fishes and cetaceans. Fishery Bulletin 69:791-798. 

Jones, J.B. 1991. Movements of albacore tuna (Thunnus alalunga) in the south 

Pacific: evidence from parasites. Marine Biology 111:1-9. 

Jones, S. 1962. The phenomenal fish mortality in the Arabian Sea in 1957--A 

speculation on the possible identity of the species concerned. Symposium 

on Scombroid Fishes, Marine Biological Association of India, Part II:713- 

718. 

Justine, J.-L., A. Lambert and X. Mattei. 1985. Spermatozoon ultrastructure 

and phylogenetic relationships in the monogeneans (Platyhelminthes). 

International Journal of Parasitology 15:601-608. 

351

352 


_____ and X. Mattei. 1987. Phylogenetic relationships between the families 

Capsalidae and Dionchidae (Platyhelminthes, Monogenea, Monopisthocotylea) 

indicated by the comparative ultrastructural study of spermiogenesis. 

Zoologica Scripta 16:111-116. 

*Kabata, Z. 1970. Crustacea as enemies of fishes. T.F.H. Publications, Inc. 

Neptune City, New Jersey, USA. 171 p. 

_____ 1979. Parasitic Copepoda of British Fishes. Ray Society, London, 

England, 468 p., 2030 figs. 

_____ 1988. Copepoda and Branchiura. Pages 4-127 In L. Margolis and Z. 

Kabata [Eds.] Guide to the parasites of fishes of Canada. Part II Crustacea. 

Canadian Special Publications of Fisheries and Aquatic Sciences 101:184 

p. 

Kawahara, E., J.S. Nelson and R. Kusuda. 1986. Fluorescent antibody 

technique compared to standard media culture for detection of pathogenic 

bacteria for yellowtail and amberjack. Fish Pathology 21:39-45. 

Khalil, L.F., A. Jones and R.A. Bray (Eds.). 1994. Keys to the Cestode 

Parasites of Vertebrates. CAB International, Oxford, England, 751 p. 

Kensley, B. and M. Schotte. 1989. Marine isopod crustaceans of the Caribbean. 

Smithsonian Institution Press, Washington, DC, 308 p. 

*Klemm, R. 1984. Cookie cutter shark. Sea Frontiers 28(2):9. 

Koratha, K.J. 1955. Studies on the monogenetic trematodes of the Texas coast. 

II. Descriptions of species from marine fishes of Port Aransas. University 

of Texas Institute of Marine Science Publication 4:251-278. 

Lee, J.J., S.H. Hutner and E.C. Bovee (Eds.). 1985. An Illustrated Guide to the 

Protozoa. Academic Press, Lawrence, Kansas, 629 p. 

Lester, R.J.G., A. Barnes and G. Habib. 1985. Parasites of skipjack tuna, 

Katsuwonus pelamis: Fishery implications. Fishery Bulletin 83:343-356. 

Lewis, A.G. 1967. Copepod crustaceans parasitic on teleost fishes of the 

Hawaiian Islands. Proceedings of the U.S. National Museum 121:1-204. 

Llewellyn, J. 1956. The host-specificity, micro-ecology, adhesive attitudes, and 

comparative morphologies of some trematode gill parasites. Journal of the 

Marine Biology Association of the United Kingdom 35:113-127. 

_____ 1957. The mechanism of the attachment of Kuhnia scombri (Kuhn, 

1829)(Trematoda: Monogenea) to the gills of its host Scomber scombrus 

L., including a note on the taxonomy of the parasite. Parasitology 47:30- 

39. 

Linton, E. 1897a. Notes on larval cestode parasites of fishes. Proceedings of the 

U.S. National Museum 19:787-824, 8 pl. 

_____ 1897b. Notes on cestode parasites of fishes. _____ 20:423-456, 8 pl. 

_____ 1900. Fish parasites collected at Woods Hole in 1898. Bulletin of the 

U.S. Commission on Fish and Fisheries 19:267-304. 

_____ 1901. Parasites of fishes of the Woods Hole region. ____ 19:405-492. 

_____ 1905. Parasites of fishes of Beaufort, North Carolina. Bulletin of the 

U.S. Bureau of Fisheries for 1904. 24:321-428, 34 pl.

BIBLIOGRAPHY 

_____ 1907. Notes on parasites of Bermuda fishes. Proceedings of the U.S. 

National Museum 33:85-126. 

_____ 1908,1910. Helminth fauna of the Dry Tortugas. I. Cestodes. II. 

Trematodes. Carnegie Institution of Washington Publication 102:157-190; 

133:11-98. 

_____ 1924. Notes on cestode parasites of sharks and skates. Proceedings of 

the U.S. National Museum 64:1-114, 13 pl. 

_____ 1940. Trematodes from fishes mainly from the Woods Hole region, 

Massachusetts. _____ 88:1-172. 

_____ 1941. Cestode parasites of teleost fishes of the Woods Hole region, 

Massachusetts. _____ 90:417-441, 3 pl. 

Lom, J. and I. Dyková. 1992. Protozoan Parasites of Fishes. Elsevier Science 

Publishers, Amsterdam, Netherlands, 315 p. 

Love, M.S. and M. Moser 1983. A checklist of parasites of California, Oregon, 

and Washington marine and estuarine fishes. NOAA Technical Report 

NMFS SSRF-777:576 p. 

MacCallum, G.A. 1913. Notes on four trematodes parasites of marine fishes. 

Centralblatt fur Bakteriologie Parasitenkunde und Infektionskrankheiten., 

1 Abteilung Originale 70:407-416. 

_____ 1915. Some new species of ectoparasitic trematodes. Zoologica 1:393- 

410. 

_____ 1917. Some new forms of parasitic worms. Zoopathologica 1:43-75. 

_____ 1921. Studies in helminthology. _____ 1:137-284. 

_____ and W.G. MacCallum. 1916. The family Koellikeriadae (Didymozoidae). 

Zoologische Jahrbuecher Abteilung fuer Systematik 39:141-168. 

Madhavi, R. 1982. Didymozoid trematodes (including new genera and species) 

from marine fishes of the Waltair coast, Bay of Bengal. Systematic 

Parasitology 4:99-124. 

*McClane, A.J. (Ed.). 1974a,b. McClane's field guide to salt water fishes of 

North America. [and] McClane's New Standard Fishing Encyclopedia and 

International Angling Guide. 2nd Edition, Holt, Rinehart and Winston, 

New York, USA, 283 p.(a), 1156 p.(b) 

McIntosh, A. 1934. A new blood trematode, Paradeontacylix sanguinicoloides 

n. g., n. sp. from Seriola lalandi, with a key to the species of the Family 

Aporocotylidae. Parasitology 26:463-467. 

McKenzie, K. 1983. The selection of parasites for use as biological tags in 

population studies of bluefin tuna. ICCAT, Collected Volume of Scientific 

Papers 18:834-838. 

McMahon, J.W. 1964. Monogenetic trematodes from some Chesapeake Bay 

fishes. Part II: the superfamily Diclidophoroidea. Chesapeake Science 5: 

124-133. 

Mamaev, Y.L. 1982. Monogeneans of the subfamily Grubeinae Price, 1961 

(family Mazocraeidae) [in Russian]. Parazitology 16:457-463. 

Manooch, C.S., III and W.T. Hogarth. 1983. Stomach contents and giant 

trematodes from wahoo Acanthocybium solanderi collected along the 

353

354 


south Atlantic and Gulf coasts of the United States. Bulletin of Marine 

Sciences 33:227-238. 

_____, D.L. Mason and R.S. Nelson. 1984. Food and gastrointestinal parasites 

of dolphin Coryphaena hippurus collected along the southeastern and Gulf 

coasts of the United States [of America]. Bulletin of the Japanese Society 

of Scientific Fisheries 50:1511-1525. 

Manter, H.W. 1931. Some digenetic trematodes of marine fishes of Beaufort, 

North Carolina. Parasitology 23:396-411. 

_____ 1940a. Digenetic trematodes of fishes from the Galapagos Islands and 

the neighboring Pacific. Alan Hancock Pacific Expeditions, Volume 2: 

329-497. 

_____ 1940b. Gasterostomes (Trematoda) of Tortugas, Florida. Papers from 

the Tortugas Laboratory of the Carnegie Institution of Washington 33:1- 

19. 

_____ 1947. The digenetic trematodes of marine fishes of Tortugas, Florida. 

American Midland Naturalist 38:257-416. 

_____ 1970. A new genus of trematode (Digenea; Gorgoderidae) from the 

ureter of tuna fish (Thunnus thynnus maccoyii) in Australia. Transactions 

of the Royal Society of South Australia 94:147-150. 

Margolis, L., Z Kabata, and R.R. Parker. 1975. Catalogue and synopsis of 

Caligus, a genus of Copepoda (Crustacea) parasitic on fishes. Bulletin of 

the Fisheries Research Board of Canada 192:117 p. 

_____, J.O. Corlis, M. Melkonian, D.J. Chapman (Eds). 1989. Handbook of 

Protoctista. Jones and Bartlett, Boston, USA, 914 p. 

Menzies, R.J. and P.W. Glynn. 1968. The common marine isopod crustacea 

of Puerto Rico. Studies on the fauna of Curaçao and other 

Caribbean Islands 27:1-110. 

Mignucci-Giannoni, A.A. 1996. Marine mammal strandings in Puerto Rico and 

the United States and British Virgin Islands. PhD Dissertation, University 

of Puerto Rico, Mayagüez, Puerto Rico, 247 p. 

Millemann, R.E. 1956. Notes on the genus Hexostoma (Monogenea: Hexostomatidae) 

with a redescription of H. euthynni Meserve, 1938. Journal of 

Parasitology 42:316-319. 

Murchelano, R.A., L. Despres-Patanjo and J. Ziskowski. 1986. A histopathologic 

evaluation of gross lesions excised from commercially important 

North Atlantic marine fishes. NOAA Technical Report NMFS 37:14 p. 

Muroga, K., T. Nakai and T. Nishizawa. 1994. Viral diseases in Japanese 

seawater pen culture. Page V-2 In: International Symposium on Aquatic 

Animal Health, Abstracts, Seattle, Washington, USA, 298 p. 

Nahhas, F.M. 1993. Some acanthocephala and Digenea of Marine fish from 

Grand Cayman, Cayman Islands, British West Indies. Journal of the 


_____ and R.M. Cable. 1964. Digenetic and aspidogastrid trematodes from 

marine fishes of Curaçao and Jamaica. Tulane Studies in Zoology and 

Botany 11:167-228.

BIBLIOGRAPHY 

_____ and K. Carlson. 1994. Digenetic trematodes of marine Fishes of 

Jamaica, West Indies. Hofstra University Marine Laboratory, Ecological 

Survey of Jamaica Publication 2:60 p. 

_____ and R.B. Short. 1965. Digenetic trematodes of marine fishes from 

Apalachee Bay, Gulf of Mexico. Tulane Studies in Zoology and Botany 

12:39-50. 

Nasir, P. and J.L. Fuentes Zambrano. 1983. Some Venezuelan monogeneans 

[in Spanish]. Rivista di Parassitologia 44:335-380. 

Neish, G.A. and G.C. Hughes. 1980. Fungal Diseases of Fishes. TFH Publications, 

Neptune City, New Jersey, 159 p. 

Nigrelli, R.F. 1938. Parasites of the swordfish Xiphias gladius L. American 

Museum Novitates 996:1-16. 

_____ 1939. Didymocystis coatesi, a new monostome from the eye muscles of 

the wahoo, Acanthocybium solanderi (C. and V.). Transactions of the 

American Microscopical Society 58:170-178. 

_____ and H.W. Stunkard. 1947. Studies on the genus Hirudinella, giant 

trematodes of scombriform fishes. Zoologica 31:185-196. 

Nikolaeva, V.M. 1968. Studies of the helminth fauna of Thunnus albacroes and 

Histiophoridae in the Gulf of Mexico [in Russian]. Pages 150-157 In: Z. 

B. Yankovskaya (Ed.) (Studies of Central American Seas). Kiev: Nankova 

Dunka (2). 

_____ 1985. Trematodes--Didymozoidae fauna, distribution and biology. 

Pages 67-73 In: W.J. Hargis, Jr. (Ed.) Parasitology and pathology of 

marine organisms of the world ocean. NOAA Technical Report NMFS 

25:135 p. 

_____ and A.M. Parukhin. 1968. Study of the helminths of fish in the Gulf of 

Mexico [in Russian]. Pages 126-149 In: Z.B. Yankovskaya (Ed.) (Studies 

of Central American Seas). Kiev: Nankova Dunka (2). 

Norris, D.E. and R.M. Overstreet. 1976. The public health implications of 

larval Thynnascaris nematodes from shellfish. Journal of Milk and Food 

Technology 39:47-54. 

Olsen, L.S. 1952. Some nematodes parasitic in marine fishes. Publications of 

the Institute of Marine Science, University of Texas 2:173-215. 

Ortiz, E.A.R., E.H. Williams, Jr., and L. Bunkley-Williams. 1995. A record of 

paper nautilis (Argonauta argo and Argonauta hians) in Puerto Rico. 

Caribbean Journal of Science 31:340-341. [in a dolphin stomach] 

Overstreet, R.M. 1969. Digenetic trematodes of marine teleost fishes from 

Biscayne Bay, Florida. Tulane Studies in Zoology and Botany 15:119-176. 

*_____ 1978. Marine maladies? Worms, germs, and other symbionts from the 

Northern Gulf of Mexico. Mississippi-Alabama Sea Grant Consortium 

MASGP-78-021:140 p. 

_____ and G.M. Meyer. 1981. Hemorrhagic lesions in stomach of rhesus 

monkey caused by a piscine ascaridoid nematode. Journal of Parasitology 

67:226-235. 

355

356 


Palko, B. J., G. L. Beardsley and W. J. Richards. 1982. Synopsis of the 

biological data on dolphin-fishes, Coryphaena hippurus Linnaeus and 

Coryphaena equiselis Linnaeus. NOAA Technical Report, NMFS Circular 

443, FAO Fisheries Synopsis 130: 28 p. 

Palm, H. W. 1995. Study of the systematics of tapeworms (Cestoda: Trypanorhyncha) 

of Atlantic fishes. [in German]. Berichte aus dem Institut für 

Meereskunde an der Christian-Albrechts-Universität, Kiel, Nr. 275:238p. 

Parker, R.R. 1969. Validity of the binomen Caligus elongatus for a common 

parasitic copepod formerly misidentified with Caligus rapax. Journal of 

the Fishery Research Board of Canada 26:1013-1035. 

Pearse, A.S. 1949. Observations of flatworms and nemerteans collected at 

Beaufort, N. C. Proceedings of the U.S. National Museum 100:25-38. 

_____ 1951. Parasitic crustacea from Bimini, Bahamas. _____ 101:341-372. 

_____ 1952a. Parasitic crustacea from the Texas coast. Institute of Marine 

Science, Publication 2:5-42, 151 figs. 

_____ 1952b. Parasitic Crustaceans From Alligator Harbor, Florida. Quarterly 

Journal of the Florida Academy of Sciences 15:187-243. 

Perez-Vigueras, I. 1935. Tristomum poeyi n. sp. (Trematoda) a parasite of 

Makaira ampla Poey (Pisces) [in Spanish]. Memorias de la Sociedad 

Cubana de Historia Natural 9:43-44. 

_____ 1942. Helminthological notes [in Spanish]. Revista de l'Universidad de 

Habana (40-42):193-223. 

_____ 1958. Additions to the Cuban helminth fauna [in Spanish]. Memorias de 

la Sociedad Cubana de Historia Natural 24:17-38. 

Pillai, N.K. 1971. Notes on some copepod parasites in the collection of the 

British Museum (N.H.), London. Journal of the Marine Biological Association 

of India 11:149-174. 

Pinkus, G.S., C. Coolidge and M.D. Little. 1975. Intestinal aniskiasis: First case 

report from North America. American Journal of Medicine 59:114-120. 

Plumb, J.A. 1994. Health maintenance of cultured fishes: principal microbial 

diseases. CRC Press Boca Raton, Florida, USA, 254 p. 

Price, E.W. 1938. North American monogenetic trematodes. II. The families 

Monocotylidae, Microbothridae, Acanthocotylidae and Udonellidae (Capsaloidea). 

Journal of the Washington Academy of Science 28:109-126. 

_____ 1939. North American monogenetic trematodes III. The family Capsalidae 

(Capsaloidea). Journal of the Washington Academy of Science 29: 63- 

92. 

_____ 1951. A new North American Monogenetic trematode Capsala manteri, 

n. sp. Proceedings of the Helminthological Society of Washington 18:24- 

25. 

_____ 1960. The giant marlin, Makiara marlina Jordan and Evermann, a new 

host for Capsala pricei Hidalgo, 1959, with a review of the subfamily 

Capsalinae. Libro. Homenaje al Dr. E. Caballero y Caballero, Mexico, p. 

237-244.

BIBLIOGRAPHY 

_____ 1961. North American monogenetic trematodes. VIII. The family 

Hexostomatidae. Proceedings of the Helminthological Society of Washington 

29:1-18. 

_____ 1962. North American monogenetic trematodes. X. The family Heteraxinidae. 

Journal of Parasitology 48:402-418. 

*Prince, E.D. 1984. Don't throw back recaptured tagged billfish or tuna--SAVE 

IT FOR SCIENCE. Marlin Magazine 3(2):50-54. 

Ragan, M.A., C.L. Googin, R.L. Cawthorn, L. Cerenius, A.V.C. Jamieson, 

S.M. Plourde, T.G. Rand, K. Söderhall and R.R. Gutell. 1996. A novel 

clade of protistan parasites near the animal-fungal divergence. Proceedings 

of the National Academy of Sciences 93:(in press). 

Raju, G. 1960. A case of hermaphroditism and some other gonadal abnormalities 

in the skipjack Katsuwonus pelamis (Linnaeus). Journal of the 

Marine Biological Association of India 2:95-102, 5 figs. 

Rand, T.G. 1996. Fungal diseases of fish and shellfish. Pages 297-313 In: 

Howard/Miller (Eds.) The Mycota VI. Human and animal relationships. 

Springer-Verlag, New York, USA, 399 p. 

Raptopoulou, F.A. and R.H. Lambertsen. 1987. Parasite-associated pathology 

of the dolphinfish, Coryphaena hippurus L., from Florida waters. Journal 

of Fish Diseases 10:379-384. 

Rees, G. 1969. Cestodes from Bermuda fishes and an account of Acompsocephalum 

tortum (Linton, 1905) gen. nov. from the lizard fish Synodus 

intermedius (Agassiz). Parasitology 59:519-548. 

_____ 1970. Some helminth parasites of fishes of Bermuda and an account of 

the attachment organ of Alcicornis carangis MacCallum, 1917 (Digenea: 

Bucephalidae). _____ 60:195-221. 

Robins, C.R., R.M. Bailey, C.E. Bond, J.R. Brooker, E.A. Lachner, R.N. Lea 

and W.B. Scott. 1991. Common and Scientific Names of Fishes from the 

United States and Canada. 5th Edition, American Fisheries Society, 

Special Publication 20:183 p. 

Rohde, K. 1978. Monogenea of Australian Marine Fishes. The Genera 

Dionchus, Sibitrema and Hexostoma. Seto Marine Biological Laboratory 

24:349-367. 

_____ 1994. Ecology of marine parasites. CAB International, Wallingford, 

Berkshire, England, 298 p. 

_____ 1987. Grubea australis n. sp. (Monogenea, Polyopisthocotylea) from 

Scomber australasicus in southeastern Australia, and Grubea cochlear 

Diesing, 1858 from S. scombrus and S. japonicus in the Mediterranean and 

western Atlantic. Systematic Parasitology 9:29-38. 

_____ 1989. Kuhnia sprostonae Price, 1961 and K. scombercolias Nasir & 

Fuentes Zambrano, 1983 (Monogenea: Mazocraeidae) and their microhabitats 

on the gills of Scomber australasicus (Teleostei: Scombridae), and 

the geographic distribution of seven species of gill Monogenea of Scomber 

spp. Systematic Parasitology 14:93-100. 

357

358 


_____ 1991. Size differences in hamuli of Kuhnia scombri (Monogenea, 

Polyopisthocotylea) from different geographical areas not due to 

differences in host size. International Journal of Parasitology 21:113-114. 

_____ and N. Watson. 1985. Morphology, microhabitats and geographic 

variation of Kuhnia spp. (Monogenea, Polyopisthocotylea). International 


Rose, C.D. 1966. The biology and catch distribution of the dolphin, Coryphaena 

hippurus (Linnaeus), in North Carolina waters. PhD Dissertation, 

University of North Carolina, 153 p. 

Ruszkowski, J.S. 1932. Studies of the life cycle and structure of marine 

cestodes. The life cycle of the tetrarhynchid Grillotia erinaceus (van Ben., 

1858) [in French]. Académie Polonaise des Sciences et des Lettres, 

Comptes Rendus Mensuels des Séances Mathématiques et Naturelles, 

Cracovie (9):6. 

Saunders, D.C. 1958. Blood parasites of the marine fishes of the Florida Keys. 

Yearbook of the American Philosophical Society, p. 261-266. 

_____ 1958. The occurrence of Haemogregarina bigemina Laveran and 

Mesnil and H. dasyatis n. sp. in marine fish from Bimini, B.W.I. 

Transactions of the American Microscopical Society 77:404-412. 

_____ 1959. Haemogregarina bigemina Laveran and Mesnil from marine 

fishes of Bermuda. _____ 78:374-379. 

_____ 1964. Blood parasites of marine fish of southwest Florida, including a 

new haemogregarine from the menhaden, Brevoortia tyrannus (Latrobe). 

_____ 83:218-225. 

_____ 1966. A survey of the blood parasites of the marine fishes of Puerto 

Rico. _____ 85:193-199. 

*Schell, S.C. 1970. How to Know the Trematodes. William C. Brown 

Company Publishers, Dubuque, Iowa, 355 p. 

_____ 1985. Handbook of Trematodes of North America North of Mexico. 

University Press of Idaho, Moscow, Idaho, 263 p. 

*Schmidt, G.D. 1970. How to Know the Tapeworms. William C. Brown 

Company Publishers, Dubuque, Iowa, 266 p. 

_____ 1986. CRC Handbook of Tapeworm Identification. CRC Press, Boca 

Raton, Florida, 675 p. 

Schwartz, F.J. 1977. Effects of sharksucker, Echeneis naucrates, disk on scaled 

and scaleless fishes and sea turtles. American Society of Biologists 

Bulletin 24:84. 

Scott, T. and A. Scott. 1913. The British parasitic Copepoda. Volumes I-II, Ray 

Society, London, England, 256 p., 72 pl. 

Shaffer, R. V. and E. L. Nakamura. 1989. Synopsis of biological data on the 

cobia Rachycentron canadum (Pisces: Rachycentridae). NOAA Technical 

Report NMFS 82 and FAO Fisheries Synopsis 153:21 p. 

Shiino, S.M. 1959. Copepod parasites of eastern Pacific fishes [in German]. 

Report of Faculty of Fisheries, Prefectural University of Mie 3:267-333.

BIBLIOGRAPHY 

_____ 1960. Copepods parasitic on the fishes collected on the coast of 

Province Shima, Japan. Report of Faculty of Fisheries, Prefectural University 

of Mie 3:471-500. 

Siddall, M.E., D.S. Martin, D. Bridge, S.S. Desser, and D. Cone. 1995. The 

demise of a phylum of protists: phylogony of Myxozoa and other parasitic 

Cnidaria. Journal of Parasitology 81:961-967. 

Siddiqi, A.H. and R.M. Cable. 1960. Digenetic trematodes of marine fishes of 

Puerto Rico. Scientific Survey of Porto Rico and the Virgin Islands 

17:257-369. 

Silas, F.G. 1962. Parasites of scombroid fishes. Part I. Monogenetic trematodes, 

digenetic trematodes, and cestodes. Pages 799-875 In: Symposium 

on Scombroid Fishes, Marine Biological Association of India, Part III: 

799-1236. 

_____ and A.N.P. Ummerkutty. 1962. Parasites of scombroid fishes. Part II. 

Copepoda. Pages 876-993 In: _____ Part III:799-1236. 

Simmons, D.C. 1969. Maturity and spawning of skipjack tuna (Katsuwonus 

pelamis) in the Atlantic Ocean, with comments on nematode infestation of 

the ovaries. U.S. Fish and Wildlife Service, Special Scientific Report 

Fisheries 580:1-17. 

Sindermann, C.J. 1990. Principal Diseases of Marine Fish and Shellfish. 2nd 

Edition. Academic Press, San Diego, California, USA. Volume 1, 521 p., 

Volume 2, 516 p. 

_____ 1993. Interactions of pollutants and disease in marine fish and shellfish. 

Chapter 16, Pages 451-482 In: J.A. Couch and J.W. Fournie (Eds.) 

Pathobiology of marine and estuarine organisms. Advances in Fisheries 

Science. CRC Press, Boca Raton, Florida, USA. 

Skinner, R.H. 1978. Some external parasites on Florida fishes. Bulletin of 

Marine Science 28:590-595. 

Smith, J.L.B. 1950. The sea fishes of southern Africa. Central News Agency, 

Capetown, South Africa, 550 p. 

Sniesko, S.F. (Ed.). 1970. A Symposium on Diseases of Fishes and Shellfishes. 

American Fisheries Society, Special Publication No. 5:526 p. 

Sogandares-Bernal, F. 1959. Digenetic trematodes of marine fishes from the 

Gulf of Panama and Bimini, British West Indies. Tulane Studies in 

Zoology 7:70-117. 

_____ and R.F. Hutton. 1959. Studies on helminth parasites from the coast of 

Florida. IV. Digenetic trematodes of marine fishes of Tampa, Boca Ciega 

Bays, and the Gulf of Mexico. Quartly Journal of the Florida Academy of 

Sciences 21:259-273. 

Speare, P. 1990a. Relationships among black marlin, Makaira indica, in eastern 

Australia coastal waters, inferred from parasites. Australian Journal of 

Marine and Freshwater Research 45:535-549. 

_____ 1990b. Parasites of the Pacific sailfish (Istiophorus platypterus): A 

preliminary investigation of their useful in stock discrimination. Pages 95- 

102 In: R. H. Stroud [see Barse and Hocutt 1990]. 

359

360 


Sproston, N.G. 1945. The genus Kuhnia n. g. (Trematoda: Monogenea). An 

examination of the value of some specific characters, including factors of 

relative growth. Parasitology 36:176-190. 

Steele-Llinás, R.M. 1982. Some parasitic copepods of marine fishes of Puerto 

Rico and other adjacent areas of the Caribbean. MS Thesis, University of 

Puerto Rico, Mayaguez, Puerto Rico, 88 p. 

Strasburg, D.W. 1959. Notes on the diet and correlating structures of some 

central Pacific echeneid fishes. Copeia 1959:244-248. 

_____ 1959. An instance of natural mass mortality of larval frigate mackerel in 

the Hawaiian Islands. Journal of Conservation 24:255-263. 

Sumner, F.B., R.C. Osborn and L.J. Cole. 1913. A biological survey of the 

waters of Woods Hole and vicinity. Section 3. A catalogue of the marine 

fauna. U.S. Bureau of Fisheries Bulletin 31:545-794. 

Tetra Tech, Inc. 1992. Characterization of Use Impairments of the U.S. Virgin 

Islands and Puerto Rico. Final Report to Marine and Wetlands Protection 

Branch, USEPA, New York, 428 p. 

Thatcher, V.E. 1959. A report on some monogenetic trematode parasites of 

Louisiana marine fishes. Proceedings of the Louisiana Academy of 

Sciences 22:78-82. 

_____ 1993. Neotropical Trematodes [in Portuguese]. Instituto Nacional de 

Pesquisas da Amazõnia, Manaus, Brazil 553 p. 

Vervoort, W. 1965. Three new species of Bomolochidae (Copepoda, Cyclopoida) 

from tropical Atlantic tunnies. Zoologische Verhandelingen, Leiden 

76:3-40. 

Vigeras, P. 1935. Tristomas poeyi n. sp. (Trematoda) parasitic on Makaira 

ampla Poey (Pisces) [in Spanish]. Memorias de la Sociedad Cubana de 

Historia Natural 9:43-44. 

Wallace, D.H. and E.M. Wallace. 1942. Observations on the feeding habits of 

the white marlin, Tetrapturus albidus Poey. Publications of the 

Chesapeake Biological Laboratory 50:3-10. 

Walters, V. 1980. Parasitic copepoda and monogenea as biological tags for 

certain populations of Atlantic bluefin tuna. ICCAT Collection Volume 

Science Papers IX(2):491-498. 

Ward, D.R. and C. Hackney. 1991. Microbiology of marine food products. 

Van Norstrand Reinhold, New York, USA, 450 p. 

Ward, H.L. 1954. Parasites of marine fishes of the Miami region. Bulletin of 

Marine Science of the Gulf and Caribbean 4:244-261. 

Ward, J.W. 1962. Helminth parasites of some marine animals, with special 

reference to those from the yellow-fin tuna, Thunnus albacares (Bonnaterre). 

Journal of Parasitology 48:155. 

Wardle, R.A. and J.A. McLeod. 1952. The zoology of tapeworms. The 

University of Minnesota Press, Minneapolis, Minnesota, USA, 780 p. 

Watertor, J.L. 1973. Incidence of Hirudinella marina Garcia, 1770 (Trematoda: 

Hirudinellidae) in tunas from the Atlantic Ocean. Journal of Parasitology 

59:207-208.

BIBLIOGRAPHY 

Watson, C., R.E. Bourke and R.W. Brill. 1988. A comprehensive theory on 

the etiology of burnt tuna. Fishery Bulletin 86:367-372. 

Williams, E.H., Jr. 1978. Conchoderma virgatum (Spengler) (Cirripedia, 

Thoracica) in association with Dinemoura latifolia (Steenstrup and Lütken) 

(Copepoda, Caligidea) a parasite of the shortfin mako, Isurus oxyrhynchus 

Rafinesque (Pisces, Chondrichthyes). Crustaceana 34:109-111. 

_____ 1982. Disease factors which should be considered in fish kill investigations. 

Pages 42-43 In: D.K. Atwood (Ed.) Unusual mass fish mortalities in 

the Caribbean and Gulf of Mexico. Atlantic Oceanographic and Meteorological 

Laboratories, Miami, Florida, 46 P. 

_____ 1983. New host records for some nematode parasites of fishes from 

Alabama and adjacent waters. Proceedings of the Helminthological Society 


_____ and T.E. Bowman. 1994. Lironeca Leach, 1818 (Isopoda, Cymothoidae): 

proposed conservation as the correct spelling. Bulletin of Zoological 

Nomenclature 51:224-226. 

_____ and L. Bunkley-Williams. 1994. Four cases of unusual crustacean-fish 

associations and comments on parasitic processes. Journal of Aquatic 

Animal Health 6: 202-208. 

_____ and _____ 1995. Book Review: Keys to the cestode parasites of 

vertebrates, edited by L.F. Khalil, A. Jones, and R.A. Bray. 1994. CAB 

International, Oxon, United Kingdom, 751 p. Fisheries Review 40:574. 

*_____ and _____ 1997a. Fish parasitic isopods. Chapter In: G.L. Hoffman 

(Ed.) Parasites of North American Freshwater Fishes. Cornell University 

Press (in press). 

*_____ and _____ 1997b. Marine major ecological disturbances: Destructive 

indicators of global changes? Chapter In: B. McKay, Greenpeace (Ed.) 

Global Changes. Oxford University Press (in press). 

_____, _____ and T.G. Rand. 1994. Some copepod and isopod parasites of 

Bermuda marine fishes. Journal of Aquatic Animal Health 6:279-280. 

_____, _____ and C.J. Sanner. 1994. Some copepod and isopod parasites of 

Colombian marine fishes. _____ 6:362-364. 

_____, _____, M.J. Dowgiallo and W.G. Dyer. 1991. Influence of collection 

methods on the occurrence of alimentary canal helminth parasites in fish. 


_____ and J.L. Gaines, Jr. 1974. Acanthocephala of fishes from marine and 

brackish waters of the Mobile Bay region. Journal of Marine Sciences of 

Alabama 2:135-148. 

_____, R.R. Lankford, G. van Buurt, D.K. Atwood, J.C. Gonzalez, G.C. McN. 

Harvey, B. Jimenez, H.E. Kumpf and H. Walters. 1982. Summary report 

of the IOCARIBE steering committee for developing region-al 

contingencies for fish kills. UNESCO, Paris, Report Number CARI/ FK/- 

1/3, 20 p. 

*_____ and C.J. Sindermann. 1992. Effects of disease interactions with 

exotic organisms on the health of the marine environment. Pages 71-77 

In: M.R. DeVoe (Ed.) Introductions and Transfers of Marine Species, 

361

362 


Achieving a Balance Between Economic Development and Resource 

protection. South Carolina Sea Grant, Charleston, South Carolina, 204 p. 

_____ and L.B. Williams. 1986. The first association of Conchoderma virgatum 

(Spengler)(Cirripedia: Thoracica) with a euryphorid copepod in the 

mouth of a fish. Galaxea 5:209-211. 

*_____ and _____ 1987. Caribbean Mass Mortalities: A problem with a 

solution. Oceanus 30(4):69-75. 

Williams, H.H. and A. Jones. 1976. Marine helminths and human health. 

Commonwealth Institute of Helminthology Misc. Publication No. 3:47 p. 

Wilson, C.B. 1905. North American parasitic copepods belonging to the family 

Caligidae. Part I. The Caliginae. Proceedings of the U.S. National Museum 

28:479-572. 

_____ 1908. North American parasitic copepods: New genera and species of 

Caliginae. _____ 33:593-627, pl. 49-56. 

_____ 1911. North American parasitic copepods belonging to the Family 

Ergasilidae. _____ 39:263-400. 

_____ 1917a. North American parasitic copepods belonging to the Lernaeopodidae 

with a revision of the entire family. _____ 47:565-729, 29 pl. 

_____ 1917b. North American parasitic copepods belonging to the Lernaeidae 

with a revision of the entire family. _____ 53:1-150, 20 pl. 

_____ 1932. The copepods of the Woods Hole region. Massachusetts. U.S. 

National Museum Bulletin Number 158:1-635. 

_____ 1935a. New parasitic copepods. Smithsonian Miscellaneous Collections 

91(19):1-9. 

_____ 1935b. Parasitic copepods from the Dry Tortugas. Papers from the 

Tortugas Laboratory 29:329-347, 6 pl. 

Yamaguti, S. 1939. Parasitic copepods from fishes of Japan, part 5: Caligoida, 

III. Volumen Jubilare Pro Prof. Sadao Yoshida 3:443-487. 

_____ 1959-1963. Systema Helminthum, Vol. II, 1959. The Cestoda of Vertebrates 

(860 p.). Vol. III, 1961. The Nematodes of Vertebrates (1261 p.). 

Vol. IV, 1963. Monogenea and Aspidocotylea (699 p.). Vol. V, 1963. 

Acanthocephala (421 p.). 1963c. Parasitic Copepoda and Branchiura of 

Fishes (1104 p.). Interscience Publishers, New York, USA. 

_____ 1968. Monogenetic trematodes of Hawaiian fishes. University of Hawaii 

Press, Honolulu, Hawaii, USA, 288 p. 

_____ 1970. Digenetic trematodes of Hawaiian fishes. Keigaku Publishers, 

Tokyo, Japan, 436 p. 

_____ 1971. Synopsis of digenetic trematodes of vertebrates. Vols. 1-2, 

Keigaku Publishing Company, Tokyo, Japan, 1074 p. 

_____ 1975. A synoptical Review of Life Histories of Digenetic Trematodes of 

Vertebrates: With Special Reference to the Morphology of Their Larval 

Forms. Satyu Yamaguti, Kyoto, Japan, 590 p. 

Young, R.T. 1954. A note on the life cycle of Lacistorhynchus tenuis (van 

Beneden, 1858), cestode of the leopard shark. Proceedings of the 

Helminthological Society of Washington 21:111.

ABOUT THE AUTHORS 

Cutting-edge science thrives on mysteries, and Dr. Ernest H. (Bert) 

Williams, Jr. finds an ample supply at the crossroads between Caribbean 

Aquatic Animal Health and Worldwide Disturbances. Bert received his doctorate 

in Fisheries Management from Auburn University in 1974 and has been 

a Professor in the Department of Marine Sciences UPR ever since. His interests 

include not only fish parasites but large-scale marine ecological disturbances. 

Bert was an Officer in the Alabama and Puerto Rico Army National 

Guards. He is a Member of the Board of the Global Coral Reef Alliance and the 

Caribbean Stranding Network, a Member of the U.S. National Research Council 

Non-Indigenous Marine Species Panel, Headed a UNESCO/United Nations 

Committee on Caribbean Fish Kills and is a Cofounder and Country Representative 

of the Caribbean Red Tide and Mass Mortality Network. He was Coordinator 

of the Marine Ecological Disturbance Information Center (unfunded at 

present), and Executive Director of the Association of Marine Laboratories of 

the Caribbean for 15 years (1977-91). Bert is an Associate Editor of the Journal 

of Aquatic Animal Health, and is on the Review Board of the Caribbean Journal 

of Science. A fluke, copepod and isopod have been named in Bert's honor; and 

he received an honorable mention in the 1993 Rolex Awards for Enterprise for 

his work with marine ecological disturbances. 

Dr. Lucy Bunkley-Williams completed her Doctorate in Fish Pathology 

at Auburn University in a near record pace of 2 years by 1984. She supported 

herself as a Research Associate in DMS from 1987 until 1995 when she became 

an Assistant Professor in the Biology Department, and continues to support her 

research at UPR with external research grants. In 1991, she completed a 

Shrimp Disease Course at the University of Arizona with Sea Grant support to 

make this expertise available in Puerto Rico. 

Lucy is an award-winning underwater and scientific photographer. Her 

photographs are on display in the Smithsonian and have appeared around the 

world in books and magazines, and her underwater video footage has been used 

in national and international TV programs. Lucy has been honored as one of 

the Outstanding Women Graduates of Auburn University, by being featured in 

a U.S. News and World Report article, and by having 6 new species named for 

her (protozoan, fluke, spiny-headed worm, copepods, isopod). 

Recent emergencies for Lucy and Bert included investigating massive kills 

of coral reef fishes in Barbados, mortalities of sea urchins throughout Puerto 

Rico and south Florida, and dying tilapia and shrimp in aquaculture facilities; 

diagnosing a scoliosis condition in local largemouth bass, Micropterus salmoides 

(Lacepéde), which may infect humans; isolating and naming a new primary 

pathogen which causes Tilapia-Wasting Disease, describing the new disease, and 

tracing its origin and international distribution in hopes of stopping its spread; 

and responding to a plan submitted to the government, to import a new exotic 

fish, with recommendations to protect Puerto Rico from its diseases. Despite 

381

382 


this typically frantic but productive schedule, they have found time to characterize 

the parasites and diseases of local freshwater fishes in books in English 

and Spanish to make this information available in a friendly and useable format 

for everyone. 

Lucy and Bert also solve aquatic animal health problems for sport fishermen, 

the Department of Natural and Environmental Resources (DNER), the 

University of Puerto Rico (UPR) and the local public; and represent Puerto Rico 

in international fish kill investigations as part of the Sportfish Restoration 

Project DNER/UPR. They have solved disease problems for aquaculture 

projects under separate grants, and for the Caribbean Stranding Network. They 

have "Doctored" all sorts of creatures from microscopic fairy shrimp to whales. 

Lucy has known the frustration of having a beautiful Atlantic spotted dolphin, 

Stenella frontalis (Cuvier), she was trying to save, die in her arms; and the joy 

of rehabilitating and releasing marine mammals and sea turtles. 

Lucy and Bert have published more than 180 scientific papers and they 

have also written popular articles explaining coral reef bleaching, mass 

mortalities and fish isopods. They are preparing a chapter about marine 

ecological disturbances for the Greenpeace volume on "Global Change". Their 

work has been featured in a marine biology textbook, a children's science book, 

in numerous national and international newspaper and magazine articles and 

TV programs, and they were featured in a national "Network Earth" television 

program filmed in La Parguera, Puerto Rico. Future endeavors include a 

textbook "Marine Major Ecological Disturbances" (MMED), and a guidebook 

to isopod parasites of world fishes. 

Bert and Lucy have "buddied" on more than 2500 hours of underwater 

research scuba dives throughout the West Indies and parts of the Pacific. Bert 

has known the terror of going "nose-to-nose" with a 2000 pound great white 

shark in open mid-water. Lucy is preparing a popular book describing their 

experiences living and working underwater, as Co-principal Investigators and 

Aquanauts, on 5 week-long Hydrolab Undersea Habitat, NOAA research 

missions (1979-85), in St. Croix, U.S. Virgin Islands. 

Bert and Lucy were Visiting Foreign Researchers at the Sesoko Marine 

Science Center, University of the Ryukyus, in 1985-86, where they conducted 

underwater studies around Okinawa and many adjacent islands, that resulted in 

20 scientific publications about fish parasites, fish associates, fishes and birds. 

They also visited almost every fisheries and marine sciences lab. in Japan to 

examine isopod parasites of fishes.

parasites of offshore big game fishes of puerto rico - Uprm

Create successful ePaper yourself

Delete template?

Save as template?