6 - Sphaeromatidae::“Cute As Buttons”

- « 

.4 i r - « - - f -- - 

' ^HSTSaP 

4 

! * • ; * ! 

-9 * 

.• 

• 

I 

M**

Guide to the 

MARINE ISOPOD 

CRUSTACEANS 

of the Caribbean

Guide to the 

MARINE ISOPOD 

CRUSTACEANS 

of the Caribbean 

Brian Kensley and 

Marilyn Schotte 

SMITHSONIAN INSTITUTION PRESS 

WASHINGTON, D.C., AND LONDON

© 1989 by the Smithsonian Institution 

All rights reserved 

Designer: Linda McKnight 

Editor: Nancy Dutro 

Library of Congress Cataloging-in-Publication Data 

Kensley, Brian Frederick. 

Guide to the marine isopod crustaceans of the 

Caribbean / Brian Kensley 

and Marilyn Schotte. 

p. cm. 

Bibliography: p. 

Includes index. 

ISBN 0-87474-724-4 (alk. paper) 

1. Isopoda—Caribbean Sea— 

Classification. 2. Crustacea—Caribbean Sea— 

Classification. I. Schotte, Marilyn. II. Title. 

QL444.M34K434 1989 

595.3'7209153'35—dcl9 88-38647 

British Library Cataloging-in-Publication Data 

available 

Manufactured in the United States of America 

10 9 8 7 6 5 4 3 2 1 

98 97 96 95 94 93 92 91 90 89 

°° The paper used in this publication meets the 

CIP 

minimum requirements of the American National 

Standard for Performance of Paper for Printed 

Library Materials Z39.48-1984

Contents 

1 Introduction 

1 HISTORIC BACKGROUND 

3 GEOGRAPHIC AREA COVERED IN THIS GUIDE 

4 ARRANGEMENT OF THE GUIDE AND HOW TO USE IT 

5 ACKNOWLEDGMENTS 

7 Glossary of Technical Terms 

13 Marine Isopods of the Caribbean 

13 ORDER ISOPODA 

15 SUBORDER ANTHURIDEA 

73 SUBORDER ASELLOTA 

107 SUBORDER EPICARIDEA 

114 SUBORDER FLABELLIFERA 

236 SUBORDER GNATHIIDEA 

243 SUBORDER MICROCERBERIDEA 

246 SUBORDER ONISCIDEA 

251 SUBORDER VALVIFERA 

261 Zoogeography 

261 FAUNAL PROVINCES 

262 ANALYSIS OF THE ISOPOD FAUNA 

266 THE BAHAMAS 

269 BERMUDA 

269 CAVE ISOPODS 

275 Appendix 

277 Literature Cited 

293 Index

Introduction 

The title of this work will no doubt raise several questions in many readers' 

minds: why the Caribbean? why not the Caribbean and the Gulf of Mexico? 

why only the marine isopods? just what is the "Caribbean area"? We hope 

that the answers to some of these (and other) questions will become 

apparent. 

There are several works that already deal with the isopods of the Carib 

bean, as part of a wider treatment of North American isopods (e.g., 

Richardson, 1905; Schultz, 1969). Why then this "Isopods of the Carib 

bean"? As partial answer, the following: many new records of isopods from 

the Caribbean region (in its broadest sense) have appeared in scattered pub 

lications in the last few decades. The time has come to pull these together in a 

single work. The number of marine laboratories in the area has increased, 

with more and more students exploring especially the shallow marine en 

vironment. A single work on a relatively speciose and abundant group of 

invertebrates would be useful to such investigators, as they build up a com 

prehensive view of the biology of the region. Concepts of the taxonomy of 

several isopod groups have changed radically over the last few years; again, 

there is obvious utility in having these changes summarized in a single 

source. New species and records are continuously being found. Having a 

single baseline work decreases the time needed for investigating and estab 

lishing the validity of such records. 

HISTORIC BACKGROUND 

The history of isopod taxonomic research in the Caribbean really starts with 

a worldwide monographic work on the Cirolanidae by Hansen in 1890. In 

cluded here were about 12 species from the Danish West Indies, now the 

U.S. Virgin Islands. Since then a few major works on Caribbean isopods 

have appeared, such as Moore's report on the isopods of Puerto Rico (1901) 

and Menzies and Glynn's report on the same area (1968). Some areas have 

received considerable attention, such as the aforementioned Puerto Rico and, 

more recently, Belize. A list of 116 species of isopods from Cuba (including 

Oniscidea) has been published (Ortiz, Lalana, and Gomez, 1987). At the 

other extreme, there are no records from a number of localities, especially the 

1

Figure 1. Map of area covered by this guide.

INTRODUCTION 3 

islands of the southeastern chain of the Lesser Antilles. In total there are 

about 40 publications, varying from descriptions of single species to longer 

works, that deal with isopods from the Caribbean. These publications will be 

encountered in the following guide, under the specific taxa. 

GEOGRAPHIC AREA COVERED IN THIS GUIDE 

The accompanying map (Figure 1) shows the area for which records are 

included in this guide. 

While it may seem logical to include the Gulf of Mexico, and while there 

are several isopod species common to both areas, this has not been done. 

There are relatively few isopod records from the Gulf; undoubtedly a great 

deal of taxonomic work awaits the careful collector in this area. Also, from a 

zoogeographic point of view, separation of the Gulf may be justified. 

Bermuda, on the other hand, situated in the northwestern Atlantic several 

hundred miles off the coast of the United States, is included. This island, 

although remote from the Caribbean, is swept by waters that earlier have 

passed through the Caribbean. Zoogeographically, the shallow-water Ber- 

mudan and Caribbean faunas have much in common. 

While perhaps not strictly in the Caribbean Sea, the Bahamas and the 

Florida Keys are included here, their shallow-water marine faunas being 

overwhelmingly Caribbean in nature. 

Turning to depths limits, within the area under discussion, species from 

the intertidal to 200 meters have been dealt with in some detail. This arbi 

trary cutoff depth was selected because most Caribbean isopod species in 

habit relatively shallow depths. About 30 species have been recorded from 

below 200 meters in the Caribbean, many of these known only from the type 

material. A list of species of this very poorly known deeper fauna is included 

here. Without doubt, many species in the deeper waters of the Caribbean 

await discovery. 

A fascinating group of isopods, while not strictly shallow-water marine 

forms, is included. These are the true cave forms, found mainly in the sub 

orders Anthuridea and Flabellifera. Given the history of the Caribbean from 

the Quaternary to the present, it is not surprising that caves are common 

throughout the region. These caves may be well inland and contain only 

freshwater, but are more commonly anchialine, that is, having some (fre 

quently subterranean) link to the sea. Less common, and of lesser interest 

from an isopod taxonomist's point of view, are the fully marine caves in 

direct communication with the sea or, indeed, under the surface of the sea 

itself.

4 INTRODUCTION 

ARRANGEMENT OF THE GUIDE AND HOW TO USE IT 

A short introduction to the Crustacea Isopoda is provided, followed by a 

glossary of descriptive terms and morphological features used throughout the 

guide (see Figure 2). 

Keys and diagnoses to the suborders and all lower taxa follow. For ease of 

usage, except in the keys, all taxa are presented in alphabetical order, regard 

less of their phylogenetic relationships. 

Diagnoses are provided for all suborders, families, genera, and species. 

The only exceptions to this are in the suborders Epicaridea and Oniscidea. 

Within each suborder, a key to the families occurring in the Caribbean is 

provided. Similarly, within each family and genus, keys are provided to the 

relevant genera and species, respectively. 

In whatever context, where an author and date appear, a reference to these 

is provided in the Literature Cited section. In some cases, reference is made 

to useful publications such as revisions of families or genera. 

As this is not a textbook on the Isopoda, biological information is generally 

kept to a minimum. In the case of individual species, however, what little 

ecological information is available, is provided. For general texts on biology, 

internal anatomy, physiology, and reproduction, the reader is referred to 

works such as Kaestner (1967), Waterman (1960), Bliss (1982-1985), and 

Schram (1986). 

Within each species discussion, a diagnosis is given, along with maximum 

(total) middorsal lengths for males and females, where known. The diag 

noses are not exhaustive, but provide only the information needed to distin 

guish the species. Diagnoses thus vary in length from the statement of a 

single feature to a paragraph concerning several features, depending on the 

understanding and complexity of the taxonomy of the group. In the longer 

diagnoses, morphological features are dealt with in order from anterior to 

posterior on the animal's body. Records are given, rather than geographical 

distribution, as our knowledge of many species is woefully incomplete. These 

records are given in a roughly north-to-south order; records outside of the 

Caribbean region, as here defined, are given on a separate line. A few species 

not yet recorded from the Caribbean are included, in the strong likelihood 

that they will eventually be found here. The records include depth distribu 

tion information in meters, where known. Records were taken from pub 

lished papers; in addition, the collections of the United States National 

Museum of Natural History, Smithsonian Institution, were scoured, and 

many unpublished records from this source are also included. In the "Re 

marks" section, ecological information such as substrate preferences is given. 

Hosts of parasitic species are given. Formal synonymies are not provided,

INTRODUCTION 5 

but nomenclatural comments are included in the few cases where a species 

may be known under a more commonly used name. Usually, a figure of the 

entire animal of each species is given. Diagnostic features are usually illus 

trated. Unless otherwise stated, all illustrations are original and by the au 

thors, and were made from actual specimens. 

Common and scientific names of fishes that are hosts to parasitic isopods 

are taken from the American Fisheries Society special publication no. 12 

(Robins et al., 1980). 

Finally, a word of warning. Difficulties may be experienced in using the 

keys, for which there may be any of several reasons: characters seen in the 

animal may not clearly conform to those in the key (in which case refer to the 

figures, as well as to good recent descriptions or diagnoses); your material 

may be a new record for the region; or you may have an undescribed species 

(in which case refer to more comprehensive treatments of the group). 

ACKNOWLEDGMENTS 

Much of the material covered in this work comes from the many collectors 

who have deposited specimens from several regions of the Caribbean in the 

collections of the National Museum of Natural History, Smithsonian Institu 

tion. Rather than risk the unwitting omission of a name, we thank all of these 

individuals collectively. Without their efforts, our knowledge of the Carib 

bean fauna would be the poorer. 

Material was borrowed from several institutions. We thank the following 

scientists for their assistance in this connection: Jan Stock and Dirk Platvoet 

of the Instituut voor Taxonomische Zoologie, University of Amsterdam; 

Jean Just and Torben Wolff of the Zoological Museum, University of 

Copenhagen; Jacques Forest of the Museum National d'Histoire Naturelle, 

Paris; Richard Heard of the Gulf Coast Research Laboratory, Ocean 

Springs, Mississippi; Willard Hartman of the Peabody Museum of Natural 

History, Yale University; Herbert W. Levi of the Museum of Comparative 

Zoology, Harvard University; Harold S. Feinberg of the American Museum 

of Natural History, New York; John E. Miller of the Harbor Branch Founda 

tion, Florida; and Paula M. Mikkelsen of the Indian River Coastal Museum 

at Fort Pierce, Florida. 

Bruce Collette of the National Marine Fisheries Laboratory at the 

Smithsonian Institution assisted with fish names used in this work, for which 

we are grateful. 

We thank the staff of the Scanning Electron Microscope Laboratory of the 

Smithsonian Institution, and especially Susann Braden, who produced the 

electron micrographs used here.

6 INTRODUCTION 

We are grateful to the Smithsonian Research Opportunities fund, admin 

istered by David Challinor, then Assistant Secretary for Research, and to the 

Smithsonian's Caribbean Coral Reef Ecosystems program administered by 

Klaus Riitzler of the Department of Invertebrate Zoology, for the funding of 

several fieldtrips to the Caribbean. The second author acknowledges a finan 

cial award from the Smithsonian's Women's Committee for a grant to facili 

tate fieldwork. 

Several individuals have provided encouragement, advice, suggestions, 

critical comments, and missed references, all of which have vastly improved 

this work. In this regard we are especially grateful to Thomas E. Bowman, C. 

W. Hart, Jr., Horton H. Hobbs, Jr., and Molly K. Ryan, all of the Depart 

ment of Invertebrate Zoology, National Museum of Natural History, 

Smithsonian Institution; Dan Adkison, Bass Harbor, Maine (who provided 

considerable assistance with the Epicaridea); Richard Heard of the Gulf 

Coast Research Laboratory, Mississippi; and Paul Delaney of the Los An 

geles County Museum of Natural History. 

This is Contribution Number 248 of the Caribbean Coral Reef Ecosystems 

(CCRE) program, Smithsonian Institution, supported in part by the Exxon 

Corporation.

Glossary of Technical Terms 

AESTHETASC. Thin-walled sensory seta usually found on flagellum of 

antennule 

AMBULATORY (as applied to pereopods). Used for walking. 

ANCHIALINE. An aqueous habitat near the sea; referring to saltwater or brackish 

pools fluctuating with the tides, but with no surface connection to the sea. 

ANGULATE. Having an angle or an angular shape. 

ANTENNA. Paired appendage of the third cephalon segment; sometimes referred 

to as antenna 2. 

ANTENNULE. Paired appendage of the second cephalon segment; sometimes 

referred to as antenna 1. 

APICAL. Relating to the apex or tip. 

APPENDAGE. An articulated structure used for feeding, locomotion, sensory 

reception, e.g., mouthparts, antennae, pereopods, pleopods, uropods. 

ARTICLE. A single section of an appendage, with an articulation at one or both 

ends. 

BASIS. Article of appendage adjoining coxa proximally, and carrying endopod 

distally, i.e., article 2 of pereopod. 

BIARTICULATE. Composed of two articles. 

BIDENTATE. Having two teeth. 

BIFID. Divided into two lobes or parts by a cleft. 

BILOBED. Composed of two lobes. 

BIRAMOUS. Composed of two rami or branches. 

BIUNGUICULATE. Having two claws, as in a bifid dactylus. 

CARINA. A keel, or an acute ridge. 

CARINATE. Having one or more carinae or acute ridges. 

CARPOCHELATE. Having a chela or pincerlike structure formed by the seventh 

(dactylus) and fifth (carpus) articles of an appendage. 

CARPUS. Article 5 of pereopod. 

CEPHALON. Anterior region of body or head; more correctly the cephalothorax 

in isopods, as the first pereonal segment is usually fused with the head. 

CHELA. Distal pincerlike part of appendage, often formed by a mobile and an 

immobile finger. 

CHELATE. Having a chela; modified to form a pincer. 

CLAVATE. Club shaped; having one end thickened. 

CLYPEUS. Platelike structure of cephalon, anterior to upper lip or labrum, 

sometimes fused with frontal lamina. 

CONGLOBATE. Able to roll up into a ball, as in some sphaeromatid and 

oniscidean isopods. 

CONSPECIFIC. Belonging to the same species. 

CONTIGUOUS. Touching.

frontal lamina 

clypeus 

labrum 

peduncle 

uropod 

flageHum 

I 

rrjn 

antennule 

14 

> * antenna 

maxi 

maxilla 2 

pereopods 

ischium 

Vy —^ carpus 

merus 

pleopods 

propodus 

maxilliped 

dactylus 

F i g u r e 2 . S c h e m a t i c r e p r e s e n t a t i o n o f a n i s o p o d i l l u s t r a t i n g m o r p h o l o g i c a l t e r m s .

GLOSSARY 9 

COPULATORY STYLET. Structure situated on endopod of pleopod 2 in males, 

used for transfer of spermatophore in some species; also referred to as appendix 

masculina. 

CORDATE. Heart shaped in outline. 

COXA. Basal article of an appendage, attached to sternite, sometimes expanded 

into a lateral coxal plate. 

CRENULATE. Having a scalloped edge with rounded teeth, usually used to refer 

to the margin of a structure. 

DACTYLUS. Terminal (7th) article of a pereopod or thoracic appendage. 

DENTATE. Edged with teeth. 

DENTICLE. A small tooth. 

DENTICULATE. Having fine teeth. 

DIGITIFORM. Fingerlike. 

DISTAL. Situated away from the base or point of origin or attachment. 

ECDYSIS. Molting of the integument. 

EMARGINATE. Having the margin concave. 

ENDITE. Medially directed lobe of coxa or basis of an appendage, especially the 

maxilliped. 

ENDOPOD. Inner ramus of a biramous appendage. 

ENTIRE. Complete; usually referring to the margin of a structure that is smooth. 

EPIMERON. Lateral part of a somite. 

EPIPOD. Lateral extension of a protopodite. 

EXCAVATE. Hollowed out. 

EXOPOD. Outer ramus of a paired appendage. 

FALCATE. Sickle shaped; curved and tapering to a point. 

FLAGELLUM. Distal part of antenna or antennule, usually multiarticulate, 

occasionally reduced to one or a few articles. 

FRONTAL LAMINA. Platelike structure of the cephalon immediately anterior to, 

and sometimes fused with, clypeus. 

GENICULATE. Bent at an abrupt angle, as in the body of many arcturid isopods. 

GRANULATE. Having the appearance of bearing beadlike or grainlike 

protuberances; usually applied to the description of a surface. 

HIRSUTE. Bearing hairs (elongate hairs in the case of most isopods). 

HYPOGEAN. Underground. 

IMMERSED. Sunken into, as with one structure into another. 

INCISOR. Cutting process of the mandible, usually dentate, sometimes modified 

for piercing. 

INDURATE. Hardened, usually by calcium carbonate or sclerotized protein. 

INTEGUMENT. Outer covering, e.g., the exoskeleton. 

INTERSTITIAL. Relating to interstices; living in the interstices of sand grains, 

gravel, or rubble. 

ISCHIUM. Article 3 of pereopod. 

LABIUM. Lower lip; usually consisting of a pair of lobes posterior to the mouth. 

LABRUM. Unpaired projection anterior to mouth, attached to the clypeus; upper 

lip. 

LACINIA MOBILIS. Small, usually toothed, process articulating at base of incisor 

in left or both mandibles. 

LAMELLAR. In the shape or structure of a thin plate or lamella.

10 GLOSSARY 

LAMINA DENTATA. Serrate platelike structure in the mandible of anthurideans, 

formed by the fusion of spines of the spine-row. 

LANCEOLATE. Lance shaped; narrow and tapering to a point. 

LINGUIFORM. Tongue shaped. 

MANGA. Young of some peracaridean crustaceans (including isopods), lacking 

last thoracic appendage at time of release from broodpouch. 

MANDIBLE. First pair of mouthparts, functioning as jaws, often sclerotized. 

MARSUPIUM. Structure in which eggs are retained by female; the broodpouch. 

MAXILLA (1 and 2). Two sets of paired mouthpart appendages immediately 

posterior to mandible. 

MAXILLIPED. First paired appendage of the thorax; usually incorporated into the 

mouthparts. 

MEDIAN. At, near, or directed toward the middle or midline. 

MERUS. Article 4 of pereopod. 

METAMORPHOSED. Transformed; changed in appearance, structure, or function. 

MESIAL. Near or toward the middle or midline. 

MOLAR. Grinding, and sometimes piercing, structure of the mandible. 

MULTIARTIGULATE. Composed of many articles. 

NATATORY. Adapted for swimming. 

OBSOLETE. Becoming vestigial, and losing original function. 

OMMATIDIA. Individual visual components of the compound eye. 

OOSTEGITE. Medially directed lamellar structure arising from coxa of pereopod 

in the female, forming part of the broodpouch or marsupium. 

OPERCULIFORM. In the form of a cover or lid. 

OVATE. Egg shaped or oval. 

PALM. Cutting edge of the propodus, often defined proximally by a spine, in a 

subchelate appendage. 

PALP. Articulated ramus consisting of one to three articles in mandible, of up to 

five articles in the maxilliped. 

PECTINATE. Having teeth like a comb. 

PEDUNCLE. Stalk or proximal part of an appendage, as in antennae. 

PENIAL RAMI. Paired submedian process on sternite 7 of male. 

PEREON. Middle or thoracic region of the body, consisting of seven segments or 

pereonites, first fused with cephalon in isopods. 

PEREONITE. Segment of the pereon. 

PEREOPOD. Paired appendage of the pereon, consisting of seven articles when 

unmodified. 

PILOSE. Covered with short hairs or setae. 

PLEON. Posterior or abdominal region of the body, primitively consisting of six 

segments or pleonites, and bearing paired pleopod and uropod appendages. 

PLEONITE. Segment of the pleon. 

PLEOPOD. Paired appendage of the pleon, five pairs being present in the 

primitive condition. 

PLEOTELSON. Structure resulting from the fusion of the telson and one or more 

pleonal segments. 

PLICATE. Pleated or folded. 

PRANIZA. Juvenile, immature stage of gnathiideans. 

PREHENSILE. Adapted for holding or clinging.

PRODUCED. Extended or lengthened. 

PROPODUS. Article 6 of pereopod. 

GLOSSARY 11 

PROTANDROUS. In hermaphroditic forms, becoming a functional male producing 

spermatozoa before becoming a functional female producing eggs. 

PROTOGYNOUS. In hermaphroditic forms, becoming a functional female 

producing eggs before becoming a functional male producing spermatozoa. 

PROTOPODITE. Proximal part of an appendage, consisting of the coxa and basis. 

PROXIMAL. Situated near the point of attachment. 

PYLOPOD. First pereopod of the Gnathiidea, modified to form part of the 

mouthparts. 

RAMUS. Branch of an appendage. 

RENIFORM. Kidney shaped. 

RETICULATE. Resembling or forming a network. 

RETINACULAE. Small hooks on an appendage, used to link the left and right 

members of a pair of appendages. 

ROSTRUM. Anterior middorsal projection of cephalon. 

SAGITTATE. Arrow shaped. 

SGLEROTIZED. Hardened, usually with chitin. 

SERRATE. Edged with toothlike projections as in a saw. 

SETIFEROUS. Bearing setae. 

SETOSE. Bearing setae. 

SINUATE. Having a wavy margin. 

SINUOUS. Having curves. 

SOMITE. Body segment, usually having a pair of appendages. 

SPATULATE. Shaped like a spatula. 

SPICATE. Shaped like a spike. 

SPINE-ROW. Row of spines situated between the incisor and molar processes of 

the mandible. 

SPINOSE. Bearing spines. 

STATOCYST. Small saclike sensory organ, often containing granules, used to 

indicate to the animal its orientation. 

STYGOBIONT. Cave organism. 

STYLIFORM. Having a long, slender, stilettolike shape. 

SUB-. A prefix indicating ''almost" or "just less than," e.g., submarginal—almost 

on the margin. 

SUBGHELATE. Having a subchela, forming a pincerlike structure, especially by 

the dactylus folding back on the propodus. 

SUTURE. A line indicating an area of articulation, or of incomplete fusion. 

SYMPOD. Proximal part of an appendage, often formed by the fusion of the coxa 

and basis. 

TELSON. Terminal part of the body, usually bearing the anus. 

THORAX. Tagma or body region between the cephalon and the abdomen. 

TRAGHEATE. Bearing tubular respiratory trachea (more correctly pseudotrachea) 

on pleopods, as in Oniscidea. 

TRICUSPID. Bearing three cusps or points. 

TRIDENTATE. Having three teeth. 

TRIFID. Divided into three parts or lobes. 

TRILOBED. Divided into three lobes.

12 GLOSSARY 

TRISINUATE. Having three curves. 

TRIUNGUIGULATE. Bearing three claws, as in a trifid dactylus. 

TRUNCATE. Having the appearance of having been abruptly cut off. 

TUBERCULATE. Bearing knoblike or wartlike prominences or tubercles. 

UNIARTIGULATE. Composed of one article. 

UNIRAMOUS. Having one ramus or branch. 

UNIUNGUICULATE. Having a single claw, as in a dactylus. 

UROPOD. Paired pleonal appendage of the last pleonite, usually situated at the 

base of the telson.

Marine Isopods of the Caribbean 

Phylum Arthropoda 

Superclass Crustacea Pennant, 1777 

Class Malacostraca Latreille, 1806 

Subclass Eumalacostraca Grobben, 1892 

Superorder Peracarida Caiman, 1904 

Order Isopoda Latreille, 1817 

DIAGNOSIS Body usually dorsoventrally depressed, occasionally sub- 

cylindrical, rarely bilaterally compressed. Carapace lacking. Antennules and 

antennae uniramous (scale on antenna in some asellotes may represent rudi 

mentary second ramus). Eyes sessile (although situated on nonmobile stalks 

in some asellotes). Mouthparts consisting of one pair of mandibles, two pairs 

of maxillae, one pair of maxillipeds; latter appendages of first thoracic seg 

ment fused with cephalon. Mandible usually with palp consisting of one to 

three articles; incisor, lacinia mobilis, and molar usually present; lacinia mo- 

bilis often differing on left and right sides, sometimes absent from right man 

dible; molar variable. Maxilliped usually consisting of palp of no more than 

five articles, lamellar endite often with coupling hooks, lamellar epipod. Per- 

eonites usually separate, although pereonite 1 sometimes fused with 

cephalon. Coxae of pereopods variously fused with, and forming expanded 

lateral processes of, pereonites. Pereopod 1 forming additional mouthpart 

(pylopod) only in Gnathiidea. Pereopods generally similar, ambulatory; 

pereopods 1-3 secondarily variously modified and becoming subchelate or 

prehensile; pereopods 4-7 occasionally modified, becoming natatory or pre 

hensile. Pereopod 7 occasionally not developed (neotenous condition). 

Broodpouch or marsupium formed by varying number of oostegites attached 

ventrally and medially to coxae of pereopods; eggs held in anterior or pos 

terior pockets or internal pouches in gnathiids and some sphaeromatids. 

Pleon consisting of six pleonites, free or variously fused, plus telson; if one or 

more pleonites fused with telson, resulting structure referred to as pleotelson. 

13

14 1SOPODA 

Key to suborders of Isopoda 

1. Parasitic on crustaceans; body of 9 nearly always asymmetrical 

Epicaridea 

Free-living or parasitic on fishes; body of 9 bilaterally symmetrical, or 

if parasitic, 9 somewhat distorted 2 

2. Body more or less bilaterally compressed Phreatocoidea 

Body more or less dorsoventrally depressed or subcylindrical 3 

3. With six pereonites and five pairs of pereopods Gnathiidea 

With seven pereonites and six or seven pairs of pereopods 4 

4. Body usually more than six times longer than wide, subcylindrical, 

uropods never operculiform 5 

Body usually less than six times longer than wide, usually 

dorsoventrally depressed; if subcylindrical, uropods operculiform . . 6 

5. Uropodal exopod often folding dorsally over pleotelson; rarely 

interstitial forms : Anthuridea ÎC 

Uropods terminal, exopod lacking; minute interstitial forms 

6. Antennules minute; terrestrial forms, with pleopods tracheate 

Microcerberidea 

Oniscidea 

Antennules rarely minute; aquatic forms, pleopods never tracheate . . 7 

7. Uropods ventral, operculiform, covering pleopods Valvifera r' 

Uropods never operculiform over pleopods 8 

8. Uropods lateral or ventrolateral, forming tailfan with pleotelson; 

pleopods 1 and 2 rarely operculiform Flabellifera(? I v 

Uropods terminal or subterminal; pleopods 1 and 2 variously 

operculiform Asellota ,•;••- 

The suborder Phreatocoidea contains freshwater forms, and has a 

Gondwanian distribution, primarily in the Southern Hemisphere. 

Pleopods (on pleonites 1—5) biramous, lamellar, primarily for respiration; 

anterior pleopods occasionally operculiform. Pleopod 2 in male (and occa-

ANTHURIDEA 15 

sionally also pleopod 1 in Oniscidea and Ascllota) with endopod bearing 

copulatory stylet. One pair of uropods on pleonite 6. Young leave brood- 

pouch as manca, i.e., resembling adult but lacking pereopod 7; in Epi- 

caridea, manca stage represented by epicaridium stage; latter transforms 

into microniscium and then cryptoniscium stage, before becoming adult. 

Suborder Anthuridea Leach, 1814 

DIAGNOSIS Body generally elongate and subcylindrical. Eyes absent in 

some genera. Antennular peduncle of three articles; antennal peduncle of five 

articles. Mandible with palp of one to three articles, or absent; body of mand 

ible either styliform and lacking molar and lacinia mobilis, or with molar 

variously specialized or reduced, lacinia mobilis absent, and spine-row modi 

fied to form platelike lamina dentata. Maxilla 1 with inner ramus reduced, 

outer ramus slender. Maxilla 2 rudimentary. Maxilliped variable, with palp 

of one to five articles, endite present, modified, reduced, or absent. Pereonite 

1 free. Pereopod 1, or pereopods 1-3 subchelate; pereopods 4-7 generally 

ambulatory. Pleonites 1-5 free or fused, pleonite 6 partly or completely fused 

with telson. Pleopods 1-5 similar, or pleopod 1 variously modified to form 

operculum. Uropodal exopod often folded dorsally over pleotelson. 

Pleotelson with pair of statocysts, with single statocyst, or lacking statocysts. 

REMARKS Protogyny has been demonstrated in several species of An 

thuridea. The order of development in these cases is: egg, manca (both in the 

broodpouch), immature subadult, ovigerous female, premale, male, with 

varying numbers of molts between each stage. At least one molt takes place 

between ovigerous female and premale, the latter being distinguished by the 

loss of the oostegites and by the elongation of, and acquisition of more flagel 

lar articles in, the antennule. One or two molts take place between premale 

and sexually mature male, the latter being characterized by the possession of 

elongate antennular flagella bearing dense whorls of aesthetascs, a more set 

ose and/or spinose pereopod 1, and sometimes by an elongation of the pleon 

and uropods. In some genera, the males have somewhat atrophied 

mouthparts, suggesting that they do not feed at this stage. As a result of this 

seemingly widespread protogyny, sex ratios are strongly biased toward 

females, and in several species males are not yet known. 

The number of families in the suborder Anthuridea has not been settled. 

At present, three families are recognized. Doubtless, further families will be 

defined and the genera reshuffled.

16 ANTHURIDEA • ANTHUR1DAE 

Key to families of Anthuridea 

1. Mouthparts adapted for piercing and sucking, together forming 

conelike structure Paranthuridae p£i 

Mouthparts adapted for cutting, lamina dentata and molar usually 

present 2 

2. Pereopod 1 subchelate, with propodus expanded; pleonites generally 

fused; if free, much shorter than wide Anthuridae A 

Pereopods 1—3 subchelate, subsimilar; pleonites free, often as long as 

wide Hyssuridae pC& 

Family Anthuridae Leach, 1814 

DIAGNOSIS Mouthparts adapted for cutting. Pereopod 1 usually markedly 

different from remaining pereopods, subchelate with propodus more or less 

inflated. Exopod of pleopod 1 operculiform, covering remaining pleopods. 

Pleonites 1—5 fused, with fusion marked ventrolateral^ by short slits, occa 

sionally with dorsal grooves marking lines of fusion, or free; if free, length of 

each pleonite much less than width. Pleotelson with pair of statocysts, or 

single medial statocyst, or lacking statocysts. 

Key to genera of Anthuridae 

1. Pleopod 1, both rami contributing to operculum 2 

Pleopod 1, only exopod operculiform 6 

2. Antennal peduncle bearing serrate process Licranthura M> 

Antennal peduncle lacking any serrate process 3 

3. Pleopod 1 in 9, rami to some degree fused Eisothistos ?Z 

Pleopod 1 in 9, rami free 4 

4. Pereopods 1 and 2 subchelate, of similar size, propodi not noticeably 

inflated 5 

Pereopod 1 much larger and propodus more expanded, than 

pereopod 2 Minyanthura

Amakusanthura 17 

5. Integument noticeably pitted; mandible on one side lacking molar, on 

other side with spicate molar Apanthuroides P- 2° 

Integument not pitted; mandibles similar on both sides Chalixanthura p.

18 ANTHURIDEA • ANTHURIDAE 

illipedal palp of three articles; endite small or absent. Pereopod 1, propodal 

palm with step or tubercle. Pereopods 4-7, carpus triangular. Pleonites 

short, pleonites 1-4 fused, boundaries demarked by complete dorsal folds, 

pleonites 4 and 5 separated only by lateral fold, not demarked dorsally; 

pleonite 6 dorsally demarked from telson. Pleopod 1 exopod operculiform. 

Uropodal exopod often notched or excavate distally. Pleotelson with two 

basal statocysts. 

Key to species of Amakusanthura 

1. Telson thickened, with raised area at midlength, widening and sloping 

posteroventrally; uropodal exopod distally shallowly excavate 

magnified 

Telson dorsally flat, not thickened; uropodal exopod distinctly notched 

or barely excavate distally 2 

2. Integument pigmented; uropodal endopod length 1.5 times basal width 

signata 

Integument not pigmented 3 

3. Uropodal exopod distinctly notched 4 

Uropodal exopod, outer margin weakly excavate; mandibular palp 

article 3 bearing three spines lathridia 

4. Mandibular palp article 3 bearing three spines signified 

Mandibular palp article 3 bearing two spines geminsula 

Amakusanthura geminsula (Kensley, 1982) 

Figure 3A-E 

DIAGNOSIS 9 : 8.1 mm. Shallow middorsal pit on pereonites 4-6, appearing 

chalky white in life. Antennular flagellum of three articles. Antennal 

flagellum of two articles. Mandibular palp article 3 bearing two spines. Max- 

illiped with short slender endite; terminal palp article set obliquely at outer 

distal angle of penultimate article. Pereopod 1, propodal palm with step; 

carpus distally a rounded lobe. Uropodal exopod with notch. Pleotelson 

tapering in posterior half to subtruncate apex. 6: 4.8 mm. Eyes larger than 

9. Antennular flagellum of 8-9 articles. Pereopod 1, carpus with distal lobe

Amakusanthura geminsula 19 

Figure 3. Amakusanthura geminsula: Ay 9; By pereopod 1; Cy telson; Dy uropodal 

sympod and endopod; E> uropodal exopod. Amakusanthura lathridia: F, pereopod 1; 

Gy telson; H, uropodal exopod; /, uropodal sympod and endopod. 

narrowly rounded; propodal palm with rounded tubercle; irregular band of 

setae on mesial surface. 

RECORDS Carrie Bow Cay and Twin Cays, Belize, intertidal to 1.5 m; 

Jamaica.

20 ANTHURIDEA • ANTHUR1DAE 

Amakusanthura lathridia (Wagele, 1982) 

Figure 3F-I 

DIAGNOSIS 9 2.7 mm. Antennular flagellum of three articles. Antennal 

flagellum of three articles. Mandibular palp article 3 bearing three spines. 

Maxilliped lacking endite. Pereopod 1, propodal palm with distal step; 

carpus with triangular posterodistal lobe. Uropodal exopod apically acute, 

outer margin slightly excavate, sinuate; endopod length slightly less than 

twice basal width. Pleotelson elliptical, widest at about midlength, margins 

weakly serrate. 

RECORDS Cuba, interstitial beach sand just above water line. 

Amakusanthura magnifica (Menzies and Frankenberg, 1966) 

Figures 4, 5 

DIAGNOSIS 9: 13.9 mm. Maxilliped with short endite, terminal article of 

palp short, set obliquely at outer distal angle of penultimate article. Pereopod 

1, propodal palm with strong tubercle; carpus with low distal sclerotized 

lobe. Uropodal exopod ovate, with shallow distal notch; endopod twice 

longer than wide. Pleotelson with posterior margin evenly rounded to sub- 

truncate; dorsally with raised area at midlength, broadening and sloping 

B 

D 

C 

Figure 4. Amakusanthura magnifica: A> 

pereopod 1; B, telson; C, uropodal sympod 

and endopod (setae omitted); D> uropodal 

exopod (setae omitted)

Figure 5. Amakusanthura magnijica: A, 

p l e o n i t e s i n l a t e r a l v i e w ; B } p l e o n i n d o r s a l 

v i e w . 

Amakusanthura signata 21 

a w a y p o s t e r i o r l y . 6 : 1 0 . 3 m m . A n t e n n u l a r f l a g e l l u m o f 2 4 a r t i c l e s . P e r e o p o d 

1 , c a r p u s w i t h s c l e r o t i z e d d i s t a l r o u n d e d l o b e ; p r o p o d a l p a l m w i t h s t r o n g 

s c l e r o t i z e d t u b e r c l e l o n g e r t h a n i n 2 ; n u m e r o u s s e t a e o n m e s i a l s u r f a c e . 

P l e o p o d 2 , c o p u l a t o r y s t y l e t o f e n d o p o d n o t r e a c h i n g b e y o n d r a m u s . 

R E C O R D S O f f G e o r g i a , 1 7 - 1 3 7 m ; o f f F l o r i d a , 7 - 1 1 m ; C u b a ; G u l f o f 

M e x i c o . 

A m a k u s a n t h u r a s i g n a t a ( M e n z i e s a n d G l y n n , 1 9 6 8 ) 

F i g u r e 6 A - E 

D I A G N O S I S 2 : 4 . 9 m m . I n t e g u m e n t w i t h s t r o n g p a t c h e s o f p i g m e n t o n 

c e p h a l o n ; p i g m e n t s p a r s e o n p e r e o n i t e s ; t w o b a r s o n p l e o n i t e 6 . A n t e n n u l a r


Figure 6. Amakusanthura signata: A, 9; B, telson; C, pereopod 1; D, uropodal 

sympod and endopod; E> uropodal exopod. Amakusanthura signijica: F, telson; G, 

uropodal sympod and endopod; H, uropodal exopod; I, pereopod 1. 

flagellum of three articles. Antennal flagellum of three articles. Mandibular 

palp article 3 bearing three spines. Maxilliped with short endite not reaching 

base of palp article 2. Pereopod 1, carpus with strong subacute sclerotized 

lobe distally; propodal palm with transparent convex flange and step.

Anthomuda stenotelson 23 

Uropodal exopod with strong distal notch; endopod length about 1.5 times 

basal width. Pleotelson widest in posterior half, tapering posteriorly to shal- 

lowly notched apex. 6: 4.5 mm. Eyes larger than in 9 . Antennular flagellum 

of nine articles. Pereopod 1, carpus with strong distal lobe, propodal palm 

with step, convex transparent flange, and numerous setae on mesial surface. 

Pleopod 2, copulatory stylet of endopod reaching well beyond ramus. 

RECORDS Cuba; Puerto Rico, intertidal to 1.5 m; Carrie Bow Cay, Belize, 

intertidal to 24 m. 

Amakusanthura significa (Paul and Menzies, 1971) 

Figure 6F—I 


flagellum of four or five articles. Mandibular palp article 3 bearing three 

spines. Maxilliped lacking endite. Pereopod 1, propodal palm with step; 

carpus distally rounded. Uropodal exopod ovate, with distal notch; endopod 

length slightly more than twice basal width. Telson elliptical, posterior mar 

gin narrowly rounded. 

RECORDS Off Venezuela, 95 m. 

Anthomuda Schultz, 1979 

DIAGNOSIS Eyes present. Mandibular palp of three articles. Maxillipedal 

palp of four articles; endite reaching well beyond base of palp article 2. Per- 

eopods 1 and 2 similar, subchelate. Pleonites 1-6 short, free; pleonite 6 dor- 

sally demarked. Telson with single semiovate hollow representing statocyst. 

Anthomuda stenotelson Schultz, 1979 

Figure 7A 

DIAGNOSIS 9 8.8 mm. Antennular flagellum of three or four articles. An 

tennal flagellum of six articles. Pereopods 1 and 2 similar, propodi somewhat 

inflated; pereopod 1, propodal palm straight, unarmed; pereopod 2 propodal 

palm with three sensory spines. Pereopods 4—7 with rectangular carpi. 

Pleonite 6 free, posterior margin broadly bilobed. Uropodal exopod elongate- 

elliptical, distally narrowly rounded; endopod length 2.5 times basal width, 

distally broadly rounded. Telson narrowly lanceolate, posteriorly narrowly 

rounded, with single open hollow statocyst. 

RECORDS Off Bermuda, 90 m.

Figure 7. Anthomuda stenotelson: A, 9. Apanthura cracenta: B, 9; C, telson; D, 

uropodal endopod and sympod; E, uropodal exopod. Apanthura crucis: F, uropodal 

exopod; G, uropodal endopod and sympod. Apanthura harringtoniensis (from Wagele, 

1981): H, uropodal endopod and sympod; /, uropodal exopod.

Apanthura Stebbing, 1900 

Apanthura cracenta 25 

DIAGNOSIS Integument sometimes pigmented. Eyes present. 9: Antennu 

lar flagellum of three articles. Antennal flagellum of two to four articles. 

Mandibular palp of three articles, terminal article bearing distal spines. 

Maxillipedal palp of three articles; endite small, or lacking. Pereopod 1, pro- 

podal palm usually with step or tubercle; propodus inflated. Pereopods 4-7, 

carpus triangular. Pleonites 1-5 fused; pleonite 6 dorsally demarked. 

Pleopod 1, exopod operculiform. Uropodal exopod ovate, sometimes distally 

notched or excavate. Pleotelson with two basal statocysts. 6 antennular 

flagellum of about 10 articles. 

REMARKS Differentiation of species, especially if long preserved and the 

pigmentation is lost, depends on subtle features of the mandible, maxilliped, 

pereopods, and pleotelson. 

Key to species of Apanthura 

1. Uropodal exopod distally notched or excavate 2 

Uropodal exopod distally entire; uropodal endopod length subequal to 

width cruris 

2. Uropodal exopod distally notched; uropodal endopod length 2.4 times 

basal width cracenta 

Uropodal exopod faintly excavate or sinuate; uropodal endopod length 

about twice basal width harringtoniensis 

Apanthura cracenta Kensley, 1984 

Figure 7B-E 

DIAGNOSIS 9: 4.6 mm. Antennular flagellum of three articles. Antennal 

flagellum of two articles. Mandibular palp article 3 bearing four spines. 

Maxilliped with short rounded endite; terminal palp article set at oblique 

angle on, and less than half length of penultimate article. Pereopod 1, carpus 

triangular with acute sclerotized tip overlapping base of palm; propodal 

palm with rounded tubercle near midlength. Uropodal exopod deeply 

notched; endopod ovate, length 273 basal length. Pleotelson lanceolate. 6: 

3.8 mm. Antennular flagellum of six articles. Antennal flagellum of two arti 

cles. Pleopod 2, copulatory stylet of endopod reaching by half its length be 

yond ramus.

26 ANTHUR1DEA • ANTHURIDAE 

RECORDS Turks and Caicos Islands, 1 m; Carrie Bow Cay, Belize, intcrti- 

dal on reef crest, 2 m. 

Apanthura crucis (Barnard, 1925) 

Figure 7F,G 

DIAGNOSIS 9: 5.9 mm. Antennular flagellum of four articles. Antennal 

flagellum of four articles. Mandibular palp article 3 with eight spines. Max- 

illiped lacking endite; terminal palp article more than half length of penulti 

mate article. Pereopod 1, propodal palm with low rounded tubercle at mid- 

length. Uropodal exopod ovate, outer margin evenly convex; endopod 

subcircular. Pleotelson anteriorly narrow, widest at midlength, posterior 

margin broadly rounded. 6: 6.2 mm. Antennular flagellum of 12 or 13 arti 

cles. Antennal flagellum of four articles. Pereopod 1, propodal palm with low 

rounded tubercle; dense band of setae on mesial surface. Copulatory stylet on 

endopod of pleopod 2 reaching a third of its length beyond ramus. 

RECORDS Turks and Caicos Islands, 1 m; St. Croix, U.S. Virgin Islands, 8 

m. 

Apanthura harringtoniensis Wagele, 1981 

Figure 7H,I 

DIAGNOSIS 9 6.0 mm. Antennular flagellum of four articles. Antennal 

flagellum of four articles. Mandibular palp article 3 bearing two spines. 

Maxillipedal endite short, not reaching base of palp article 2. Pereopod 1, 

propodal palm stepped; carpus distally bluntly triangular. Uropodal exopod 

apically acute, outer margin distally sinuate; endopod length about twice 

basal width. Pleotelson with posterolateral margins faintly denticulate; apex 

subtruncate and bearing several setae. 

RECORDS Harrington Sound, Bermuda. 

Apanthuroides Menzies and Glynn, 1968 

DIAGNOSIS Eyes present. Mandibular palp of three articles; body of mand 

ible anteriorly produced, molar absent on left side, present as narrow 

spikelike process on right. Maxillipedal palp of three articles; endite present, 

reaching palp article 2. Pereopod 1, propodus barely expanded, palm un-

Chalixanthura 27 

armed. Pereonite 7 less than half length of pereonite 6. Pereopods 4-7, carpi 

with free anterior margin shorter than posterior margin. Pleonites 1—5 fused, 

6 fused with telson. Pleopod 1, both rami forming operculum. 

Apanthuroides millae Menzies and Glynn, 1968 

Figure 8 

DIAGNOSIS 9: 4.5 mm. Integument with diffuse brown pigmentation and 

numerous shallow pits. Antennular flagellum of four articles. Antennal 

flagellum of seven articles. Pleotelson with strong middorsal ridge. 6: 5.0 

mm. Integumental pigment stronger than in 9, in irregular brown patches. 

Eyes larger than in 9 . Antennular flagellum of six or seven articles. Antennal 

flagellum of seven articles. Pereopod 1, propodal palm armed with five 

fringed spines. 

RECORDS Carrie Bow Cay, intertidal to 30 m; Puerto Rico, intertidal. 

REMARKS The highly modified mandible probably indicates some spe 

cialized form of feeding, but this has yet to be discovered. 

Chalixanthura Kensley, 1984 

DIAGNOSIS Eyes present, enormously enlarged, especially ventrally in

Figure 8. Apanthuroides millae: A, $; B, right mandible; C, left 

maxilliped; E, uropodal exopod; F, pleopod 1; G, pereopod 1.

Chalixanthura lewisi Kensley and Snelgrove, 1987 

Figure 9A-D 

Cortezura 29 

DIAGNOSIS 9: 3.1 mm. Integument lacking pigment. Antennular flagellum 

of three articles. Antennal flagellum of seven articles. Maxillipedal endite 

large, apically acute. Pereopod 1 propodus slightly expanded, palm with few 

(3) simple setae. Pereopod 7 propodus with two elongate anterodistal fringed 

spines. Uropodal and telsonic margins serrate. 6: 2.2 mm. Antennular 

flagellum of 11 articles. Antennal flagellum of six articles. Maxilliped re 

duced, lacking endite. Pereopod 1, propodal palm with two spines plus seven 

spines on mesial surface. Pereopod 7 as in 9. 

RECORDS Barbados, in Madracis mirabilis coral, 9—15 m. 

Chalixanthura scopulosa Kensley, 1984 

Figure 9E-J 

DIAGNOSIS 9: 2.5 mm. Antennular flagellum of three articles. Antennal 

flagellum of seven articles. Maxillipedal endite short, not reaching base of 

palp article 4. Pereopod 1, propodal palm unarmed. Pereopod 7, propodus 

with single elongate anterodistal fringed spine. Uropodal exopod margin de 

eply incised; endopod ovate, margins serrate. Telson elongate-ovate, pos 

terior margin serrate. 6: 2.6 mm. Eyes considerably larger and with more 

ommatidia than in 9. Antennular flagellum of seven articles. Antennal 

flagellum of four articles. Pereopod 1 propodal palm with three sensory 

spines. Uropodal exopod deeply incised. 

RECORDS Carrie Bow Cay, Belize, 0.1 m. 

Cortezura Schultz, 1977 

DIAGNOSIS Eyes small, weakly pigmented. Antennular flagellum of two 

short articles. Antennal flagellum of single article. Mandibular palp of two 

articles. Maxillipedal palp of two articles, terminal article small; short endite 

present. Pereopod 1, propodus inflated. Pereopods 4-7 with carpus having 

anterior margin shorter than posterior. Pleonites 1—5 fused; pleonite 6 dor- 

sally demarked. Pleopod 1, exopod operculiform. Pleotelson with two basal 

statocysts. 

*

Figure 9. Chalixanthura lewisi: A, 6; By 9; C, uropodal exopod; D, pleopod 1. 

Chalixanthura scopulosa: E, 6] F, 9; G, maxilliped; H, pereopod 1; 7, pleopod \\J, 

uropodal exopod.

Cortezura confixa (Kensley, 1978) 

Figure 10 

Cyathura (Cyathura) 31 

DIAGNOSIS 9 13.4 mm. Antennular peduncle articles 1 and 2 each with 

clump of ventrally directed setae. Antennal peduncle article 2 and antennu 

lar peduncle article 1 locked together. Mandibular palp of two articles, distal 

article twice length of proximal. Pereopod 1, propodus expanded, especially 

posteriorly; palm concave, with strong tubercle and irregular band of setae 

on mesial surface. Uropodal exopod ovate, apically rounded; endopod length 

1.5 times greatest width. Telson ovate, posteriorly faintly narrowed, prox- 

imally thickened, with faint ridges diverging posteriorly. 

RECORDS Gubagua Island, Venezuela, 4-10 m. 

REMARKS This species was described under the generic name Venezanthura 

Kensley (1978). The only other known species of Cortezura is the type of the 

genus, C. penascoensis Schultz, 1977, from California. 

Cyathura Norman and Stebbing, 1886 

DIAGNOSIS Eyes present or absent. Antennular flagellum of one to three 

articles in 9 . Antennal flagellum of one to three articles. Mandibular palp of 

three articles. Maxillipedal palp of two articles; endite reduced or absent. 

Pereopod 1, propodus inflated. Pereopods 4-7, carpi triangular. Pleopod 1 

exopod operculiform. Pleonites 1-5 short, fused; pleonite 6 fused with telson, 

sometimes dorsally demarked. Pleotelson with two basal statocysts. 

Key to subgenera of Cyathura 

1. Pleonite 6 not dorsally demarked from telson; articulation of uropodal 

exopod very short; exopod not adpressed to telson dorsally 

Stygocyathura 

Pleonite 6 dorsally demarked from telson; articulation of uropodal 

exopod relatively elongate; exopod adpressed dorsally to telson 

Cyathura (Cyathura) Norman and Stebbing, 1886 

Cyathura 

DIAGNOSIS Integument usually strongly pilose or setose; eye and body pig-

Figure 10. Cortezura confixa: A, 9; B, cephalon and pereonite 1, lateral view 

(pereopod 1 removed); C, antennule; D, pleon, lateral view; E, mandible; F, 

maxilliped; G, pereopod 1; H, pereopod 7.

Cyathura (Stygocyathura) 33 

mentation usually present. Pereopod 1, propodal palm armed with tubercle. 

Pleopod 1, protopod with retinaculae. Uropodal exopod articulation relative 

elongate along lateral margin of sympod; exopod well developed, ovate. 

Pleonite 6 dorsally demarked from telson. Marine or estuarine forms. 

Cyathura (Cyathura) cubana Negoescu, 1979 

Figure 11A,B 

DIAGNOSIS Ovigerous 9: 7.0 mm. Antennular flagellum of two articles. An- 

tennal flagellum of one article. Pereopod 1, propodal palm with rounded lobe 

in proximal half Maxillipedal palp with distal article 0.34 times length of 

proximal article; small rounded endite present. Dorsal pigmentation consist 

ing of irregular brown mottling. 6: 5.5 mm. Antennular flagellum of four 

t 

articles. Antennal flagellum of three articles. Pereopod 1, propodal palm with 

rounded lobe in proximal half. Copulatory stylet elongate-cylindrical, api- 

cally narrowed and flexed. 

RECORDS Cuba, in mangroves, 2.5-7.0 m; Salt Creek, Belize, in man 

groves, 1.5 m. 

Cyathura (Stygocyathura) Botosaneanu and Stock, 1982 

DIAGNOSIS Eye and body pigmentation absent. Body sparsely pilose or set 

ose. Tendency toward elongation of some appendages, especially propodus of 

pereopods 2—7. Pereopod 1, propodal palm lacking strong tubercle. Pleopod 

1, protopod lacking retinaculae. Pleonite 6 fused with telson, not dorsally 

demarked. Uropodal exopod with very short articulation on sympod, not 

adpressed dorsally to telson. Cave or hypogean forms. 

REMARKS The ten species of Stygocyathura from the area covered in this work 

are morphologically very similar, with specific differences, although real, 

being very subtle. A dichotomous key would be cumbersome and require 

considerable dissection of mouthparts. The copulatory stylet of the male pro 

vides a valuable specific feature but males are not always available. Instead 

of a key, we have provided a list of species with their total lengths and lo 

calities (Table 1). Given the very restricted distribution of these cave species, 

material from localities not listed here should be treated as potentially un- 

described, and the material compared with descriptions, especially those of 

Botosaneanu and Stock (1982).


Figure 11. Cyathura (Cyathura) cubana: A, 9; By pereopod 1,9. Cyathura 

(Stygocyathura) cuborientalis: C, pereopod 1, 9; D, telson. Cyathura (Stygocyathura) 

curassavica: E, pereopod 1, 9; F9 telson. Cyathura (Stygocyathura) hummetincki: G, 

pereopod 1, 9; H, telson. Cyathura (Stygocyathura) motasi: I, telson; J, pereopod 1, 

9. 

Cyathura (Stygocyathura) cuborientalis Botosaneanu and Stock, 1982 

Figure 11C,D 

DIAGNOSIS 9 6.8 mm. Pereopod 1, propodal palm straight, bearing about

Cyathura (Stygocyathura) hummelincki 35 

TABLE l. CARIBBEAN SPEGIES OF Cyathura (Stygocyathura), THEIR TOTAL LENGTHS 

(MM) AND LOCALITIES 

C. (S.) cuborientalis 

C. (S.) curassavica 

C. (S.) hummelincki 

C. (S.) motasi 

C. (S.) orghidani 

C. (S.) parapotamica 

C. (S.) salpiscinalis 

C. (S.) sbordonii 

C. (S.) specus 

C. (S.) univam 

9 6.8 

6 7.0, 9 9.2 

6 4.75, 9 8.5 

6 6.8, 9 10.0 

9 8.0 

ovig. 9 3.6, 9 4.1 

6 5.6, 9 7.3 

6, 9 9.0 

6 18.0, 9 19.8 

9 10.0 

Cuba 

Curasao 

Aruba 

Haiti 

Cuba 

Jamaica 

Haiti 

Vera Cruz, Mexico 

Cuba 

Venezuela 

10 pectinate marginal spines; low triangular ridge present. Pleotelson evenly 

tapering to notched apex; angle of apex about 90°. 

RECORDS Oriente Province, Cuba, interstitial in river alluvia. 

Cyathura (Stygocyathura) curassavica Stork, 1940 

Figures 11E,F; 12 

DIAGNOSIS 6 7.0 mm, 9 9.2 mm. Pereopod 1, propodal palm gently con 

vex, bearing about 10 pectinate spines. Pleotelsonic margins in anterior two- 

thirds subparallel, tapering gently to finely notched apex; angle of apex less 

than 90°. 

RECORDS Curagao, from pits and wells. 

Cyathura (Stygocyathura) hummelincki Botosaneanu and Stock, 1982 

Figure 11G,H 

DIAGNOSIS 6 4.75 mm, 9 8.5 mm. Pereopod 1, propodal palm gently sinu 

ate, bearing 11-23 marginal pectinate spines. Pleotelsonic margins faintly 

concave in midregion, posterior margin evenly convex, apex with slight 

notch. 

RECORDS Aruba, in pits, wells, and temporary water sources.


Figure 12. Cyathura 

(Stygocyathura) curassavica: 

pleon. 

Cyathura (Stygocyathura) motasi Botosaneanu and Stock 

Figure 111 J 

? 

1982 

DIAGNOSIS 6 . 8 m m , 9 1 0 . 0 m m . P e r e o p o d 1 , p r o p o d a l p a l m g e n t l y s i n u 

a t e , b e a r i n g 1 1 - 1 8 m a r g i n a l p e c t i n a t e s e t a e . P l e o t e l s o n i c m a r g i n s t a p e r i n g 

e v e n l y t o s l i g h t l y n o t c h e d a p e x ; a n g l e o f a p e x l e s s t h a n 9 0 ° . 

RECORDS Haiti, from wells. 

Cyathura (Stygocyathura) orghidani Negoescu Vladescu, 1983 

Figure 13A,B 

DIAGNOSIS 8.0 mm. Pereopod 1, propodal palm slightly convex, bearing 

about 11 marginal pectinate setae. Pleotelson, angle of apex obtuse, with 

small notch. 

RECORDS Pinar del Rio Province, Cuba, from freshwater lake in cave. 

C y a t h u r a ( S t y g o c y a t h u r a ) p a r a p o t a m i c a B o t o s a n e a n u a n d S t o c k , 1 9 8 2 

Figure 13D,E 

DIAGNOSIS m m ( o v i g . 9 3 . 6 m m ) . P e r e o p o d 1 , p r o p o d a l p a l m 

a t e , b e a r i n g e i g h t m a r g i n a l p e c t i n a t e s e t a e . P l e o t e l s o n g e n t l y t a p e r i n g t o 

n o t c h e d a p e x ; a n g l e o f a p e x a b o u t 9 0 ° . 

R E C O R D S J a m a i c a , f r o m r i v e r a l l u v i a .

Figure 13. Cyathura (Stygocyathura) orghidani (from Negoescu, 1983): A, telson; B, 

pereopod 1,9. Cyathura (Stygocyathura) sbordonii: C, pereopod 1, 9 (from Argano, 

1971). Cyathura (Stygocyathura) parapotamica: D> telson; E, pereopod 1, 9. Cyathura 

(Stygocyathura) salpicinalis: F, pereopod 1, 9; G, telson. Cyathura (Stygocyathura) 

specus: H, 9; I, telson and uropods; J, pereopod 1, 9. Cyathura (Stygocyathura) 

univam: K, telson.


Cyathura (Stygocyathura) salpiscinalis Botosaneanu and Stock, 1982 

Figure 13F,G 

DIAGNOSIS 6 5.6 mm, 9 7.3 mm. Pereopod 1, propodal palm sinuate, 

bearing up to 15 marginal pectinate setae, and with distinct triangular ridge. 

Pleotelson gently tapering, with slight apical eminence. 

RECORDS Haiti, from alluvia of lake. 

Cyathura (Stygocyathura) sbordonii Argano, 1971 

Figure 13C 

DIAGNOSIS 6 and 9 9.0 mm. Pereopod 1, propodal palm convex, bearing 

up to 16 marginal pectinate setae. Pleotelson with angle of apex obtuse. 

RECORDS Vera Cruz, Mexico, from freshwater in cave. 

Cyathura (Stygocyathura) specus Bowman, 1965 

Figure 13H-J 

DIAGNOSIS 6 18.0 mm, 9 19.8 mm. Pereopod 1, propodal palm almost 

straight, bearing up to 15 marginal pectinate setae. Pleotelson with angle of 

apex obtuse. 

RECORDS Las Villas Province, Cuba, from freshwater lake in cave. 

Cyathura (Stygocyathura) univam Botosaneanu, 1983 

Figure 13K 

DIAGNOSIS 9 10.0 mm. Pereopod 1, propodal palm gently convex, bearing 

22 marginal pectinate setae. Pleotelson posteriorly broadly rounded, apex 

emarginate. 

RECORDS Peninsula de Morocoy, Venezuela, from phreatic water in cave. 

Eisothistos Haswell, 1884 

DIAGNOSIS Mouthparts forwardly produced, tailfan spiny, indurate. Eyes 

present, larger and with more ommatodia in 6 than in 9. Pereonites some 

times elongate. Mandible with strong incisor, reduced lamina dentata, palp 

and molar lacking. Maxilliped lacking endite, palp slender, of three to five

Eisothistos teri 39 

articles. Pereopods 1-3 not subchelate, propodi relatively elongate, mini 

mally expanded. Pleopod 1 9, rami fused, together forming operculum; 6 

rami separate. Pleonites free, short, longer in 6 than in 9. Telson lacking 

statocysts. 

REMARKS Wagele (1979) first recorded species of Eisothistos preying on ser- 

pulid polychaete worms in their tubes. 

The genus contains about 12 species in the Pacific, Indian Ocean, Carib 

bean, Antarctic, and Mediterranean. 

Key to species of Eisothistos 

1. 9 telson with middorsal spines; 6 pereopod 1 propodal palm with 19 

or 20 spines teri 

9 telson lacking middorsal spines; 6 pereopod 1 propodal palm with 

11 spines petrensis 

Eisothistos petrensis Kensley, 1984 

Figure 14A-E 

DIAGNOSIS 9: 4.0 mm. Antennular flagellum of six articles. Antennal 

flagellum of six articles. Telson posteriorly faintly bilobed, margin strongly 

serrate; faint anterior middorsal ridge; middorsally unarmed. Pereopod 1, 

propodal palm unarmed. Pleopod 1 rami fused for V12 of length. 6: 2.0 mm. 

Antennular flagellum of eight articles. Antennal flagellum of six articles. Per 

eopod 1 propodal palm with 11 fringed spines. Telson narrower than in 9, 

with middorsal ridge running almost entire length. 

RECORDS Carrie Bow Cay, Belize, 0.1-36 m; Looe Key, Florida, 5-6 m; 

Turks and Caicos Islands, 1.0 m; St.Thomas, U.S. Virgin Islands, 7-10 m. 

Eisothistos teri Kensley and Snelgrove, 1987 

Figure 14F-H 

DIAGNOSIS 9: 3.2 mm. Basal antennular peduncle article bearing triangu 

lar apically rounded laminate process; flagellum of seven articles. Antennal 

flagellum of six articles. Mandible with biserrate lamina dentata. Pereopod 1 

propodal palm unarmed. Pleopod 1 rami fused for 3 A of length. Uropodal 

exopod with one or two strong slightly recurved spines on dorsal surface.


Figure 14. Eisothistos petrensis: Ay 9; By uropodal sympod and endopod; Cy 

uropodal exopod; D, pleopod 1; Ey pereopod 1, 6. Eisothistos ten: Fy telson and 

uropod; Gy pleopod 1; Hy pereopod 1, 6. 

Telson with eight or nine slightly recurved middorsal teeth becoming longer 

posteriorly. 6: 2.0 mm. Antennular flagellum of eight articles. Antennal

Haliophasma curri 41 

flagellum of six articles. Pereopod 1 propodal palm with 19 or 20 fringed 

spines. Uropodal exopod lacking dorsal teeth. Telson lacking middorsal 

teeth; posterior margin incised into 12 acute or narrowly rounded teeth. 

RECORDS Barbados, in Madracis mirabilis coral, 9—15 m. 

Haliophasma Haswell, 1881 

DIAGNOSIS Eyes present. Integument often indurate, with scattered pitting. 

Antennular flagellum usually of two articles. Antennal flagellum of 4—7 arti 

cles. Mandibular palp of three articles. Maxillipedal palp of two articles, 

article 2 smaller than article 1. Pereopod 1, propodus expanded. Pereopods 

4-7, carpi roughly rectangular. Pleopod 1 exopod operculiform. Pleonites 1 — 

5 short, fused; pleonite 6 usually demarked from telson. Latter with two 

basal statocysts. 6 often with more elongate form than 9. Antennular 

flagellum multiarticulate. Eyes larger. 

Key to species of Haliophasma 

1. Telson posteriorly narrowly rounded; dactylus of pereopod 1 dentate 

Telson posteriorly broadly rounded; dactylus of pereopod 1 entire 

Haliophasma curri Paul and Menzies, 1971 

Figure 15A-C 

valeriae 


flagellum of five articles. Mandibular palp article 3 with five spines. Max 

curri 

illipedal palp, terminal article small, set obliquely at distolateral angle of 

article 1. Pereopod 1, carpus triangular, posterodistal margin crenulate, dis- 

tally rounded; propodal palm crenulate, with low rounded proximal lobe. 

Pleonite 6 dorsally demarked, narrow, with middorsal point in posterior 

margin. Uropodal exopod elongate, outer margin sinuate, dentate; endopod 

ovate, distally narrowed, length twice greatest width, outer margin dentate. 

Telson parallel sided, posterior margin broadly rounded. 

RECORDS Off Venezuela, 95 m; Culebra Island, Bay of Panama, intertidal.


Figure 15. Haliophasma curri: A, 9; B, pereopod 1; C, uropod. Haliophasma valeriae: 

D, 9; E, pereopod 1; F, uropod. 

Haliophasma valeriae Paul and Menzies, 1971 

Figure 15D-F 

DIAGNOSIS 9 6.5 mm. Body long and slender. Antennular flagellum of

Malacanthura 43 

three articles. Antennal flagellum of six articles. Mandibular palp article 3 

with four spines. Pereopod 1, carpus triangular, tipped with acute tooth; 

propodus elongate, palm with about seven teeth, five fringed spines on mesial 

surface; unguis of dactylus strongly flexed; margin of dactylus with three 

strong triangular teeth. Uropodal exopod elongate, apically acute, margins 

serrate; endopod length little more than twice greatest width, outer margin 

serrate, apex acute. Telson elongate-elliptical, apically narrowly rounded; 

strong middorsal longitudinal rounded ridge running almost entire length. 


REMARKS Wagele (1981) made this species the type of his new genus Ne- 

manthura, based primarily on the elongate form of the body and appendages. 

Haliophasma irmae Paul and Menzies, 1971, from the same locality as the 

above species, is probably the same species. 

Licranthura Kensley and Schotte, 1987 

DIAGNOSIS Serrate process on antennal peduncle article 3. Mandibular 

palp of three articles; molar lacking. Maxillipedal endite short. Pereopod 1 

larger than pereopods 2 and 3. Pereopods 4—7, carpi triangular. Pleonites 

short, free. Pleopod 1, both rami forming operculum. Pleotelson lacking 

statocysts. 

Licranthura amyle Kensley and Schotte, 1987 

Figure 16 

DIAGNOSIS 9 3.8 mm. Eyes small, pigmented. Antennular flagellum of 

three articles. Antennal peduncle article 3 with lamellar expanded process, 

serrate on mesial margin; flagellum of six articles. Maxillipedal palp of five 

articles; very short endite. Pereopod 1, propodal palm unarmed. Uropodal 

and telsonic margins serrate. 

RECORDS Carrie Bow Cay, Belize, 0-25 m, in coral rubble. 

Malacanthura Barnard, 1925 


palp of three articles, terminal article usually broadly ovate. Pereopod 1, 

propodus expanded. Pereopods 4-7 with carpi roughly rectangular. Pleopod


Figure 16. Licranthura amyle: A, 9; B, antennule and antenna; C, telson and 

uropod. 

1, exopod operculiform. Pleonites 1—5 short, fused; pleonite 6 dorsally de- 

marked. Pleotelson with two basal statocysts.

Malacanthura caribbica Paul and Menzies, 1971 

Figure 17 

Mesanthura 45 

DIAGNOSIS 9 27.1 mm. Integument moderately indurate. Antennular 

flagellum of seven articles. Antennal flagellum of four articles. Mandibular 

palp, article 3 with comb of 11 spines. Maxillipedal palp, terminal article 

broadly ovate, penultimate article with row of seven spines on mesial margin. 

Pereopod 1 propodus expanded, palm straight, with few spines on mesial 

margin. Uropodal exopod barely reaching base of endopod, narrow, apically 

acute, outer margin sinuate, serrate; endopod set obliquely on sympod, mar 

gin serrate, apically acute. Telson lanceolate, apically narrowly rounded, 

with strong longitudinal middorsal carina. 

RECORDS Off Venezuela, 95 m; off Colombia, 42-44 m. 

REMARKS Malacanthura cumanensis Paul and Menzies, 1971, described from 

the same locality as M. caribbica, was shown to be the latter species (Kensley, 

1980). 

Mesanthura Barnard, 1914 

DIAGNOSIS 9: Dorsal integument with (usually) species-specific pigment 

pattern; pigment persistent in alcohol. Mandibular palp of three articles, 

terminal article with row of spines, number of which specific for species. 

Maxilliped with endite either very reduced or absent; palp of three articles, 

with terminal article usually about half length of penultimate article, suture 

transverse. Pereopod 1, propodus expanded, palm often with step. Pereopods 

2 and 3, propodi not expanded. Pereopods 4—7, carpi roughly triangular, 

with anterior margin shorter than posterior margin. Pleonites 1—5 fused, 

pleonite 6 dorsally demarked. Pleopod 1, exopod operculiform. Telson with 

two basal statocysts. 6: Eyes larger than in 9. Antennular flagellum of sel 

dom more than 10 articles bearing numerous aesthetascs. Mouthparts, espe 

cially body of mandible, reduced. Pereopod 1, propodus bearing dense band 

of spines on mesial surface near palm. Pigment pattern more diffuse than in 

9, extending onto ventral surface. 

REMARKS Mesanthura is a relatively large genus of about 30 species, re 

corded from most tropical and temperate seas, in shallow habitats. The 

males of few species have been recorded; by themselves, males are difficult to 

identify as the dorsal pigment pattern characteristic of the female breaks 

down and spreads onto the ventrum.

Figure 17. Malacanthura caribbica: A, 9; 

maxilliped; E, uropod; F, pleopod 1.

Key to species of Mesanthura (9 only) 

Mesanthura bivittata 47 

1. Pigment in tiny evenly scattered chromatophores over body; cephalon 

with solid patch of pigment; mandibular palp article 3 with seven 

spines punctillata 

Pigment not evenly scattered over body 2 

2. Pereonites 4 and 5 with patch of pigment, sometimes with unpigmented 

area in middle of patch 3 

Pereonites 4 and 5 lacking fairly solid patch of pigment 6 

3. Pereonites 4 and 5 lacking unpigmented area in pigment patch; five 

transverse lines on pigment on pleon; mandibular palp article 3 with 

six spines paucidens 

Pereonites 4 and 5 with unpigmented area in pigment patch; pleon 

lacking transverse pigment lines 4 

4. Pigment in obvious double longitudinal bands on pereon and pleon; 

mandibular palp article 3 with nine spines bivittata 

Pigment not in obvious double bands 5 

5. Unpigmented area in middle of pigment patch of pereonites 1-3; 

mandibular palp article 3 with 10 spines pulchra 

No unpigmented area in middle of pigment of pereonites 1-3; 

mandibular palp article 3 with eight spines looensis 

6. Pigment of pereon in fine reticulate lines; mandibular palp article 3 

with six spines reticulata 

Pigment of pereon not in fine reticulate lines; mandibular palp article 3 

with four spines 7 

7. Pigment in more or less complete rings on pereonites 1—6 hopkinsi 

Pigment in strong transverse posterior bars on pereonites 4-7 . .fasciata 

Mesanthura bivittata Kensley, 1987a 

Figures 18A, 20A-D 

DIAGNOSIS 6 5.2 mm, ovigerous $ 7.8 mm. Pigment in obvious double 

longitudinal bands on pereon and pleon. Mandibular palp article 3 with nine 

spines. Maxilliped lacking endite. Pereopod 1, propodal palm with rounded 

lobe. 

RECORDS Twin Cays, Belize, under red mangroves, 1-2 m.

Mesanthura 

W® 

A-O 

I, 

• • 

V > : < < 

^ 

A 

rr- 

/v\' 

a * 

Mesanthura bivittata; B, Mesanthura fasciata 

^ 

«-tf 

4 

-1' 

V 

C

Mesanthura fasciata 49 

Figure 19. A, Mesanthura paucidens; B, Mesanthura pulchra; C, Mesanthura punctillata; 

D, Mesanthura reticulata. 

Mesanthura fasciata Kensley, 1982 

Figures 18B, 20E-H 

DIAGNOSIS 9 4.5 mm. Pigment in triangular patch on head, open irregular 

rings on pereonites 1-3; pereonites 4—7 with strong transverse posterior bar;

Figure 20. Mesanthura bivittata: A, mandible; By maxilliped; C, pereopod 1; D, 

uropodal exopod. Mesanthura fasciata: E, mandible; F, maxilliped; G, pereopod 1; 

H, uropodal exopod. Mesanthura hopkinsi: I, mandible; J, maxilliped; K, pereopod 1; 

L, uropodal exopod.

Mesanthura paucidens 51 

five bars on pleon. Mandibular palp article 3 with four spines. Maxilliped 

with very reduced endite. Pereopod 1, propodal palm with step. 

RECORDS Looe Key, Florida, 5-6 m; Cozumel, Mexico; Jamaica; Carrie 

Bow Cay, Belize, 0.2-6 m. 

Mesanthura hopkinsi Hooker, 1985 

Figures 18C, 20I-L 

DIAGNOSIS 9 2.4 mm. Pigment in triangle on cephalon; in irregular rings 

on pereonites 1-6; four transverse bars on pleon. Mandibular palp article 3 

with four spines. Maxilliped lacking endite. Pereopod 1, palm of propodus 

lacking step. 

RECORDS Looe Key, Florida, 0.5 m; Florida Middlegrounds, Gulf of Mex 

ico, 55 m. 

Mesanthura looensis Kensley and Schotte, 1987 

Figures 18D, 21A-D 

DIAGNOSIS 9 10.0 mm. Pigment in solid patches on cephalon, pereonites, 

pleon, uropods and telson; pereonites 4—6 with open central area in patch. 

Mandibular palp article 3 with eight spines. Maxilliped lacking endite. Per 

eopod 1, propodal palm with step. 

RECORDS Looe Key, Florida, 1 m. 

Mesanthura paucidens Menzies and Glynn, 1968 

Figures 19A, 2 IE-1 

DIAGNOSIS 9: 6.6 mm. Pigment in roughly rectangular to ovate patches on 

cephalon and pereonites; in five transverse lines connected laterally on pleon. 

Mandibular palp article 3 with six spines. Maxilliped with short narrow en 

dite. Pereopod 1 propodal palm with step. 6: 6.4 mm. Antennular flagellum 

of seven articles. Pigment more diffuse than in 9 but retaining five pleonal 

bars. 

RECORDS Looe Key, Florida, 5-6 m; Carrie Bow Cay, Belize, 15.2 m; Pu 

erto Rico, intertidal; Jamaica.


Figure 21. Mesanthura looensis: A, mandible; B, maxilliped; C, pereopod 1; D, 

uropodal exopod. Mesanthura paucidens: Ey mandible; F9 maxilliped; G, pereopod 1, 

9; H, pereopod 1, 8; I> uropodal exopod. Mesanthura pulchra: J, mandible; K, 

maxilliped; L, pereopod 1, 6; M, pereopod 1, 9; JV, uropodal exopod. 

Mesanthura pulchra Barnard, 1925 

Figures 19B, 21J-N 

DIAGNOSIS 9: 9.3 mm. Pigment pattern in roughly rectangular patches on 

cephalon and pereonites 1-6, with open oval middorsal area in patch. Man 

dibular palp article 3 with 10 spines. Maxilliped lacking endite. Pereopod 1, 

propodal palm with step. 6: 5.4 mm. Antennular flagellum of eight articles.

Minyanthura corallicola 53 

RECORDS Egmont Key, Florida, 18.3-36.6 m; Looe Key, Florida, 0.5-12 

m; Dry Tortugas; Turks and Caicos Islands, 1 m; Puerto Rico, intcrtidal to 

1.5 m; Carrie Bow Cay, Belize, intertidal to 1.5 m; St.Thomas and St.John's, 

U.S. Virgin Islands; Cozumel, Mexico. 

REMARKS Menzies and Glynn (1968) recorded this species as M. decorata 

from Puerto Rico, while Menzies and Kruczynski (1983) recorded it as M. 

Jloridensis. 

Mesanthura punctillata Kensley, 1982 

Figures 19C, 22A-F 

DIAGNOSIS 9: 6.4 mm. Pigment in solid patch between eyes, rest of pig 

ment on pereon and pleon with chromatophores scattered, diffuse, not in any 

regular pattern. Mandibular palp article 3 with seven spines. Maxilliped 

lacking endite; terminal palp article semicircular. Pereopod 1 propodal palm 

with step. 6: 4.5 mm. Antennular flagellum of 10 articles. 

RECORDS Turks and Caicos Islands, 1 m; Carrie Bow Cay, 0.2-20 m. 

Mesanthura reticulata Kensley, 1982 

Figures 19D, 22G-J 

DIAGNOSIS 9 6.1 mm. Dorsal pigment pattern a network of 

chromatophores arranged in fine lines. Mandibular palp article 3 with six 

spines. Maxilliped lacking endite. Pereopod 1 propodal palm with step. 

RECORDS Carrie Bow Cay, 10-24 m. 

Minyanthura Kensley, 1982 

DIAGNOSIS Mandible lacking molar and palp. Maxillipedal palp of five ar 

ticles; large apically acute endite present. Pereopods 4-7, carpi rectangular. 

Pleopod 1, both rami forming operculum. Pleonites 1-5 fused; pleonite 6 

fused with telson, not dorsally demarked. Telson with two basal statocysts. 

Minyanthura corallicola Kensley, 1982 

Figure 23 

DIAGNOSIS 9: 1.7 mm. Antennular flagellum of one article. Antennal

Figure 22. Mesanthura punctillata: A, mandible, 9; B, mandible 8; C, maxilliped; D, 

pereopod I, 9; E, pereopod 1, 6\ F9 uropodal exopod. Mesanthura reticulata: G, 

mandible; H, maxilliped; /, pereopod \\ J, uropodal exopod.

Minyanthura corallicola 55 

Figure 23. Minyanthura corallicola: A, 9; B, telson and uropod; C, pleopod 1; D, 

mandible; E, maxilliped; F, pereopod 1. 

flagellum of four articles. Pereopod 1, propodus somewhat inflated. Uropo- 

dal rami and posterior telsonic margin serrate. 6: 1.3 mm. Eyes larger than 

in 9. Antennular flagellum of three articles. Antennai flagellum of two 

articles. 

RECORDS Carrie Bow Cay, Belize, 6-24 m; Barbados, 9-15 m; Jamaica.


Pendanthura Menzies and Glynn, 1968 

DIAGNOSIS 9 : Integument with some red-brown dorsal pigmentation. Eyes 

small, pigmented. Antennular flagellum of two articles. Antennal flagellum 

of one article. Mandibular palp of single reduced article; incisor, molar, and 

lamina dentata present. Maxillipcdal palp of single broad article; small tri 

angular endite present. Pereopod 1, propodus expanded. Pereopods 4-7, 

carpi short, triangular, lacking free anterior margin. Pleonites 1-5 very 

short, fused, 6 fused with telson. Pleopod 1, exopod operculiform. Telson 

basally broad, with two statocysts at about midlength. 6: Antennular ped 

uncle of four articles. Antennal flagellum of one article. Pereopod 1, pro 

podus with dense clump of spines on mesial surface. Eyes only slightly larger 

than in 9. 

REMARKS The genus comprises three species, two from the Caribbean and 

one from the Pacific, all of which have been taken from shallow coral reefs. 

Key to species of Pendanthura 

1. Dorsal pigmentation over entire body; pereopod 1, propodal palm with 

rounded lobe tanaiformis 

Dorsal pigmentation on cephalon, pereonite 2, and pleon; pereopod 1, 

propodal palm lacking rounded lobe hendleri 

Pendanthura hendleri Kensley, 1984 

Figure 24A-E 

DIAGNOSIS 9: 3.3 mm. Dorsal pigmentation limited to cephalon, pereonite 

2, and very short pleon. Reduced mandibular palp with two setae. Pereopod 

1, propodus expanded, palm gently convex, lacking rounded lobe, with four 

spines on mesial surface near palmar margin. 6: 2.8 mm. Pigmentation as in 

9. Pereopod 1, propodal palm gently convex, lacking rounded lobe. 

RECORDS Carrie Bow Cay, Belize, 9-23 m; Twin Cays, Belize, 0-2 m; Pan 

ama, 30 m. 

Pendanthura tanaiformis Menzies and Glynn, 1968 

Figure 24F-H 

DIAGNOSIS Ovigerous 9: 2.9 mm. Dorsal pigmentation a dense red-brown

Figure 24. Pendanthura hendleri: A, 2; B, mandible; C, maxilliped; D, pereopod 1, 

6\ Ey pereopod 1, 2. Pendanthura tanaiformis: Fy pereopod 1, 2; G, 2; H> mandible.

58 ANTHURIDEA • HYSSURIDAE 

reticulation over entire dorsum. Mandibular palp with one seta. Pereopod 1, 

propodus expanded, palm with rounded lobe, mesial surface with six spines 

near palmar margin. 6: 2.8 mm. Pigmentation as in 9. Propodal palm with 

rounded lobe. 

RECORDS Bermuda; Carrie Bow Cay, Belize, intertidal to 1 m; Puerto Rico, 

intertidal; Cozumel, Mexico. 

REMARKS Kensley (1984) characterized this common species of the interti 

dal of the reef crest as stress-tolerant, breeding throughout the year. 

Skuphonura Barnard, 1925 

DIAGNOSIS Antennal flagellum of single article. Cephalon with midventral 

toothlike process at base of mouthparts. Mandibular palp of three articles. 

Maxillipedal palp of three articles; endite lacking. Pereopod 1, propodus ex 

panded. Pereopods 4—7, carpi triangular. Pleopod 1, exopod operculiform. 

Pleonites 1—5 fused; pleonite 6 dorsally demarked. Pleotelson with two basal 

statocysts. 

Skuphonura laticeps Barnard, 1925 

Figure 25 

DIAGNOSIS 6 6.0 mm. Cephalon wider anteriorly than posteriorly, with 

anterolateral lobes extending well beyond rostrum. Antennular flagellum of 

two articles. Pereonite 1 with strong midventral forwardly directed tooth. 

Pereopod 1, carpus with posterodistal angle produced into triangular lobe; 

propodus expanded, palmar margin proximally convex, numerous setae on 

mesial surface. Pleopod 2, copulatory stylet of endopod reaching beyond 

rami. Pleonite 6 dorsally demarked, posterior margin middorsally incised. 

Uropodal exopod ovate, with distal notch; endopod length slightly more than 

basal width. Pleotelson widest at midlength, posteriorly narrowly rounded, 

with broadly rounded longitudinal raised area. 

RECORDS St. Thomas, U.S. Virgin Islands, 8-40 m. 

Family Hyssuridae Wagele, 1981 

DIAGNOSIS Maxillipedal palp usually of five articles; endite present. Per 

eopods 1-3 subsimilar, often all three pairs subchelate. Pleopod 1 similar to

Figure 25. Skuphonura laticeps: A, 6; B, cephalon and pereonite 

antennule, 8\ D, pereopod 1, 6; E, antenna; F, maxilliped.


following pleopods, not operculiform. Pleonites 1-5 freely articulating, rela 

tively elongate. Pleotclson lacking statocysts. 

Key to genera of Hyssuridae 

1. Pleotelson not covered by uropodal exopods; maxillipedal rami basally 

free Kupellonura 

Pleotelson covered completely by uropodal exopods; maxillipedal rami 

basally fused Xenanthura 

Kupellonura Barnard, 1925 

DIAGNOSIS Mandibular palp of three articles. Maxillipedal palp of five arti 

cles; large endite present. Pereopods 1-3 similar, propodi somewhat ex 

panded. Pereopods 4-7, carpi triangular. Pleonites 1—5 elongate, free, sube- 

qual. Pleopods 1—5 similar. 

Kupellonura imswe (Kensley, 1982) 

Figures 26, 27 

DIAGNOSIS 9: 3.4 mm. Eyes present. Antennular flagellum of four articles. 

Antennal flagellum of seven articles. Maxillipedal endite reaching to base of 

article 3. Pereopods 1-3 similar; pereopod 1, carpus posterodistally acute; 

propodus expanded, palm straight, unarmed; carpus triangular, free anterior 

margin shorter than posterior. Pleonite 6 with middorsal incision in posterior 

margin. Uropodal exopod, outer margin serrate, distally narrowly rounded; 

endopod length twice greatest width. Telson widest at midlength, 

posterolateral margin serrate, apically broadly rounded. 6: 2.7 mm. Eyes 

larger than in 9. Mouthparts reduced. Pereopod 1, propodal palm armed 

with row of eight fringed spines. 

RECORDS Carrie Bow Cay, Twin Cays, Glover's Reef, Belize, 0-6.0 m; 

Montego Bay, Jamaica, 1 m. 

Xenanthura Barnard, 1925 

DIAGNOSIS Mandibular palp of single article. Maxillipedal rami fused; ar 

ticulations of palp articles obscure. Pereopods 1—3 similar. Pereopods 4-7, 

carpi triangular.

Figure 26. Kupellonura imswe: A, 9; B, cephalon and pereonite 1, 6; C, maxilliped; 

Dy pereopod 1, 9; E, pereopod 1, 6.


Figure 27. Kupellonura imswe: A phalon, dorsal view; B, posterior pleon, d 

view; C, pleon, ventral view; D> cephalon and pereonites 1 and 2, lateral view 

Xenanthura brevitelson Barnard, 1925 

Figure 28 

D I A G N O S I S 9 : 4 . 0 m m . O m m a t i d i a o f e y e s i n l o n g i t u d i n a l r o w o f t h r e e o r 

f o u r g r o u p s . A n t e n n u l a r flagellum o f t h r e e a r t i c l e s . A n t e n n a l f l a g e l l u m o f 

t h r e e a r t i c l e s . M a n d i b u l a r p a l p o f s i n g l e s h o r t a r t i c l e b e a r i n g s i n g l e s e t a . 

M a x i l l i p e d a l r a m i b a s a l l y f u s e d , p a l p o f t h r e e o r f o u r o b s c u r e l y s e p a r a t e d 

a r t i c l e s . P e r e o p o d s 1 - 3 s i m i l a r , c a r p u s o f p e r e o p o d 2 t r i a n g u l a r , w i t h s t r o n g 

t r i a n g u l a r p r o j e c t i n g l o b e p o s t e r o d i s t a l l y ; p r o p o d i e x p a n d e d , p a l m i n c i s e d 

i n t o s e v e r a l r o u n d e d l o b e s . U r o p o d a l e x o p o d s c i r c u l a r , o v e r l a p p i n g d o r s a l l y 

a n d c o v e r i n g t e l s o n ; e n d o p o d p r o j e c t i n g b e y o n d e x o p o d s v e n t r a l l y , m e s i a l 

m a r g i n w i t h s t e p , d i s t a l l y r o u n d e d . T e l s o n s h o r t e r t h a n u r o p o d a l e x o p o d , 

p o s t e r i o r m a r g i n t r u n c a t e t o f a i n t l y c o n c a v e . 6 : 3 . 5 m m . A n t e n n u l a r 

flagellum o f s e v e n a r t i c l e s . 

R E C O R D S O f f G e o r g i a , 2 0 - 1 4 5 m ; o f f F l o r i d a , 8 - 1 0 m ; T u r k s a n d C a i c o s 

I s l a n d m ; S t . T h o m a s , U . S . V i r g i n I s l a n d s , 5 0 - 6 0 m ; G u l f o f M

Figure 28. Xenanthura brevitelson: A, 9;B, pereopod 1; C, uropodal endopod 

telson; E, uropodal exopod.

64 ANTHURIDEA • PARANTHURIDAE 

Family Paranthuridae Menzies and Glynn, 1968 

DIAGNOSIS Mouthparts together forming somewhat elongate cone adapted 

for piercing and sucking. Mandible with styliform incisor, lacking lamina 

dentata and molar. Maxilla slender, styliform, distally serrate. Maxilliped 

elongate; number of palp articles usually reduced. Pereopod 1, or pereopods 

1-3 subchelate. Pleopod 1 exopod operculiform. Pleonites short, free or 

Key to genera of Paranthuridae 

1. Pereopod 7 lacking in adult 2 

Pereopod 7 present in adult 3 

2. Eyes present Colanthura 

Eyes absent Curassanthura 

3. Antennular and antennal flagellum of more than 10 

articles Accalathura 

Antennular and antennal flagella with fewer than 10 articles 4 

4. Antennal flagellum a single (rarely two or 3) flattened article; 

maxillipedal palp of one or two articles Paranthura 

Antennal flagellum of seven articles; maxillipedal palp of three articles 

Accalathura Barnard, 1925 

Virganthura 

DIAGNOSIS Eyes present. Antennular and antennal flagella multiarticulate, 

each of more than 10 articles. Mandibular palp of three articles. Maxillipedal 

Key to species of Accalathura 

1. Uropodal exopod elongate-narrow; endopod length twice basal width 

crenulata 

Uropodal exopod ovate, apically subacute; endopod length 1.5 times 

basal width setosa

Colanthura tenuis 65 

palp of two articles, endite almost reaching end of palp. Pereopod 1 subche- 

late, propodus inflated, larger than pereopods 2 and 3. Pereopods 4-7, carpi 

with anterior and posterior margin subequal. Pleonites free, short. Pleopod 1, 

exopod operculiform. Telson with single statocyst. 

Accalathura crenulata (Richardson, 1901) 

Figure 29A-D 

DIAGNOSIS 9: 16.0 mm. Antennular flagellum of about 26 articles. Anten 

nal flagellum of about 18 articles. Uropodal exopod narrow, parallel sided. 

Telson apically subacute. 6: 15.0 mm. Pleopod 2, copulatory stylet of endo- 

pod apically acute, with subapical "heel." 

RECORDS OfFNorth Carolina, 30 m; off Georgia, 20 m; Cuba; Puerto Rico, 

intertidal; Cozumel, Mexico; Carrie Bow Cay, Twin Cays, Belize, intertidal 

to 6 m; west coast of Florida, Gulf of Mexico, 55 m. 

Accalathura setosa Kensley, 1984 

Figure 29E-H 

DIAGNOSIS 9: 8.5 mm. Antennular flagellum of 11 articles. Antennal 

flagellum of 13 articles. Uropodal rami, margins closely setose; exopod ovate, 

outer margin sinuate, apex subacute; endopod ovate, length 1.25 times great 

est width. Telson apically rounded. 6: 7.0 mm. Pleopod 2, copulatory stylet 

of endopod apically strongly bifid. 

RECORDS Carrie Bow Cay, Belize, intertidal to 0.5 m. 

Colanthura Richardson, 1902 

DIAGNOSIS Integument with minute squamae. Mandible lacking palp. 

Maxillipedal palp articles fused except for minute terminal article. Pereopod 

1 subchelate, propodus expanded. Pereopods 2 and 3 subchelate but smaller 

than pereopod 1. Pereopods 4-6, carpi rectangular. Pereonite 7 very short, 

pereopod 7 lacking. Pleotelson lacking statocyst. 

Colanthura tenuis Richardson, 1902 

Figure 30A-C 

DIAGNOSIS 9: 3.5 mm. Eyes present. Integument diffusely brown in color. 

Antennular flagellum of four articles. Antennal flagellum of single article.

Figure 29. Accalathura crenulata: A, Q,B, uropodal sympod and endopod; C, 

uropodal exopod; D, telson. Accalathura setosa: E, 9; F, uropodal sympod and 

endopod; G, uropodal exopod; H, telson.

Curassanthura bermudensis 67 

Pereopod 1, propodus with mesial surface bearing proximal row of six spines. 

Pleonites 1—5 short, fused, boundaries marked dorsally by folds. Telson pos 

teriorly broadly rounded. 6: 3.5 mm. Antennular flagellum of five articles. 

RECORDS Bermuda, intertidal to 0.5 m. 

Curassanthura Kensley, 1981 

DIAGNOSIS Eyes lacking. Mandibular palp of three articles. Maxillipedal 

palp of five articles; short endite present. Pereopod 1 subchelate. Pereopods 

2-6 similar, carpi rectangular. Pereopod 7 lacking. Pleonites 1-5 free; 

pleonite 6 dorsally demarked. Pleopod 1, exopod operculiform. Pleotelson 

with single statocyst. Interstitial littoral forms. 

REMARKS Three species of this interstitial genus are known, two from the 

Caribbean, and one from the upper sublittoral gravels of a lava tunnel on 

Lanzarote, Canary Islands. 

Key to species of Curassanthura 

1. Telson with posterior fourth abruptly narrowed; uropodal exopod 

length 4.5 times greatest width bermudensis 

Telson tapering, but posterior fourth not abruptly narrowed; uropodal 

exopod length 2.5 times greatest width halma 

Curassanthura bermudensis Wagele, 1985 

Figure 30G,H 

DIAGNOSIS 9 3.0 mm. Pereopod 1 propodal length 2.5 times proximal 

width, palm with proximal strongly recurved hooklike tooth. Uropodal ex 

opod slender, parallel sided, 4.5 times longer than greatest width. Telson 

constricted in posterior fourth. 

RECORDS Church Cave, Bermuda, in shore sediments.

68 ANTHURIDEA • PARANTHURIDAE 

Figure 30. Colanthura tenuis: A, 9; B3 uropod; C, pereopod 1. Curassanthura halma: 

D, 9; E, telson; F} uropodal exopod. Curassanthura bermudensis: G3 telson; H3 

uropodal exopod (from Wagele, 1985). 

Curassanthura halma Kensley, 1981 

Figure 30D-F 

DIAGNOSIS 9 2.3 mm. Pereopod 1, propodal length about 1.7 times proxi 

mal width, palm with eight fringed spines and basal tridentate lobe. Uropo-

Paranthura antillensis 69 

dal exopod triangular, length 2.5 times greatest width, shorter than sympod. 

Telson tapering but not abruptly narrowed in posterior fourth. 

RECORDS Curagao, in shore sediments 1.5 m above tide line; Bonaire, in 

shore sediments above tide line. 

Paranthura Bate and Westwood, 1868 

DIAGNOSIS Eyes present. Antennular flagellum shorter than peduncle. An- 

tennal flagellum usually of single flattened setose article. Mandibular palp of 

three articles, article 3 with comb of spines. Maxillipedal palp of one or two 

articles; endite small to absent. Pereopod 1, propodus inflated, larger than 

that of pereopods 2 and 3. Pereopods 4-7, carpi rectangular. Pleonites short, 

more or less distinct. Pleopod 1, exopod operculiform. Telson lacking 

statocyst. 

REMARKS This is the largest of the paranthurid genera, with over 50 names 

in the literature. Many of these are poorly described. Species of Paranthura are 

common in the shallow waters of the temperate and tropical seas. 

Key to species of Paranthura 

1. Telson posteriorly truncate; uropodal exopod rectangular, margins 

serrate infundibulata 

Telson posteriorly rounded; uropodal exopox ovate, margins entire . . 2 

2. Uropodal endopod longer than wide 3 

Uropodal endopod as long as wide antillensis 

3. Uropodal exopod elongate-elliptical; telson parallel sided for half length 

jloridensis 

Uropodal exopod ovate, outer margin sinuate; telson evenly elliptical 

Paranthura antillensis Barnard, 1925 

Figure 31A-F 

barnardi 

DIAGNOSIS 9 5.1 mm. Antennular flagellum of four articles. Mandibular 

palp, article 3 with five spines. Maxillipedal palp of single article three times 

longer than basal width; short endite present. Pereopod 1, propodus ex-

Figure 31. Paranthura antillensis: Ay 9; B, telson; Cy uropodal exopod; D, uropodal 

sympod and endopod; Ey pereopod 1, 9; F, pereopod 2, 9. Paranthura barnardi: Gy 

9; H, telson; /, pereopod 1, 9;Jy uropodal exopod; K, uropodal sympod and 

endopod.

Paanthura infundibulata 71 

panded, palm bearing row of setae, mesial surface near palmar margin with 

convex flange and row of about 10 spines. Pereopod 2, propodal palm with 

five stout sensory setae. Pleonite 6 free, posterior margin bilobed. Uropodal 

exopod ovate, outer margin sinuous; endopod almost circular, as wide as 

long. Telson posteriorly rounded. 

RECORDS St. Johns, St. James, U.S. Virgin Islands, 32 m; Carrie Bow Cay, 

Belize, intertidal to 1.5 m. 

Paranthura barnardi Paul and Menzies, 1971 

Figure 31G-K 

DIAGNOSIS 9 6.0 mm. Mandibular palp article 3 with eight spines. Max- 

illipedal palp of single article, length four times basal width. Pereopod 1, 

propodal palm concave, with convex flange and row of about 17 spines on 

mesial surface. Uropodal exopod broadly ovate, apically subacute; endopod 

ovate, length about 1.5 times basal width. Telson evenly elliptical, apex 

evenly rounded. 


Paranthura Jloridensis Menzies and Kruczynski, 1983 

Figure 32A-E 

DIAGNOSIS 9 6.3 mm. Integument with sparse irregular pigmentation. 

Mandibular palp article 3 with eight spines. Maxillipedal palp of single arti 

cle, length 3.5 times basal width. Pereopod 1, propodal palm with trans 

parent flange and row of 10 setae on mesial surface. Uropodal exopod 

elongate-elliptical; endopod ovate, length 1.5 times greatest width. Telson 

posteriorly broadly rounded, parallel sided for about half its length. 

RECORDS OfTSanibel Island, Florida, Gulf of Mexico, 73 m. 

Paranthura infundibulata Richardson, 1902 

Figure 32F-J 

DIAGNOSIS 9 8.2 mm, 6 6.0 mm. Integument with red-brown pigmenta 

tion; broad irregular patch between eyes running onto bases of antennules; 

pereonites 1 and 2 with anterior patches, remainder of pereonites with pos 

terior patches; strong patch on uropodal exopod, endopod, and telson. Man 

dibular palp article 3 with 11 or 12 spines. Maxillipedal palp of single article,

Figure 32. Paranthura Jloridensis: A, 9; B, pereopod 1, 9; C, uropodal sympod and 

endopod; D, uropodal exopod; Ey telson. Paranthura infundibulata: F> 9; G, telson; 

Hy pereopod 1, 9; /, uropodal exopod; J, uropodal sympod and endopod.

ASELLOTA 73 

length 2.5 times basal width. Pereopod 1, propodus with convex flange and 

row of more than 20 setae. Uropodal exopod elongate-rectangular, mesial 

and distal margins serrate; endopod roughly square, margins serrate. Telson 

parallel sided, posterior margin truncate. 

RECORDS Bermuda, 11-12 m; Carrie Bow Cay, Belize, intertidal to 1 m; 

Cozumel, Mexico; Venezuela. 

Virganthura Kensley, 1987b 


palp of three articles; endite present. Pereopods 1-3 subchelate, pereopod 1 

larger than pereopods 2 and 3; pereopods 4-7, carpi rectangular. Pleopod 1, 

exopod operculiform. Pleonites 1-5 short, distinct; pleonite 6 dorsally de- 

marked. Telson with single statocyst. 

Virganthura crassa (Barnard, 1925) 

Figure 33 


flagellum of seven articles. Maxillipedal endite reaching distal margin of first 

palp article. Pereopod 1, propodal palm slightly concave, bearing seven 

spines. Uropodal exopod ovate, outer margin sinuate; endopod distally 

rounded, articulating obliquely on sympod. 

RECORDS U.S. Virgin Islands, 30 m. 

Suborder Asellota Latreille, 1803 

DIAGNOSIS Antennules uniramous. Antennae uniramous, with scale in 

some families. Mandible usually with palp, but palp lacking in some groups. 

Pereopod 1 usually subchelate, sexually dimorphic in some groups. Coxae 

small, sometimes not all visible in dorsal view. Pleon seldom of more than 

two free pleonites plus pleotelson. Pleopod 1 absent in 9. One pair of 

pleopods in 9, and one or two pairs of pleopods in 6 forming operculum 

over remaining respiratory pleopods. Pleopod 2 in 6 usually adapted for 

copulation. Uropods usually pedunculate, but peduncle may be reduced, 

biramous or uniramous, terminal or subterminal. 

REMARKS The suborder Asellota is usually divided into four superfamilies, 

based on pleopodal arrangement. The great majority of families, however,

A 

Figure 33. Virganthura crassa: A, 9; B, maxilliped; C, uropodal endopod and 

sympod; D, telson; E, uropodal exopod. 

D

Atlantasellus cavernicolus 75 

belong to the superfamily Janiroidea, considered to be the most advanced. In 

place of a key to the four superfamilies, the chart shown in Figure 34 illus 

trates diagrammatically the arrangement of the pleopods in these groups. 

Superfamily Aselloidea Rafinesque-Schmaltz, 1815 

DIAGNOSIS 6: Pleopod 1, peduncles separate, rami uniarticulate; pleopod 2 

having biarticulate exopod and copulatory endopod; pleopod 3 biramous, 

opercular. 9: Pleopod 1 absent; pleopod 2 absent, or of single article; 

pleopod 3 biramous, opercular. 

Family Atlantasellidae Sket, 1979 

DIAGNOSIS Body resembling that of a sphaeromatid isopod. Pleon consist 

ing of two free pleonites plus pleotelson. 6: Pleopod 1 of two articles; pleopod 

2 with sympod, small exopod, and uniarticulate copulatory endopod. 9: 

Pleopods 1 and 2 absent; pleopod 3 operculate but rami not fused. 

Atlantasellus Sket, 1979 

DIAGNOSIS Eyes lacking. Antennule flattened, short and broad. Antenna 

elongate, peduncle of four (?five) articles, flagellum of four articles. Man 

dibular palp of three articles; with incisor and lacinia, molar replaced by 

brushlike process. Maxillipedal palp of five articles of similar width and 

broad endite. Pereopod 1 subchelate; pereopods 2-7 ambulatory, slender, 

dactyli biunguiculate. Uropods uniarticulate, vestigial. 

Atlantasellus cavernicolus Sket, 1979 

Figure 35 

DIAGNOSIS 6 1.1 mm. Cephalon with strong triangular rostrum, equal to 

pereonites 1-3 combined in middorsal length. Pleotelson basally broad, 

tapering to trilobed posterior margin, uropods inserted in incisions between 

median and lateral lobes. 

RECORDS Walsingham Cave, Bermuda.

PLEOPOD 1 

PLEOPOD 2 

PLEOPOD 3 

^ 

ASELLOIDEA 

cT 

n 

#1 

9 

or 

GNATHOSTENETROIDOIDEA 

V • * * * V 

1* p * * * 1 

V • * # • * 1 

\ • • • * 1 

v• * • *t 

v • * * 1 

F^ 

f 

d* 

* * * * • \ 

WJ 

09 

? 

J*-, * * * • * | 

VV-" ';:*:•'::••.••••.•.•:/ 

V • •. •,. . y 

1 '*•-"-*.*' 'J 

c. E> 

# 

« 

a 

X m 'i* ma i 

JANIROIDEA 

D> 

9 

/• V 

* • # • \ 

/ ' • ' / * " ' • " \ 

rl'- •-'•\ 

/ • " " ' • ••*• \ 

F * • • - p . - *\ 

/ . ' - • * • - " . \ 

• • • - • • » • .1 

I* - , * • • • » • • i 

'_.•.-.••. 1 

I -v-v.-.l 

Figure 34. Comparison of pleopods 1-3 in the four superfamilies of the suborder Asellota. 

« 

D> 

STENETRIOIDEA 

cr 

© 

w 

/-*- ••••'•J L>-.*: , A 

\.*•,•**•/ r-••# •••7 

V***'*l 1 1 J* * / 

/•;;;;••;>] 

9 

A 

I- • '^^**^Â 

'• •-Â 

JV - * I 

\ * • p 1 

V"'"-"-.- 

T • J 

J-.'•"•*:/ 

^^ * * * • J

Superfamily Gnathostenetroidoidea Kussakin, 1967 

Gnathostenetroid.es pugio 11 

DIAGNOSIS 6: Pleopod 1 peduncles fused, uniarticulate rami separate, op 

ercular; pleopod 2 small, with biarticulate exopod and copulatory endopod; 

pleopod 3, broad endopod biarticulate, exopod slender, uniarticulate. 9: 

Pleopod 1 absent; pleopod 2 rami fused to form operculum; pleopod 3 as in 

6. 

Family Gnathostenetroididae Kussakin, 1967 

DIAGNOSIS Uropods with short sympod, rami relatively well developed. 6: 

Pleopod 1, protopodites short, fused, rami separate, together forming oper 

culum covering remaining pleopods. Pleopod 2, separate, much smaller than 

pleopod 1. Pleopod 3, exopod elongate, slender; endopod uniarticulate, 

broad. 9: Pleopod 2, rami fused to form operculum covering remaining 

pleopods. 

Key to genera of Gnathostenetroididae 

1. Eyes absent. Rostrum anteriorly rounded/truncate Neostenetroides 

Eyes present. Rostrum anteriorly bilobed Gnathostenetroides 

Gnathostenetroides Amar, 1957 

DIAGNOSIS Eyes present. Rostrum well developed, anteriorly bilobed. Per- 

eonite 1, especially in

78 ASELLOTA • GNATHOSTENETROIDIDAE 

Figure 35. Atlantasellus cavernicolus: A, 9; B, 

pereopod 1. (From Sket, 1979). 

bearing row of nine more slender spines, posterior margin setose. 9: Oper 

culum with distal incision acute. 

RECORDS Florida Middlegrounds, Gulf of Mexico, 55 m. 

REMARKS The type of the genus, and only other species known, G. laodicense 

Amar, 1957, was recorded from the Mediterranean Sea. This species is in 

cluded here, as several of the species first recorded from the Florida Mid 

dlegrounds (Hooker, 1985) have since been recorded from the Caribbean. 

Neostenetroides Carpenter and Magniez, 1982 

DIAGNOSIS Pleonites 1 and 2 very short. Operculum of 9 subcircular. 

Pleopod 2 in 6 with elongate protopodite, copulatory organ prolonged by 

hyaline tonguelike structure. Pleopod 4, exopod ovate, wider than endopod. 

Neostenetroides stocki Carpenter and Magniez, 1982 

Figure 36D,E 

DIAGNOSIS 6 1.46 mm, 9 1.87 mm. Eyes absent. Rostrum well developed, 

anteriorly rounded/truncate. Pleotelson wider than long, lateral margins en 

tire, posterior margin regularly convex. Pereopod 1, propodus elongate, wid 

ening distally, palm poorly demarked, with few spines; dactylus stout, over 

lapping palm. Uropods unknown. 

RECORDS San Salvador, Bahamas, from Dixon Hill Lighthouse Cave.

ASELLOTA • JANIROIDEA 79 

Figure 36. Gnathostenetroides pugio: A, 8; B, mandible; C, pereopod 1. Neostenetroides 

stocki (from Carpenter and Magniez, 1982): Dy 9; E, pereopod 1. 

Superfamily Janiroidea Sars, 1899 

DIAGNOSIS 6: Pleopod 1 with elongate peduncle, occasionally fused; 

pleopod 2 with short exopod and copulatory usually elongate endopod, 

pleopods 1 and 2 together forming operculum; pleopod 3 endopod uniarticu-

80 ASELLOTA • JANIROIDEA 

late, exopod biarticulate. 9: Pleopod 1 absent; pleopod 2, rami fused to form 

operculum; pleopod 3 as in

Figure 37. Mexicope kensleyi: A, 9; B, 

pereopod 1. 

ASELLOTA • JANIRIDAE 81 

REMARKS The inability to place Mexicope in a family reflects the fact that 

the arrangement of the families and genera of the Janiroidea is still unsettled. 

While having affinities with the Pleurocopidae and the Janiridae, placement 

in either of these would require redefinition of both families, making the 

Janiridae even more of a hodgepodge of phylogenetically unrelated genera. 

Mexicope kensleyi Hooker, 1985 

Figure 37 

DIAGNOSIS 6 1.7 mm, 9 2.9 mm. Antennae slightly longer than body. Per- 

eonites and pleon laterally finely serrate. Pleotelsonic lobe between uropodal 

bases narrowly rounded, barely produced. 

RECORDS Turks and Caicos Islands, 1 m; Florida Middlegrounds, Gulf of 

Mexico, 30 m. 

Family Janiridae Sars, 1899 

DIAGNOSIS Antennae longer than antennules. Mand 

M 3 at least as wide as endite, 

markedly broader than articles 4 and 5. Pereopod 1 prehensile, subchelate, 

sexually dimorphic, larger in 6 than in 9. Pereopods 2 bulatory 

tyli biunguiculate. Coxae visible at least on three posterior pereonites. Pleon

82 ASELLOTA • JANIRIDAE 

consisting of single free pleonite (often very narrow, short and in- 

conspicuous) plus pleotelson. Uropods with well-developed sympod; usually 

biramous. 

Carpias Richardson, 1902 

DIAGNOSIS Cephalon with dorsolateral eyes, lacking rostrum. Coxae visible 

dorsally on pereonites 1-7. Pleon with one short pleonite lacking free lateral 

margins, plus broad pleotelson. Antennules and antennae well developed, 

latter with scale. Articles 2 and 3 of maxillipedal palp expanded. Pereopod 1 

sexually dimorphic, often relatively enormous and/or elongate in male, car- 

pochelate, carpus expanded, propodus variously armed or expanded, dac- 

Key to species of Carpias (based on pereopod 1 of mature 8) 

1. Propodus distally acute 2 

Propodus distally broad 4 

2. Propodus apically acute, with proximal tooth minutus 

Propodus lacking teeth 3 

3. Carpus with two strong distal teeth; propodus (when chela closed) 

reaching to merus algicola 

Carpus with two strong and one small teeth; propodus (when chela 

closed) reaching proximal half of carpus serricaudus 

4. Propodus with distinct teeth 5 

Propodus lacking distinct teeth 6 

5. Carpus distally with broadly rounded area; merus elongate-slender, 

length about four times greater than width bermudensis 

Carpus lacking broadly rounded area; merus short, broader than long 

punctatus 

6. Carpus with two distal teeth; dactylus minute triton 

Carpus with three distal teeth; dactylus small, but not obsolete 7 

7. Propodus distally truncate; carpus with middle tooth of palm distally 

faintly bilobed brachydactylus 

Propodus distally faintly bilobed; carpus with middle tooth of palm 

distally narrowly rounded harrietae

Carpias bermudensis 83 

tylus reduced or rudimentary, bearing two claws (biunguiculate). Pereopods 

2—7 similar, ambulatory, dactylus with three claws (triunguiculate). 

Uropods often longer than pleotelson, with relatively elongate sympod and 

rami. 

REMARKS This genus has been, and continues to be, a source of taxonomic 

problems. Several authors (e.g., Pires, 1980) have separated the species into 

the genera Carpias and Bagatus; others (e.g., Bowman and Morris, 1979) have 

synonymized them. In part, this uncertainty reflects the general uncertainty 

of the state of taxonomy in the family Janiridae. In this work, the genus 

Carpias is used to contain all the species described under the names Carpias 

and Bagatus. 

While these tiny asellotes are frequently extremely abundant in certain 

habitats (e.g., in reef-crest algal turfs; Kensley, 1983), difficulty is experi 

enced in identifying specimens other than mature males. The first pereopod 

of the mature male is the feature best used for species separation, but varia 

tion with maturity and geographic locality have not been investigated. With 

more detailed work, some species will undoubtedly be synonymized. 

Carpias aIgicola (Miller, 1941) 

Figure 38A,B 

DIAGNOSIS 8 2.9 mm, ovigerous 9 2.0 mm. Frontal margin straight. Per 

eopod 1 in

84 ASELLOTA • JANIR1DAE 

RECORDS Bermuda; eastern and southern coasts of Florida, 1.5-15 m. 

Carpias brachydactylus Pires, 1982 

Figure 38E 

DIAGNOSIS 6 1.6 mm. Pereopod 1 6, carpus distally between two and 

three times wider than proximal width, with strong triangular outer tooth 

defining palm, middle tooth apically faintly bifid, inner tooth rounded; pro 

podus widening to distal truncate margin, overreaching carpal palm by short 

distance, with very low tubercle at about midlength of flexor margin. 

Pleopod 1 

inner distal lobe rounded, outer lobe narrowly acute. Operculum of 9 with 

mediodistal margin gently concave. Uropod about twice length of pleotelson. 

RECORDS Biscayne Bay, Florida, intertidal to 2 m. 

Carpias minutus (Richardson, 1902) 

Figure 39A 

DIAGNOSIS 8 1.9 mm, ovigerous 9 1.8 mm. Pereopod 1

Carpias punctatus 85 

Figure 38. Carpias algicola: A, 9; B, pereopod 1,

86 ASELLOTA • JANIRIDAE 

Figure 39. Carpias minutus: A, pereopod 1, 6. Carpias punctatus: B, 6; C, pereopod 

1, 6. Carpias serricaudus: D, pereopod 1, d. Carpias triton: E, F, pereopod 1, 6. 

Pereopod 1 6, distal two-thirds parallel sided, with strong acute tooth defin 

ing palm, and second rounded tooth; propodus with three lobe-teeth on 

flexor surface, overreaching carpus by a third of its length. Pleopod 1

Carpias serricaudus Menzies and Glynn, 1968 

Figure 39D 

Joeropsis 87 

DIAGNOSIS 6 1.6 mm, 9 1.5 mm. Pereopod 1 d, palm of carpus with two 

strong outer teeth and one short inner tooth; propodus reaching back to 

proximal half of carpus, tapering distally; dactylus obsolete. Pleopod 1 d, 

outer distal lobe acute, reaching well beyond inner lobe. Pleotelsonic margins 

very faintly serrate. Uropod about 0.7 times length of pleotelson. 

RECORDS Turks and Caicos Islands, 1 m; Puerto Rico, intertidal to 1.5 m. 

Carpias triton Pires, 1982 

Figure 39E,F 

DIAGNOSIS 6 2.3 mm. Very similar to C. algicola. Pereopod 1 d, carpus 

with two strong basally broad distal teeth; propodus extending back to merus 

in adult 8y widening to broadly rounded distal margin; dactylus minute. 

Pleopod 1 d, outer distal lobe narrowly acute, reaching well beyond inner 

rounded lobe. Uropod about 1.5 times pleotelson length. 

RECORDS Carrie Bow Cay, Belize, intertidal reef crest. 

Family Joeropsidae Nordenstam, 1933 

DIAGNOSIS Cephalon free, with distinct rostrum. Molar process of mand 

ible reduced. Maxillipedal palp articles all of similar width. Antenna short, 

peduncle dilated, flagellum reduced. Pereonites similar, wider than long. 

Pereopods similar, biunguiculate. Uropods having short squat sympod and 

very reduced rami; inserted into submedian posterior notches of pleotelson. 

Joeropsis Koehler, 1885 

DIAGNOSIS Dorsolateral eyes present. Antennule, basal article widest and 

longest, often with transparent distal dentition. Antenna, peduncular articles 

3-5 somewhat dilated, article 2 often with fringe of transparent scales; 

flagellum of about six articles, together shorter than peduncle article 5. Per 

eonites similar, generally subequal in length and width. Pleotelson of single 

shield-shaped segment. Uropodal sympod usually with mesiodistal angle 

acute; rami reduced.

88 ASELLOTA • JOEROPSIDAE 

Key to species of Joeropsis 

1. Lateral margins of cephalon serrate; rostrum triangular personatus 

Lateral margins of cephalon entire; rostrum not triangular 2 

2. Body glabrous; strong band of pigment on cephalon and pereonite 4 

bifasciatus 

Body setose; pigment in reticulation over entire body 3 

3. Rostrum evenly convex; outer uropodal ramus longer than inner 

rathbunae 

Rostrum anteriorly shallowly notched; outer uropodal ramus shorter 

than inner coralicola 

Joeropsis bifasciatus Kensley, 1984 

Figure 40A-F 

DIAGNOSIS 6 2.5 mm, 9 2.4 mm. Body glabrous. Lateral margins of 

cephalon entire. Rostrum semicircular, with marginal flange of transparent 

teeth. Antennal flagellum of eight articles. Lateral margins of pleotelson ser 

rate. Apex of 9 operculum blunt. Broad band of pigment on cephalon be 

tween eyes and almost reaching posterior margin; broad band of pigment on 

pereonite 4. 

RECORDS Carrie Bow Gay, Belize, 1-6 m, often on Agaricia sp. and Porites 

sp. corals, and Halimeda sp. alga; Anguilla. 

Joeropsis coralicola Schultz and McCloskey, 1967 

Figure 40G 

DIAGNOSIS 6 2.0 mm, ovigerous 9 1.9 mm. Body setose. Lateral margins 

of cephalon entire. Rostrum anteriorly notched. Antennal flagellum of five 

articles. Lateral margins of pleotelson serrate. Apex of 9 operculum acute. 

Outer uropodal ramus shorter than inner. Pigment spread as reticulation 

over entire body. 

RECORDS Off North Carolina, on coral Oculina arbuscula; Florida Mid- 

dlegrounds, Gulf of Mexico, from sponge Agelas sp. and coral Madracis sp., 

25-33 m.

Figure 40. Joeropsis bifasciatus: A, 8; B, pleopod 1, 6; C, uropod; D, operculum, 9; 

E> antennule; F, antenna. Joeropsis coralicola: G, d. Joeropsis pe?sonatas: Hy 6. 

Joeropsis rathbunae: I, 6.

90 ASELLOTA • MICROPARASELLIDAE 

Joeropsis personatus Kensley, 1984 

Figure 40H 

DIAGNOSIS 6 2.2 mm, 9 2.0 mm. Body glabrous. Lateral margin of 

cephalon serrate. Rostrum triangular. Lateral margins of pleotelson serrate. 

Antcnnal flagellum of five articles. Apex of 9 operculum acute. Outer uropo- 

dal ramus shorter than inner. Strong band of pigment on cephalon; rest of 

body with paler reticulation of pigment. 

RECORDS Carrie Bow Cay, Belize, on Pontes sp. and Madracis sp. corals, 

and on Halimeda sp. alga, 1—20 m. 

Joeropsis rathbunae Richardson, 1902 

Figure 401 

DIAGNOSIS 6 1.9 mm, ovigerous 9 1.6 mm. Body setose overall. Lateral 

margins of cephalon entire. Rostrum evenly convex with flange of trans 

parent teeth. Antennal flagellum of three articles. Lateral margins of 

pleotelson serrate. Apex of 9 operculum acute. Outer uropodal ramus longer 

than inner. Pigment in reticulation over entire body. 

RECORDS Bermuda; Florida Keys; Turks and Caicos Islands; Puerto Rico; 

Gulf of Mexico; intertidal to 36 m. 

Family Microparasellidae Karaman, 1933a 

DIAGNOSIS Eyes lacking. Antennule much shorter than antenna. Antenna 

with scale. Pereopods all similar, with biunguiculate dactyli. Pleon of one 

free pleonite plus pleotelson. Uropods with well-developed sympod and 

rami. 

Key to genera of Microparasellidae 

1. d, plcopod 1 narrow, not overlapping external part of pleopod 2; 

maxillipedal palp of five articles Microcharon 

6, pleopod 1 broad, almost completely covering pleopod 2; 

maxillipedal palp of four articles, terminal article ending in pointed 

process Angliera

Microcharon sabulum 91 

REMARKS All the members of this family are tiny (usually less than 2 mm), 

and most are interstitial in habit, being found in marine, brackish, and fresh 

water environments. 

Angliera Chappuis and Delamare Deboutteville, 1955 

DIAGNOSIS Mandibular palp with two proximal articles inflated, terminal 

article, slender hooklike, articles lacking setae and spines. Maxillipedal palp 

of four articles, articles 1 and 3 elongate, article 2 short, article 4 with termi 

nal acute process. Pleopod 1 in 6 forming broad lamella. 

Angliera psamathus Kensley, 1984 

Figure 41A-D 

DIAGNOSIS 6 1.0 mm, 9 1.0 mm. Maxillipedal endite with seven setae on 

distal margin. Posterior four pairs of pereopods with claw on dactylus dorsal 

to unguis. Uropodal endopod subequal in length to sympod. 

RECORDS Carrie Bow Cay, Belize, interstitial in intertidal sand bank. 

REMARKS Two other species of Angliera have been recorded from the Carib 

bean area: A. dubitans Stock, 1977, from Bonaire, and A. racovitzai Coineau 

and Botosaneanu, 1973, from Cuba. The reader should refer to the original 

descriptions to distinguish the species, as differences are extremely subtle. 

Microcharon Karaman, 1934 

DIAGNOSIS Mandibular palp of three articles, two distal articles bearing 

spines and/or setae. Maxillipedal palp of five articles, articles 1, 2 and 3 

expanded. c£, pleopod 1 narrow, elongate, not obscuring pleopod 2. 

REMARKS More than 20 species of Microcharon have been described world 

wide. The genus is unusual in that the species have been found in true ma 

rine environments, in brackish habitats such as wells, and inland in 

freshwater. 

Microcharon sabulum Kensley, 1984 

Figure 41E-H 

DIAGNOSIS 6 1.4 mm, 9 1.5 mm. Antennule of five articles. Inner ramus of 

maxilla 2 with pectinate spine. Pereopodal dactyli short, biunguiculate. En-

Figure 41. Angliera psamathus: A, 6; B, right, and part of left, mandible; 

maxilliped; Dy pleopod 1, 6. Microcharon sabulum: Ey 6\ Fy mandible; G, 

maxilliped; Hy pleopod 1, d.

Munna 93 

dopod of pleopod 3 with three distal plumose setae. Uropodal sympod stout, 

longer than rami. 

RECORDS Carrie Bow Gay, Belize, interstitial in intertidal sand bank. 

REMARKS Two other species of Microcharon have been described from the 

Caribbean area: M. phreaticus Coineau and Botosaneanu, 1973, from intersti 

tial freshwater on Cuba, and M. herrerai Stock, 1977, from freshwater wells on 

Bonaire. The reader should refer to the original descriptions to distinguish 

these species. 

Family Munnidae Sars, 1899 

DIAGNOSIS Body ovate. Cephalon and all pereonites free; pleon narrower 

than rest of body, longer than broad. Eyes on lateral processes of cephalon. 

Mandible with molar and incisor present; palp present or absent. Maxillipe- 

dal palp articles 2 and 3 broader than remaining articles. Pereopod 1 prehen 

sile; pereopods 2—7 ambulatory. Uropods tiny, without sympod. Anus ex 

posed, not covered by pleopods. 

REMARKS Poore (1984) has provided the most useful and recent survey of 

the genera, and especially of Munna. 

Key to genera of Munnidae 

1. Mandibular palp present; pereopod 1 enormous in

94 ASELLOTA • MUNNIDAE 

Munna petronastes Kensley, 1984 

Figure 42A-D 

DIAGNOSIS

Uromunna reynoldsi 95 

Figure 42. Munna petronastes: A, 8\ B, uropod; C, pereopod 1, 8] D, pereopod 1 9 

(C and D same scale). Uromunna caribea: E, 6; F, larger uropodal ramus. Uromunna 

reynoldsi: G, pereopod 1, 8; Hy larger uropodal ramus; /, 8. 

Uromunna reynoldsi Frankenberg and Menzies, 1966 

Figure 42G-I 

DIAGNOSIS 6 1.5 mm, 9 1.6 mm. Propodus of pereopod 1 two or three 

times longer than wide. Operculum of 9 distally rounded. Shorter uropodal 

ramus obscured by pleotelsonic margin, with single seta. Pigmentation a

96 ASELLOTA • PARAMUNNIDAE 

broad patch between eyes on cephalon, lateral bands on pereon, and anterior 

and lateral patches on pleon. 

RECORDS Sapelo Island, Georgia, in tidal saltmarsh creek; Lake 

Ponchartrain, Louisiana; Atlantic and Pacific locks of Panama Canal. 

Family Paramunnidae Vanhoffen, 1914 

DIAGNOSIS Body broad, ovate, often with laterally produced tergal or epi- 

meral plates. Cephalon recessed into pereonite 1. Eyes, if present, on lateral 

projections of cephalon. Antennule short, usually of six articles, with single 

terminal aesthetasc. Antenna never longer than body. Mandibular palp pres 

ent or absent. Pereopod 1 prehensile; pereopods 2-7 ambulatory. Pleopod 1 

6 distally sagittate. Uropods with sympod minute or absent; rami tiny. Anus 

covered by pleopods. 

Munnogonium George and Stromberg, 1968 

DIAGNOSIS Eyes present on short lateral processes of cephalon. Antennal 

peduncular scale present. Coxal plates visible on pereonites 2—7. 

Munnogonium wilsoni Hooker, 1985 

Figure 43A,B 

DIAGNOSIS 6 0.86 mm, 9 0.98 mm. Frontal margin of cephalon broadly 

rounded. Mandibular palp absent. Uropodal endopod twice length of ex- 

opod. Lateral margins of pleotelson to uropodal insertion serrate, posterior 

margin between uropodal insertions tapering to rounded apex. 

RECORDS Florida Middlegrounds, Gulf of Mexico, 55 m. 

Family Pleurocopidae Fresi and Schiecke, 1972 

DIAGNOSIS Cephalon broader than long. Eyes (or at least ocular peduncles) 

present. Mandible with or without palp; molar truncate. Maxillipedal palp 

articles narrow, less than half width of endite. At least coxae of pereonites 5- 

7 dorsally visible. Pereopods 2-7 uni- or biunguiculate. Pleopod 1 in 6 not 

sagittate. Uropod pedunculate, inserted laterally or slightly dorso- or ven 

trolateral^ on pleotelson; biramous, or with one ramus fused with sympod.

Pleurocope 97 

Figure 43. Munnogonium wilsoni: A, 9; B, pereopod 1,

98 ASELLOTA • SANTIIDAE 

Figure 44. Pleurocope jloridensis: adult in dorsal 

view. 

Pleurocope Jloridensis Hooker, 1985 

Figure 43C 44 

DIAGNOSIS 1.15 m m , $ 0 . 9 6 m m . B o d y o v a t e , t a p e r i n g p o s t e r i o r l y . I n 

t e g u m e n t v e r y finely t u b e r c u l a t e . M e s i o d i s t a l l o b e o n a n t e n n a l p e d u n c l e a i 

t i d e 3 b e a r i n g five d i s t a l s e t a e . P e r e o n l a c k i n g l o n g d o r s a l s e t a e . P e r e o p o d 

s u b c h e l a t e , b u t a l m o s t c a r p o c h e l a t e . P l e o n c o n s i s t i n g o f s i n g l e s e g m e n t , p o s 

t e r i o r l y n a r r o w l y t a p e r e d a n d p r o d u c e d . U r o p o d a l r a m i a s l o n g a s s y m p o d . 

RECORDS T u r k s a n d C a i c o s I s l a n d s , 1 m ; C a r r i e B o w C a y , B e l i z e , 3 - 1 0 m ; 

F l o r i d a M i d d l e g r o u n d s , G u l f o f M e x i c o , 5 5 m . 

F a m i l y S a n t i i d a e W i l s o n , 1 9 8 7 

D I A G N O S I S A n t e n n f l a g e l l u m w i t h a t m o s t , t h r e e a r t i 

s c a l e s o m e t i m e s p r e s e n t . P e r e o p o d 1 p r e h e n s i l e . P e r e o p o d s 

a n t e n n u l a r 

d a c t y l i 

b i u n g u i c u l a t e . C o x a e v i s i b l e a t l e a s t o n p e r e o n i 7. P l e o n c o n s i s t i n g o f 

s i n g l e s h o r t p l e o n i t e p l u s p l e o t e l s o n . U r o p o d s p e d u n c u l a t e , b i r a m o u s , i n s e r 

t e d d o r s a l l y o r l a t e r a l l y . ( O n e s p e c i e s o f Santia p o s s e s s e s a u n i r a m o u s 

u r o p o d . ) 

S a n t i a S i v e r t s e n a n d H o l t h u i s , 1 9 8 0 

D I A G N O S I S C e p h a l o n a b o u t t w i c e w i d e r t h a n l o n g . E y e s p r e s e n t . A n t e n n u 

l a r p e d u n c l e o f t h r e e a r t i c l e s . P e r e o n i t e s l a t e r a l l y r o u n d e d , s o m e t i m e s b e a r 

i n g s h o r t l a t e r a l s p i n e s .

Santia milleri (Menzies and Glynn, 1968) 

Figure 43F-H 

ASELLOTA • STENETRIIDAE 99 

DIAGNOSIS 6 and 9 1.0 mm. Eye on short lateral process of cephalon. 

Mandibular palp of three articles. Maxillipedal palp articles all of similar 

width. Pereopod 1 barely subchelate. Uropod with sympod well developed, 

rami prominent, well developed. 

RECORDS Carrie Bow Cay, Belize, intertidal to 30 m; Puerto Rico, 1.5 m; 

San Salvador, Bahamas, 6 m; Turks and Caicos Islands, 1 m; Anguilla; Ja 

maica; Cozumel, Mexico; Gulf of Mexico. 

Brazil, 1-6 m. 

Superfamily Stenetrioidea Hansen, 1905a 

DIAGNOSIS 6: Pleopod 1 small, peduncles fused, rami separate, uniarticu- 

late; pleopod 2 small, copulatory; pleopod 3 biramous, opercular. 9 : Pleopod 

1 absent; pleopod 2 rami and peduncle fused to form operculum; pleopod 3 

as in 6. 

Family Stenetriidae Hansen, 1905a 

DIAGNOSIS 6: Pereopod 1 frequently much bigger than in 9, with distinc 

tive lobes and teeth. Pleopods 1 and 2 reduced; pleopod 1 protopodite short, 

fused, rami separate. Pleopod 2, endopod elongate, flexed, exopod short. 

Pleopod 3 exopod basally broad, distally narrowed; endopod broad, Particu 

late. 9: Pleopod 2, rami fused, short, covering base of pleopod 3. Uropod 

with short sympod, rami relatively well developed, styliform, of single article. 

Key to genera of Stenetriidae 

1. Rostrum narrowly triangular, spikelike; two free, very short pleonites 

anterior to pleotelson Stenobermuda 

Rostrum short, basally broad, anteriorly truncate or broadly rounded; 

two or three very short free pleonites anterior to pleotelson 

Stenetrium

100 ASELLOTA • STENETRI1DAE 

Stenetrium Haswell, 1881 

DIAGNOSIS Eyes present. Cephalon broader than long. Rostrum short, 

basally broad, anteriorly truncate or rounded. Pereonites 1—4 with ante 

rolateral projections; pereonites 5-7 projecting posteriorly. Pleotelson with 

sharp tooth anterior to small lateral notch. 

REMARKS If fresh material is not available, and color pattern is lost in pres 

ervation, mature male material is needed as identification is based on the 

structure of male pereopod 1. 

Key to species of Stenetrium 

1. Eyes of few (not more than 10) ommatidia, not reniform 2 

Eyes of many ommatidia, reniform 3 

2. Eyes of four ommatidia; 6 pereopod 1 small, propodus not unusually 

broad minocule 

Eyes of more than four ommatidia; 6 pereopod 1, propodus broad, 

with wide palm patulipalma 

3. Pleotelsonic margins serrate 4 

Pleotelsonic margins entire 5 

4. Rostrum convex, with fine marginal teeth; pereopod 1 c?, propodal 

palm with three straight teeth bowmani 

Rostrum truncate; pereopod 1 cJ, propodal palm with two teeth, outer 

tooth elongate, curved serratum 

5. Pereopod 1 cJ, carpus produced, apically acute stebbingi 

Pereopod 1 cJ, carpus produced, apically rounded spathulicarpus 

Stenetrium bowmani Kensley, 1984 

Figure 45 

DIAGNOSIS 8 5.0 mm, 9 5.2 mm. Rostrum convex, with tiny marginal 

teeth. Lateral lobes of cephalon acute, margins serrate. Color pattern in 

small scattered red-brown chromatophores; irregular unpigmented patches 

on cephalon. pereonite 4, and pleon; chalky-white bands on antennae and 

uropods. 8: Pereopod 1 propodus broad, palm with three teeth, outermost

Stenetrium bowmani 101 

Figure 45. Stenetrium bowmani: A, 6; B, pereopod 1, 6; C, pereopod 1, 9; D, 

pleopod \,6;E, pleopod 2, 6; F, pleopod 3; G, pleopod 4; H, operculum, 9.

102 ASELLOTA • STENETRIIDAE 

longest, slender. 9: Pereopod 1 propodus with strong denticulate spine de- 

marking palm, latter straight, with row of about seven slender spines. 

RECORDS Cozumel, Mexico; Carrie Bow Cay, Belize, 0.5-15.2 m, on algae 

and corals in reefcrest, and spur and groove zone of reef. 

Stenetrium minocule Menzies and Glynn, 1968 

Figure 46A-C 

DIAGNOSIS 8 2.8 mm, 9 3.7 mm. Eye of four ommatidia. Anterolateral 

lobes of cephalon blunt, barely produced. Rostrum poorly defined, truncate. 

8: Pereopod 1, carpus produced posterodistally into broadly rounded lobe; 

propodus broad, palm demarked by strong spine, with six low rounded teeth. 

9: Pereopod 1 propodus little broadened, palm demarked by strong denticu 

late spine, bearing several more spines. 

RECORDS Puerto Rico, intertidal to 3 m; Carrie Bow Cay, Belize, intertidal 

to 36 m, from rubble, algal turfs, and seagrass. 

Stenetrium patulipalma Kensley, 1984 

Figure 46D,E 

DIAGNOSIS 8 2.0 mm, 9 2.7 mm. Eyes of about 10 ommatidia in cluster. 

Rostrum poorly defined, truncate. Two basal articles of maxillipedal palp 

not enlarged. 8: Pereopod 1 unknown. 9: Pereopod 1 broadening to palm, 

and bearing row of about 12 small fringed spines. Color pattern: entire body 

with red-brown reticulation, dark transverse bars anteriorly on cephalon, 

pereonites 2 and 3, posteriorly on pereonites 4-7. 

RECORDS Carrie Bow Cay, Belize, 9.1—27.4 m; Barbados; Jamaica. 

Stenetrium serratum Hansen, 1904 

Figure 46F-H 

DIAGNOSIS 8 4.0 mm, ovigerous 9 4.9 mm. Rostrum truncate. Pereonites 

1-5 with acute anterolateral angles. Pleotelsonic lateral margins with five 

teeth. 8: Pereopod 1, propodus broad, palm with three teeth, outermost 

elongate, curved; dactylus reaching well beyond outermost palmar tooth. 9: 

Pereopod 1, propodus much smaller than in 8, palm bearing series of about 

nine fringed spines. Color: tiny red-brown chromatophores arranged in re-

Figure 46. tart. —«* *. * A P«-P od ' * C " "TT" ' '' ^TTH 

pLpalma: D,2;E, pereopod 1 «. ta*te —to.' F, 3; C, pereopod 1 9, H, 

pereopod 1 6.

104 ASELLOTA • STENETRUDAE 

ticulatc bands; distinctive open patches on cephalon and pereonite 1; pleon 

with two broad transverse bands. 

RECORDS Looe Key, Florida, 0.5-6 m; Turks and Caicos Islands, 1 m; 

Jamaica; Puerto Rico, intertidal to 3 m; St. Thomas, U.S. Virgin Islands; 

Carrie Bow Cay, Belize, intertidal to 15 m. 

Stenetrium spathulicarpus Kensley, 1984 

Figure 47A-C 

DIAGNOSIS 6 4.1 mm, 9 4.1 mm. Rostrum truncate. 6: Pereopod 1, merus 

and ischium each with setose fingerlike anterodistal projection; carpus with 

large spatulate and setose lobe almost reaching level of palm; propodal palm 

with large outer tooth and four or five low rounded teeth, broad band of setae 

near anterior margin; dactylus reaching slightly beyond palm, with band of 

setae along anterior margin. 9: Propodal palm straight, with row of slender 

spines; band of setae in similar position as in 6. Color: pigment in ill-defined 

and scattered reticulation; strong band on cephalon between eyes. 

RECORDS Carrie Bow Cay, Belize, intertidal to 36 m; Puerto Rico, 

intertidal. 

Stenetrium stebbingi Richardson, 1902 

Figure 47 D-H 

DIAGNOSIS 6 4.8 mm, ovigerous 9 4.1 mm. Rostrum truncate. Ommatidia 

of eye more bunched than in S. spathulicarpus. 6: Pereopod 1 variable accord 

ing to maturity; carpus produced posterodistally into narrowly triangular, 

apically acute lobe; propodal palm poorly defined, with group of two to four 

teeth near dactylar articulation. 9: Pereopod 1, palm straight, defined by 

strong outer tooth and bearing row of six or seven low rounded tubercles. 

Color: irregular reticulation of red-brown pigment, no strong band between 

eyes. 

RECORDS Bermuda, 0.5-4 m; Florida Keys, 18.3 m; Bahamas, 5 m; Turks 

and Caicos Islands, 1 m; Cuba; Jamaica; U.S. Virgin Islands, 50 m; Carrie 

Bow Cay, Belize, 0.5-36 m; Gulf of Mexico.

Figure 47. Sterutrium sPathulicarPus: A,6;B, pereopod 1, 6, many setae removed; 

C pereopod 1,9. Stenetnum stebbingi: D, 8; E, pereopod 1, 9; F, G, H, vanauon 

in pereopod 1,

106 ASELLOTA • STENETRIIDAE 

Figure 48. Stenobermuda acutirostrata: A, 9; By pereopod 1,9. 

Stenobermuda Schultz, 1979 

DIAGNOSIS Eyes of few ommatidia; rostrum narrow-based, elongate and 

spikelike. Pleon consisting of two free pleonites plus pleotelson, with 

posterolateral notch marked by tooth. 

Stenobermuda acutirostrata Schultz, 1979 

Figure 48 

DIAGNOSIS 6 4.8 mm. Eyes having five ommatidia. Spikelike rostrum 

reaching well beyond anterolateral angles of cephalon. Pereopod 1, propodus 

longer than wide, palm straight, bearing eight fringed spines, posterior mar 

gin with six spines and several setae. 

RECORDS Off Bermuda, 90 m; Turks and Caicos, 1 m.

Suborder Epicaridea Latreille, 1831 

EP1CARIDEA 107 

DIAGNOSIS Predominantly ectoparasites of marine crustaceans, feeding on 

blood. Eyes sessile, usually present in

Figure 49. A, epicaridium larva, lateral view; B> epicaridium larva, ventral view; 

C, cryptoniscium larva, ventral view (from Bonnier, 1900).

EP1GAR1DEA 109 

Figure 50. A, caridean shrimp with bopyrid parasite in branchial chamber. 

Probopyrus pandalicola: B, 9 and 6 in dorsal view, same scale; C, 6 enlarged; D, 9, 

ventral view, eggs removed from marsupium. 

cephalothorax of the host, attached dorsally to the carapace, ventrally and 

laterally in the gill chambers and on the pereopods, or in the brood 

chambers. 

The Entoniscidae are internal parasites of decapod crustaceans, being 

found in the visceral cavity, with the parasite's head in the position of the 

host's gonads or hepatopancreas. Veillet (1945) demonstrated that a pore to 

the host's branchial chamber connecting the parasite to the exterior is pres 

ent only in hosts with mature parasites, to facilitate the release of epi- 

caridium larvae. 

The Cryptoniscidae are protandrous hermaphrodites. The female is

110 

TABLE 2. CARIBBEAN EPICARIDEAN ISOPODS, THEIR HOSTS AND LOCALITIES 

Achelion occidentalis Hartnoll, 1966 

Microphrys bicornutus (Latreille) 

Stenorhynchus seticornis (Herbst) 

Jamaica 

Synalpheus pectiniger Coutiere 

North Carolina to Florida; Bahamas; 

Hispaniola; Jamaica; Bonaire; 

Curasao; Belize; Gulf of Mexico 

Aporobopyrina anomala Markham, 1973 Balanopleon tortuganus Markham, 1973 

Munida valida Smith 

Florida Keys; off Colombia; Gulf 

of Mexico 

Aporobopyrus curtatus (Richardson, 

1904) 

Petrochirus diogenes (Linnaeus) 

Petrolisthes armatus (Gibbes) 

Petrolisthes galathinus (Bosc) 

Petrolisthes marginatus Stimpson 

Porcellana sayana (Leach) 

Florida Keys; U.S. Virgin Islands; 

North Carolina 

Argeia atlantica Markham, 1977 

Sclerocrangon jacqueti (A. Milne 

Edwards) 

Bahamas; Newfoundland 

Astalione cruciaria Markham, 1975b 

Clastotoechus vanderhorsti (Schmitt) 

U.S. Virgin Islands 

Asymmetrione clibanarii Markham, 

1975d 

Clibanarius tricolor (Gibbes) 

Florida; Bahamas; Ascension 

Island 

Asymmetrione desultor Markham, 1975d 

Pagurus bonairensis Schmitt 

Pagurus longicarpus Say 

Pagurus provenzanoi Forest and de 

Saint Laurent 

Pylopagurus sp. 

North Carolina; Florida Keys; 

Curasao; Bonaire 

Azygopleon schmitti (Pearse, 1932) 

Synalpheus brooksi Coutiere 

Synalpheus hemphilli Coutiere 

Synalpheus longicarpus Coutiere 

Synalpheus mcclendoni Coutiere 

Munida simplex Benedict 

Tortuga Island 

Bopyrella harmopleon Bowman, 1956 

Synalpheus brevicarpus (Herrick) 

Synalpheus fritzmuelleri Coutiere 


Synalpheus minus Say 

Venezuela; Brazil 

Bopyrina abbreviata Richardson, 1904 

Hippolyte curacaoensis Schmitt 

Hippolyte pleuracanthus (Stimpson) 

Hippolyte zostericola (Smith) 

North Carolina to Florida; Belize; 

West Indies; Gulf of Mexico 

Bopyrinella thorii (Richardson, 1904) 

Thor Jloridanus Kingsley 

Florida; Curasao; Yucatan 

Peninsula, Mexico 

Bopyrione synalphei Bourdon and 

Markham, 1980 

Synalpheus goodei Coutiere 

Synalpheus bousfieldi Chace 

Synalpheus brevicarpus (Herrick) 


Florida; Haiti; Curasao; Gulf of 

Mexico 

Bopyrissa wolffi Markham, 1978 


Clibanarius vittatus (Bosc) 

Bermuda; North Carolina to Florida; 

Bahamas; Puerto Rico; Gulf of 

Mexico 

Cabirops sp. 

Synsynella deformans Hay 

Bermuda 

Cancricepon choprae (Nierstrasz and 

Brender a Brandis, 1925)

Domecia acanthophora (Desbonne 

and Schramm) 

Domecia hispida Eydoux and 

Souleyet 

Eriphia gonagra (Fabricius) 

Hexapanopeus angustifrons (Benedict 

and Rathbun) 

Micropanope barbadensis Rathbun 

Neopanope packardii (Kingsley) 

Neopanope texana sayi (Smith) 

Panopeus herbstii H. Milne Edwards 

Panoplax depressa Stimpson 

Paraliomera dispar (Stimpson) 

Rithropanopeus harrisii (Gould) 

Carolinas to Florida; Bermuda; 

Curagao; Gulf of Mexico 

Cancrion carolinas Pearse and Walker, 

1939 

Panopeus herbstii H. Milne Edwards 

North Carolina; Bahamas 

Dactylokepon caribaeus Markham, 

1975c 

Iliacantha liodactyla Rathbun 

Iliacantha subglobosa Stimpson 

Dominican Republic; Costa Rica- 

Panama 

Dicropleon periclimenis Markham, 1972a 

Periclimenes americanus Kingsley 

St. Lucia Island 

Diplophryxus sp. (see Markham, 1985) 

Alpheus formosus Gibbes 

Georgia; Florida; Yucatan, Mexico 

Eophrixus subcaudalis (Hay, 1917) 


Synalpheus goodei Coutiere 


Synalpheus longicarpus (Herrick) 

Synalpheus mcclendoni Coutiere 

Synalpheus pandionis Coutiere 


North Carolina to Florida; 

Yucatan Pensinsula, Mexico; 

Belize; Hispaniola; Curasao 

Gigantione mortenseni Adkison, 1984b 

Dromidia antillensis Stimpson 

Hypoconcha sabulosa (Herbst) 

Hypoconcha spinosissima Rathbun 

Florida; Haiti; Yucatan, Mexico; 

U.S. Virgin Islands; Gulf of 

Mexico 

Hemiarthrus synalphei (Pearse, 1950) 

Synalpheus fritzmuelleri Coutiere 



North Carolina to Florida; Haiti; 

Gulf of Mexico 

Leidya bimini Pearse, 1951 

Cyclograpsus interger (H. Milne 

Edwards) 

Pachygrapsus transversus (Gibbes) 

Sesarma ricordi H. Milne Edwards 

Bermuda; Florida Keys; Bahamas; 

U.S. Virgin Islands; Jamaica; 

Panama 

Leidya distorta (Leidy, 1855) 

Uca pugilator (Bosc) 

Uca spp. 

New Jersey to Florida; Guadeloupe; 

Trinidad 

Loki circumsaltanus Markham, 1972a 

Thor Jloridanus Kingsley 

Thor manningi Chace 

Southern Florida; U.S. Virgin 

Islands; Belize 

Metaphrixus carolii Nierstrasz and 

Brender a Brandis, 1931 

Hippolyte pleuracanthus Stimpson 

Southern Florida; U.S. Virgin 

Islands 

Munidion cubense Bourdon, 1972 

Munidajlinti Benedict 

Munida stimpsoni A. Milne Edwards 

Cuba; Venezuela 

Munidion irritans Boone, 1927 

Munida irrasa A. Milne Edwards 

Florida Keys; Belize 

111 

{continued)

12 

TABLE 2. {Continued) 

Munidion longipedis Markham, 1975a 

Munida longipes A. Milne Edwards 

Munida schroederi Chace 

East coast of Florida; Florida 

Keys; Cuba; Gulf of Mexico 

Parabopyrella lata (Nierstrasz and 

Brender a Brandis, 1929) 

Alpheus normanni Kingsley; 

Upogebia affinis (Say) 

Florida; U.S. Virgin Islands; 

Brazil 

Parabopyrella mortenseni (Nierstrasz 

and Brender a Brandis, 1929) 

Lysmata rathbunae Chace 

Lysmata wurdemanni (Gibbes) 

Florida; U.S. Virgin Islands; 

Venezuela 

Parabopyrella richardsonae (Nierstrasz 


Alpheus formosus Gibbes 

Alpheus heterochaelis (Say) 

U.S. Virgin Islands; Gulf of 

Mexico 

Parabopyrella thomasi (Nierstrasz and 


Tozeuma carolinense Kingsley 

U.S. Virgin Islands 

Parathelges piriformis Markham, 1972b 

Paguristes oxyophthalmus Holthuis 

Pagurus brevidactylus (Stimpson) 


Saint Laurent 

Bermuda; Bahamas; Colombia 

Parathelges tumidipes Markham, 1972b 

Allodardanus bredini Haig and 

Provenzano 

Dardanus fucosus Biffar and 

Provenzano 

Bermuda; Jamaica 

Pleurocrypta floridana Markham, 1974 

Galathea rostrata A. Milne Edwards 

Alligator Reef, Florida 

Pleurocryptella fimbriata Markham, 

1973 

Munida constricta A. Milne Edwards 

Munida miles A. Milne Edwards 

Western Caribbean; Cuba 

Probopyria alphei (Richardson, 1900a) 

Alpheus armillatus H. Milne Edwards 

Alpheus heterochaelis Say 

Alpheus normanni Kingsley 

Alpheus viridari (Armstrong) 

North Carolina to Florida; Antilles; 

Brazil; Gulf of Mexico 

Parapagurion imbricata Markham, 1978 Probopyrinella latreuticola (Gissler, 1882) 

Paguristes tortugae Schmitt 

Parapagurus sp. 

Cuba; Colombia 

Parathelges foliatus Markham, 1972b 

Clibanarius vittatus (Bosc) 


Barbados; Curasao; Trinidad 

Parathelges occidentalis Markham, 

1972b 


Iridopagurus sp. 

Pylopagurus corallinus (Benedict) 

North Carolina; Florida Keys; 

Bahamas; Venezuela 

Latreutes fucorum (Fabricius) 

Bermuda; Sargasso Sea to Azores; 

North Carolina to Florida; 

Bahamas; Antilles; Gulf of 

Mexico 

Probopyrus pandalicola (Packard, 1879) 

Macrobrachium acanthurus (Wiegmann 

Macrobrachium amazonicum (Heller) 

Macrobrachium bonelli (Nobili) 

Macrobrachium carcinus (Linnaeus) 

Macrobrachium faustinum (de Saussure 

Macrobrachium ohione (Smith) 

Macrobrachium olfersii (Wiegmann) 

Macrobrachium surinamicum Holthuis

Palaemon northropi (Rankin) 

Palaemon pandaliformis (Stimpson) 

Palaemonetes exilipes Stimpson 

Palaemonetes intermedins Holthuis 

Palaemonetes kadiakensis Rathbun 

Palaemonetes paludosus (Gibbes) 

Palaemonetes pugio Holthuis 

Palaemonetes vulgaris (Say) 

Periclimenes americanus (Kingsley) 

New Hampshire to Florida; 

Caribbean to Brazil; Pacific 

Panama 

Pseudasymmetrione sp. (see Adkison 

and Heard, 1978) 

Iridopagurus iris (A. Milne 

Edwards) 

Venezuela 

Pseudione affinis (Sars, 1882) 

Pandalus annulicornis Leach 

Pandalus bonnieri Caullery 

Pandalus leptorhynchus Kinahan 

Pandalus montagui Leach 

Plesionika antiguai Zariquiey 

Plesionika edwardsi (Brandt) 

Plesionika ensis (A. Milne Edwards) 

Plesionika heterocarpus (Costa) 

Plesionika martia (A. Milne 

Edwards) 

Bermuda; Northeastern Atlantic; 

South Africa; Java 

Schizobopyrina urocaridis (Richardson, 

1904) 

Periclimenes longicaudatus (Stimpson) 

Pontonia margarita Smith 

North Carolina to Florida; Belize; 


Stegias clibanarii Richardson, 1904 


Bermuda 

Stegophryxus hyptius Thompson, 1902 

Iridopagurus sp. 

Pagurus annulipes (Stimpson) 

Pagurus bonairensis Schmitt 


Pagurus longicarpus Say 


Saint Laurent 

Massachusetts to Florida; Curasao 

Synalpheion giardi Coutiere, 1908 

Synalpheus longicarpus Herrick 

Yucatan, Mexico 

Synsynella choprae (Pearse, 1932) 



Synalpheus minus (Say) 

Synalpheus pandionis Coutiere 

113 

Bermuda; North Carolina to Florida; 

Bahamas; Haiti; U.S. Virgin 

Islands; Gulf of Mexico 

Synsynella deformans Hay, 1917 




Bermuda; Carolinas to West Indies; 


Urobopyrus processae Richardson, 

1904 

Ambidexter symmetricus Manning and 

Chace 

Processa acutirostris Nouvel and 

Holthuis 

Processa canaliculata Leach 

Processa edulis (Risso) 

Processa Jimbriata Manning and Chace 

Processa tenuipes Manning and Chace 

Caribbean; Gulf of Mexico; Brazil; 

Mediterranean; Eastern Atlantic

1 14 FLABELLIFERA 

reduced to a simple or lobed sac, generally without appendages. The brood- 

pouch is formed by invagination of the ventral body wall. The eggs are re 

leased by the bursting of the sac. Cryptoniscids have been recorded as 

parasites of ostracods, cirripedes, mysidaceans, amphipods, isopods, and 

cumaceans. The majority feed on blood, but the females of some forms have 

been reported to be egg predators. 

Given the highly variable morphology of the epicarideans, and the neces 

sity of examining large series of specimens, keys are not provided and species 

are not treated individually here. As there is a degree of genus- and species- 

specificity for the hosts, Table 2 is provided to give a clue to the possible 

identity of a specimen. The student is then advised to consult one of the 

detailed works on the group. The most useful single work on the speciose 

Bopyridae for the area covered here is Markham (1985). 

Suborder Flabellifera Sars, 1882 

DIAGNOSIS Eyes usually well developed, reduced or absent in cave forms. 

Antennules and antennae uniramous; antennal peduncle of five or six arti 

cles. Mandible usually strong, adapted for cutting and grinding, occasionally 

for piercing; lacinia mobilis, spine-row, and molar usually present, although 

latter sometimes reduced; usually with triarticulate palp. Maxilla 1 

biramous, sometimes adapted for piercing; maxilla 2 biramous, outer ramus 

Key to families of Flabellifera 

1. Pleon consisting of four or five free pleonites plus pleotelson 3 

Pleon consisting of not more than three free pleonites plus 

pleotelson 2 

2. Pleon consisting of one or two free pleonites plus pleotelson; body 

usually dorsally strongly convex; pleopods subequal 

Sphaeromatidae 

Pleon consisting of three free pleonites plus pleotelson; body strongly 

depressed; pleopods 1—3 small, natatory, pleopods 4 and 5 large and 

broadly ovate Serolidae 

3. Uropodal rami flattened, generally not reduced 4 

Uropodal rami reduced, exopod often hooklike Limnoriidae

FLABELLIFERA • AEGIDAE 115 

4. Pereopods 4—7 prehensile, with dactyli longer than propodi; antennae 

reduced, with no clear distinction between peduncle and flagellum 

Cymothoidae 

Pereopods 4—7 ambulatory, with dactyli shorter than propodi; antennae 

normal, peduncle and flagellum clearly distinguished 5 

5. Maxilliped bearing distal recurved hooks; pereopods 1-3 strongly 

prehensile Aegidae 

Maxilliped lacking distal recurved hooks; pereopods 1-3 ambulatory or 

at most weakly prehensile 6 

6. Maxilliped lacking, or with very reduced endite; maxilla 1 a strongly 

falcate hook Corallanidae 

Maxilliped with strong endite; maxilla 1 not strongly falcate 7 

7. Mandibular incisor distally narrowed, lacinia lacking; maxilla 1 slender 

and elongate, with 3-5 distal hooked spines Tridentellidae ** 

Mandibular incisor distally broad, cusped; maxilla 1 relatively broad, 

with several distal spines and setae Cirolanidae { 

usually consisting of two lobes. Pereopods generally ambulatory, sometimes 

prehensile; pereopods 1 and 2 subchelate only in Serolidae, ancinine 

Sphaeromatidae, and some Cirolanidae; posterior pereopods sometimes sec 

ondarily natatory in some cirolanids. Pleon consisting of as many as five free 

pleonites plus pleotelson, but pleonites variously fused in several families. 

Five pairs of pleopods usually present, Uropods lateral, usually forming tail- 

fan with pleotelson. 

REMARKS This suborder contains a large group of diverse families, largely 

held together by primitive features such as the tailfan structure. Future work 

will undoubtedly show the Flabellifera to be an artificial polyphyletic group. 

Family Aegidae Leach, 1815 

DIAGNOSIS Dorsal integument usually unornamented. Coxae distinct on 

pereonites 2-7. Eyes usually present, large, often almost, or complete con 

tiguous. Mandible lacking lacinia mobilis, spine-row, and molar. Maxilla 1 

slender, with apical spines. Maxilla 2 with two terminal unequal lobes bear 

ing apical spines. Maxillipedal palp of two, three, or five articles. Pereopods 

1—3 prehensile, with dactyli strongly curved; pereopods 4—7 ambulatory.

116 FLABELLIFERA • AEGIDAE 

Pleopods biramous, bearing plumose marginal setae. Uropods forming tail- 

fan with plcotclson. Pleon of four or five free pleonitcs plus pleotelson. 

REMARKS Although these large isopods (up to 60 mm) are often referred to 

as fish parasites, Brusca (1983) prefers the term "carnivorous scavengers and 

micropredators," as they attach to fish hosts infrequently and only long 

enough to feed. When feeding, they engorge themselves on the host's blood. 

Aegids show almost no host- (or rather prey-) specificity, being opportunistic 

feeders, and are most frequently captured by bottom trawls on the ocean bed. 

In ovigerous females, the maxillipedal articles become expanded and, along 

with the anterior oostegites, cover the buccal field, thereby making feeding 

impossible. 

Key to genera and subgenera of Aegidae 

1. Maxillipedal palp of two or three articles; frontal lamina small, narrow 

Rocinela 

Maxillipedal palp of five articles; frontal lamina large, broad 2 

2. Antennular peduncle articles 1 and 2 expanded; cephalon lacking true 

rostrum, not completely separating antennular bases .... Aega (Aega) 

Antennular peduncle articles 1 and 2 not expanded; cephalon with true 

rostrum completely separating antennular bases . . . Aega (Rhamphion) 

Aega Leach, 1815 

DIAGNOSIS Eyes large, contiguous or separate. Cephalon with or without 

true rostrum. Frontal lamina broad, separating bases of antennae. Mandibu 

lar palp article 2 elongate. Maxilla 1 bearing strong apical and subapical 

spines. Maxilla 2 of two usually unequal lobes bearing stout spines. Max 

illipedal palp of four or five articles, terminal article often small, with setae or 

recurved spines; article 4 with stout recurved spines; endite small, seldom 

reaching beyond palp article 2. Pleon not much narrower than pereon. 

REMARKS Brusca (1983) published a useful account of the genus Aega in the 

Eastern Pacific.

1. Eyes contiguous 

Eyes separate . 

Key to species of Aega (Aega) 

Key to species of Aega (Rhamphion) 

Aega (A eg a) ecarinata 117 

1. Posterior margin of pleotelson distinctly dentate dentata 

Posterior margin of pleotelson at most faintly crenulate tenuipes 

Aega (Aega) deshaysiana (PL Milne Edwards, 1840) 

Figure 51A 

DIAGNOSIS 9 18.0 mm. Eyes large, contiguous. Cephalon with frontal mar 

gin acute to subacute. Frontal lamina large, shield shaped. Antennular ped 

uncle articles 1 and 2 not expanded; flagellum of more than 15 articles. 

Uropodal endopod with deep notch in lateral (outer) margin. Pleotelson with 

basal width subequal to middorsal length, triangular, lateral margins faintly 

to markedly convex, tapering to narrowly rounded to subacute apex. 

RECORDS Cuba; Yucatan Peninsula, Mexico; Gulf of Mexico. 

Azores; Cape Verde Islands; Tristan da Cunha; Mediterranean; St. Paul and 

Amsterdam Islands; Seychelles; east coast of South Africa; Philippines; Ja 

pan; Hawaii; northeast Australia; Tasmania; Cocos Islands; Costa Rica. 

REMARKS This species is more familiarly known in the Caribbean region as 

Aega antillensis Schioedte and Meinert, 1879. 

Aega (Aega) ecarinata Richardson, 1898 

Figure 51B 

deshaysiana 

ecarinata 

DIAGNOSIS 9 21.0 mm. Eyes well separated. Articles 1 and 2 of antennular 

peduncle expanded. Propodus of pereopod 3 with posterodistal lobe. Uropo

118 FLABELLIFERA • AEGIDAE 

Figure 51. A, Aega (Aega) deshayesiana; B, Aega (Aega) ecarinata; Cy Aega (Rhamphion) 

dentata; Dy Aega (Rhamphion) tenuipes. 

dal exopod narrower than endopod; latter distally truncate, lacking marginal 

notch. Pleotelson dorsally smooth, with posterior margin broadly trilobed. 

RECORDS Bahamas, 776 m; Puerto Rico; Gulf of Mexico, 176 m.

Aega (Rhamphion) dentata Schioedte and Meinert, 1879 

Figure 51C 

Rocinela 119 

DIAGNOSIS 9 7.5 mm. Eyes large, just contiguous in midline. Frontal lam 

ina distally acute. Uropodal exopod shorter than and half width of endopod; 

latter with lateral margin entire. Pleotelson with two obscure dorsal depres 

sions anteriorly; posterior margin crenulate with seven teeth. 

RECORDS Cuba. 

Aega (Rhamphion) tenuipes Schioedte and Meinert, 1879 

Figure 51D 

DIAGNOSIS 9 11.5 mm. Eyes large, contiguous. Frontal lamina distally 

broadly rounded. Uropodal exopod shorter and narrower than endopod; lat 

ter with entire lateral margin. Pleotelson dorsally smooth; posterior margin 

evenly and broadly convex, obscurely crenulate. 

RECORDS Cuba. 

Rocinela Leach, 1818 

DIAGNOSIS Cephalon with short rostrum sometimes covering antennular 

bases. Eyes well developed. Frontal lamina small, often indistinct. Mandibu 

lar palp of three articles, article 1 elongate. Maxillipedal palp of two or three 

articles. Pereopods 1—3 usually with spine-bearing expanded lobe on pos 

terior margin of propodi. 

Key to species of Rocinela 

1. Eyes contiguous oculata 

Eyes not contiguous 2 

2. Cephalon produced anteriorly into broadly rounded rostrum . . cubensis 

Cephalon lacking obvious broadly rounded rostrum 3 

3. Eyes well separated, cephalon anteriorly broadly triangular .... signata 

Eyes barely separate, cephalon anteriorly narrowly triangular insularis

120 FLABELLIFERA • AEG1DAE 

Rocinela cubensis Richardson, 1898 

Figure 52A 

DIAGNOSIS 6 16 mm. Cephalon with two small tubercles between well- 

separated eyes; rostrum broadly rounded, extending anteriorly, very ob 

vious. Flagellum of antenna with about 15 articles. Propodi of pereopods 1-3 

with two spines. Pleotelson basally wider than middorsal length, lateral mar 

gins convex, apex rounded. 

RECORDS Off Cuba, 290 m. 

Rocinela insularis Schioedte and Meinert, 1879 

Figure 52B 

DIAGNOSIS Ovigerous 9 24.5 mm. Eyes medially barely separated but not 

contiguous. Flagellum of antenna of more than 12 articles. Propodus of per 

eopods 1-3 with two to four spines on posterior lobe. Uropodal endopod 

barely reaching beyond pleotelsonic apex. Pleotelson basally slightly wider 

than middorsal length, lateral margins convex, apex rounded. 

RECORDS Florida Keys; West Indies; between Mississippi delta and west 

coast of Florida, Gulf of Mexico; 550 m. 

Rocinela oculata Harger, 1883 

Figure 52C 

DIAGNOSIS 9 21.0 mm. Eyes contiguous. Cephalon with rostrum truncate 

in dorsal view. Antennal flagellum with 12 articles. Propodi of pereopods 1-3 

with six to eight spines on lobed posterior margin. Pleotelson with basal 

width subequal to middorsal length; posterior margin broadly rounded. 

RECORDS Off Georgia; Gulf Stream off Florida, 360-400 m; Puerto Rico, 

84 m; Gulf of Mexico, 380-750 m. 

New South Wales, Queensland, Australia, 450-630 m. 

Rocinela signata Schioedte and Meinert, 1879 

Figure 52D 

DIAGNOSIS Ovigerous 9 13.0-15.0 mm. Cephalon anteriorly broadly tri 

angular, produced over bases of antennules. Eyes widely separate. Flagellum 

of antenna with 10 or 11 articles. Pereopods 1—3, propodi unarmed or with

c 

L 

r 

/ / 

* ^ 

- • * 

••rig 

b 

• ' 

1 ' * 1 

" b 1 " 

• +.• • 

• 

*^^; .'."7 

+ 

: : : 

1 • p 

w 

• 

b ' 

• - - fc l f c 

" ^ - " : ' • • - . . 

• • 

b 

• * • • * * 

- * ••'.- 

-*7^ 

b 

• 

+ 

bi 

+ 

._ b 

^^"•)b^ 

ik 

^fc ^bPÎ 

"M 

••"•"-.'IV'', 

Figure 52. ^4, Rocinela cubensis; B, Rocinela insularis; C, Rocinela oculata; D, Rocinela 

signata. 

D 

* • *' 

fc?l 

i ,r *• 

i^. 

*• + • 

* * * - 

'*••• 

+* X- * ** 

• 

« i 

+ *_+ 

*\ ^. 

* - 

• T 

+ 

> * . 

•- +

122 FLABELLIFERA • CIROLANIDAE 

single spine on posterior margin. Plcotelson with posterior margin evenly 

and broadly rounded; usually with inverted W-shaped band of pigment. 

RECORDS Florida Keys, shallow infratidal-4 m; Tortugas, from gills ofjew- 

fish Epinephelus itajara, mutton snapper Lutjanus analis; U.S. Virgin Islands, 

on mutton snapper Lutjanus analis, on yellowfin grouper Mycteroperca venenosa; 

Bahamas, on sheepshead Archosargus probatocephalus, on mutton snapper Lut 

janus analis, on blackfin snapper Lutjanus buccanella, on queen triggerfish Bat 

istes vetula, on saucereye porgy Calamus calamus; Jamaica, on French grunt 

Haemulon Jlavolineatum, hogfish Lachnolaimus maximus, on parrotfish Sparisoma 

viride; Haiti; Yucatan Peninsula, Mexico, 60—93 m, on gills of tiger shark 

Galeocerdo cuvieri; Puerto Rico, in gill slits of southern stingray Dasyatis amer- 

icana, in gill slits of nurse shark Ginglymostoma cirratum; Jamaica; Carrie Bow 

Cay and Blue Ground Range, Belize, 0.5—2 m, on jolthead porgy Calamus 

bajonado and sheepshead porgy Calamus penna, on peacock flounder Bothus lu- 

natus, on queen triggerfish Balistes vetula, on Caranx sp., on barracuda 

Sphyraena barracuda, on hogfish Lachnolaimus maximus, on mutton snapper Lut 

janus analis; Venezuela, on Orthopristis ruber, on Haemulon steindachneri; Sur 

inam, on gills of sheepshead porgy Calamus penna; Gulf of Mexico off Florida, 

shallow infratidal-55 m, on red grouper Epinephelus morio, on Lutjanus black- 

fordi, on black grouper Mycteroperca bonaci, on clearnose skate Raja eglanteria. 

Pacific records: Southern California and Gulf of California; Socorro Is 

land; Panama; Costa Rica. 

REMARKS While often taken from fish hosts (sometimes in the gill cham 

ber), this species is equally frequently found freeliving in shallow water over 

sand and coral rubble. The species will also attach itself to humans, inflicting 

a sharp bite as it tries to feed. 

Family Cirolanidae Dana, 1852 

DIAGNOSIS Eyes when present, relatively small, lateral. Frontal lamina 

present. Mandible with tridentate incisor, lacinia mobilis, blade- or sawlike 

molar, and palp. Maxillipedal palp of five articles, endite present. Coxal 

plates present on pereonites 2-7, distinctly separated by suture from tergite. 

Pereopods generally ambulatory, although anterior three pairs prehensile in 

some genera, and posterior four pairs natatory in some genera. Picon of five 

free pleonites plus pleotelson in most genera; pleonite 5 with free lateral mar 

gins or overlapped by pleonite 4. Pleopods membranous, lacking ridges or 

folds. Uropods situated at anterolateral angles of pleotelson, freely articulat 

ing, both rami well developed, mobile.

FLABELLIFERA • CIROLANIDAE 123 

REMARKS Of the many recent publications on the cirolanids, the most com 

prehensive is that of Bruce (1986) on the cirolanids of Australia. Botosa- 

neanu, Bruce, and Notenboom (1986) tabulate all the known troglobitic 

cirolanids of the world. 

Key to subfamilies of Cirolanidae 

1. Clypeus projecting; pleonite 5 with free lateral margins (except in 

Xylolana) Eurydicinae 

Clypeus flattened, not projecting; pleonite 5 lacking free lateral margin, 

overlapped by pleonite 4 2 

2. Pereopods 1-3 with ischium and merus not anterodistally produced; 

antennal peduncular articles 4 and 5 subequal; secondary unguis 

present on pereopodal dactyli Cirolaninae 

Pereopods 1—3 with ischium and merus anterodistally produced; 

antennal peducular articles 3 and 4 subequal; no secondary unguis 

on pereopodal dactyli Conilerinae 

Subfamily Cirolaninae Dana, 1852 

DIAGNOSIS Frontal lamina short, flat. Clypeus flattened, not projecting. 

Antennal peduncular articles 4 and 5 subequal, longer than articles 1-3. 

Pereopods with secondary unguis on dactyli. Penes reduced or absent. 

Pleonite 5 always overlapped by pleonite 4. Pleopod 2 in


Key to genera of Cirolaninae (Continued) 

4. Merus of pereopod 1 posterodistally produced; merus of pereopods 2 

and 3 anterodistally produced Bahalana 

Meri of pereopods 1—3 not markedly produced 5 

5. Animal able to conglobate Creaseriella 

Animal unable to conglobate 6 

6. Mandibular palp directed anteriorly Anopsilana 

Mandibular palp directed posteriorly Haptolana 

7. Pleopod 1, exopod longer and broader than endopod Oncilorpheus 

Pleopod 1, endopod longer and broader than exopod Calyptolana 

Anopsilana Paulian and Delamare Deboutteville, 1956 

DIAGNOSIS Body unable to conglobate. Eyes present or absent. Frontal 

lamina well developed, as long as broad, or longer than broad, anteriorly 

Key to species of Anopsilana 

1. Estuarine-brackish water species; integument pigmented when alive 2 

Cave species; lacking integumental pigment 3 

2. Frontal lamina distally rounded, projecting browni 

Frontal lamina distally acute, not projecting jonesi 

3. Posterior margin of pleotelson with 10 or more spines 4 

Posterior margin of pleotelson with less than 10 spines 5 

4. Posterior margin of pleotelson with 10 spines; found in cave on Cuba 

Posterior margin of pleotelson with 10—12 spines; found in cave on 

Haiti 

5. Posterior margin of pleotelson with eight spines; found in cave on 

cubensis 

acanthura 

Grand Cayman crenata 

Posterior margin of pleotelson with four spines; found in cave on Haiti 

radicicola

Anopsilana crenata 125 

somewhat expanded. Antennular peduncle of two articles. Maxillipedal en- 

dite with two coupling hooks. Pereopod 1 prehensile, pereopods 2-3 weakly 

prehensile, pereopods 4-7 ambulatory. Pleopod 2

126 FLABELLIFERA • GIROLANIDAE 

Figure 53. Anopsilana acanthura: A, 9; B, anterior cephalon. Anopsilana browni: C, 9; 

D, anterior cephalon. Anopsilana crenata: E, 9; F, anterior cephalon. Anopsilana 

cubensis: G, 9; H, anterior cephalon. 

Anopsilana cubensis (Hay, 1903) 

Figure 53G,H 

DIAGNOSIS 9 7.0 mm. Lacking eyes and integumental pigmentation. 

Frontal lamina longer than wide, anteriorly expanded, rounded. Posterior 

margin of pleotelson with 10 spines.

Bahalana 127 

RECORDS Caves in provinces of Pinar del Rio, La Habana, Matanzas, and 

on Isla de Pinos, Cuba. 

Anopsilana jonesi Kensley, 1987 

Figure 54A-D 

DIAGNOSIS 6 7.2 mm, ovigerous 9 5.9 mm. Eyes well developed and pig 

mented. Dorsal integument strongly pigmented with almost solid central 

area on pereonites 1-7. 6 cephalon with three low tubercles near posterior 

margin; pereonite 1 with four to six low tubercles. 9 lacking tubercles on 

cephalon or pereonite 1. Pereonites 2—7 with low submedian longitudinal 

ridges near posterior margin. Frontal lamina narrow, pentagonal, anteriorly 

acute, not projecting. Posterior margin of pleotelson with 9 or 10 spines. 

RECORDS Salt Creek, and Sittee River, Stann Creek District, Belize, in es- 

tuarine mangroves. 

Anopsilana radicicola (Notenboom, 1981) 

Figure 54E,F 

DIAGNOSIS 6 6.3 mm, 9 6.5 mm. Lacking eyes and integumental pigmen 

tation. Frontal lamina longer than wide, anteriorly expanded and rounded. 

Posterior margin of pleotelson with four spines. 

RECORDS Source Debarasse, a spring near Jeremie, Haiti. 

Bahalana Carpenter, 1981 

DIAGNOSIS Eyes lacking. Frontal lamina basally triangular, anteriorly nar 

rowed into poorly developed carina. Pereopods 1—3 prehensile, dactyli and 

propodi relatively elongate; pereopod 1 carpus small, almost concealed; 

merus with strong posterodistal extension almost reaching dactylar base and 

armed with spines. Pereopods 2 and 3, meri with elongate anterodistal exten 

sion, meri and carpi with shorter posterodistal extensions. Pereopods 4-7 

slender, ambulatory. Pleopod 2 in 6 with copulatory stylet articulating 

basally on endopod. Pleopods 3-5, exopods biarticulate, endopods with few 

distal marginal setae, or lacking setae. Pleonite 5 with free lateral margin, 

hardly overlapped by pleonite 4.


Key to species of Bahalana 

1. Pleopods 3—4, endopods lacking setae; maxillipedal endite with one 

coupling hook cardiopus 

Pleopods 3—4, endopods with few setae; maxillipedal endite with one or 

two coupling hooks 2 

2. Antennular peduncle, article 3 longest; maxillipedal endite with two 

coupling hooks geracei 

Antennular peduncle, article 2 longest; maxillipedal endite with one 

coupling hook mayana 

Bahalana cardiopus Notenboom, 1981 

Figure 55A,B 

DIAGNOSIS 9 10.0 mm. Maxillipedal endite with single coupling hook. Per- 

eopod 1, meral projection bearing five distal spines. Pleopods 3—5, endopods 

lacking marginal setae. Uropodal exopod half width of endopod, four spines 

on outer margin. 

RECORDS Mount Misery Cave, Mayaguana Island, Bahamas. 

Bahalana geracei Carpenter, 1981 

Figure 55C-G 

DIAGNOSIS 9 15.0 mm, 6 8 mm. Maxillipedal endite with two coupling 

hooks. Pereopod I, meral projection bearing seven distal spines. Pleopods 3- 

5, endopods with 9—13 distal marginal setae; pleopod 5 endopod with four 

distal setae. Uropodal endopod bearing few spines at outer distal margin, 

margin not serrate, lacking distinct apex; exopod half width of endopod, with 

four spines on outer margin. 

RECORDS Lighthouse Cave, San Salvador Island, Bahamas. 

Bahalana mayana Bowman, 1987 

Figure 55H 

DIAGNOSIS 9 8.4 mm, 6 10.0 mm. Clypeus acutely pointed. Antennular 

peduncle, article 2 longest. Maxillipedal endite with one coupling hook. Per-

B 

Bathynomus 129 

Figure 54. Anopsilana jonesi: A> 8\ By anterior cephalon; C, pleopod 2, 8; D, 

pleopod 3. Anopsilana radicicola: E> 8\ F> anterior cephalon. 

eopod 1, meral projection rudimentary. Pleopods 3—4, endopods with few 

marginal setae, pleopod 5 lacking marginal setae. Uropodal exopod narrow, 

V3 width of endopod; endopod with distal margin slightly concave. 

RECORDS Anchialine caves on Cozumel Island and at Tulum, Yucatan 

Peninsula, Mexico. 

Bathynomus A. Milne Edwards, 1879 

DIAGNOSIS Frontal lamina triangular; clypeus projecting anteriorly. Anten- 

nal peduncle with articles 3 and 4 subequal, article 5 longest. Maxillipedal 

endite with four to seven coupling hooks. Pereopods 1—3 with anterodistal 

margins of ischia and meri produced. Pleopods with all rami bearing margi 

nal setae; endopods bearing accessory gills at bases. Posterior margin of 

pleotelson dentate.

Figure 55. Bahalana cardiopus: A, 9\ B, pleopod 3. Bahalana geracei: C, 9; D, 

pleopod 3; E, pereopod 1; F, pereopod 2; G, anterior cephalon. Bahalana mayana: 

H, 9.

Figure 56. Bathynomus giganteus 

Bathynomus giganteus A. Milne Edwards, 1879 

Figure 56 

Calyptolana 131 

D I A G N O S I S U p t o 2 8 0 m m . L a r g e p i g m e n t e d e y e s p r e s e n t , n o t v i s i b l e d o r - 

s a l l y . A n t e n n u l e w i t h s m a l l e x o p o d d i s t a l l y o n p e d u n c u l a r a r t i c l e 3 . P l e o p o d s 

w i t h m a r g i n a l s e t a e o n a l l r a m i ; p l e o n i t e s 3 a n d 4 w i t h e p i m e r a p r o d u c e d 

p o s t e r i o r l y . P o s t e r i o r m a r g i n o f p l e o t e l s o n w i t h m e d i a n t o o t h a n d f i v e o r s i x 

t e e t h o n e a c h s i d e . 

R E C O R D S G u l f o f M e x i c o ; C a r i b b e a n ; B a h a m a s ; F l o r i d a K e y s ; 3 6 0 - 2 3 0 0 

m . 

R E M A R K S G u t - c o n t e n t a n a l y s i s o f t h e s e d e e p - w a t e r g i a n t s h a s s h o w n t h e m 

t o b e s c a v e n g e r s , c o m m o n l y f e e d i n g o n d e a d f i s h , c e p h a l o p o d s , c r a b s , a n d 

p o l y c h a e t e w o r m s . 

Calyptolana Bruce, 1985 

D I A G N O S I S A l l p e r e o p o d s a m b u l a t o r y ; e a c h d a c t y l u s w i t h s e c o n d a r y u n 

g u i s . P l e o p o d 1 o p e r c u l a t e , l o n g e r t h a n f o l l o w i n g p l e o p o d s . A l l p l e o p o d a l 

r a m i e x c e p t e n d o p o d o f p l e o p o d 5 w i t h m a r g i n a l s e t a e .

132 FLABELLIFERA • C1ROLANIDAE 

Calyptolana hancocki Bruce, 1985 

Figure 57 

DIAGNOSIS 9 3.0 mm. Body dorsally strongly convex. Cephalon with small 

rostrum curving ventrally to meet frontal lamina; latter pentagonal. Eyes 

small, well pigmented. Coxae of pereonites barely produced. Pleonite 5 lack 

ing free lateral margins. Pleotelson with broadly rounded posterior margin. 

Uropodal sympod produced along mesial margin of endopod; exopod 

slightly less than half length of endopod; latter distally rounded. 

RECORDS Dominican Republic; Aruba Island, Netherlands Antilles, 43.2 

m. 

Cirolana Leach, 1818 

DIAGNOSIS Frontal lamina usually twice as long as wide, not projecting; 

clypeus flat. Mandible with strong incisor, dentate molar, palp of three arti 

cles. Pereopods all ambulatory. Pleon consisting of five free pleonites plus 

pleotelson, pleonite 5 overlapped laterally by pleonite 4. All pleopodal rami 

Key to species of Cirolana 

1. Pleotelson posteriorly very broad to subtruncate 2 

Pleotelson posteriorly narrowed 4 

2. Uropodal endopod broad, distally rounded, lacking marginal spines 

Uropodal endopod broad, distal margin having apical tooth or 

obtruncata 

angle 3 

3. Posterior margin of pleotelson bearing spines; uropodal endopod not 

dentate minuta 

Posterior margin of pleotelson faintly crenulate, lacking spines; 

uropodal endopod strongly dentate crenulitelson 

4. Uropodal endopod evenly tapering to acute apex; uropodal exopod 

length about four times greatest width albidoida 

Uropodal endopod, outer margin convex; uropodal exopod length less 

than four times greatest width parva

Figure 57. Calyptolana hancocki: A} 9; B, ventral pleon. 

Cirolana crenulitelson 133 

bearing marginal setae except endopod of pleopod 5. Copulatory stylet on 

endopod of 6 pleopod 2 inserted proximally. 

Cirolana albidoida Kensley and Schotte, 1987 

Figure 58A-C 

DIAGNOSIS 6 7.8 mm. Integument sparsely pitted. Antenna reaching pos 

teriorly to pereonite 3. Uropodal endopod triangular, evenly tapering, mar 

gins serrate; exopod length about four times greatest width, apically acute. 

Pleopod 1 6, endopod markedly narrowed in distal half. Pleopod 2 6, copu 

latory stylet reaching beyond rami by about half its length. Pleotelson with 

sides gently convex, tapering to rounded posterior margin bearing eight 

spines and three or four small serrations anterior to first spine. 

RECORDS OffLucaya, Grand Bahama, 180-220 m. 

Cirolana crenulitelson Kensley and Schotte, 1987 

Figure 58D-F 

DIAGNOSIS 6 7.0 mm, 9 7.0 mm. Antenna reaching posteriorly to anterior 

of pereonite 3. Pleopod 2 8, copulatory stylet reaching by Vs its length be-


Figure 58. Cirolana albidoida: A> uropod; C, pleotelsonic apex. Cirolana 

crenulitelson: D, 9; Ey pleotelsonic apex; Fy uropod. Cirolana minuta: G, 6; H, 

anterior cephalon; /, uropod. 

yond rami. Uropodal endopod with mesial margin broadly convex, serrate, 

apicaily acute; exopod about 2.5 times longer than wide, mesial margin 

weakly convex, apicaily acute. Posterior margin of pleotelson truncate, 

faintly crenulate, lacking spines. 

RECORDS Carrie Bow Cay, Belize, 36 m.

Cirolana minuta Hansen, 1890 

Figure 58G-I 

Cirolana parva 135 

DIAGNOSIS 8 8.9 mm, 9 5.0 mm. Antenna reaching posteriorly to per- 

eonite 3. Pleopod 2 copulatory stylet just reaching to distal mar 

gin of rami. Uropodal endopod distally broadly rounded, margin with 

rounded teeth; exopod 2.5 times longer than wide, margin with rounded 

teeth, apically obscurely subacute. Posterior margin of pleotelson subtrun 

cate, with about eight spines. 

RECORDS Jamaica; Puerto Rico; Cozumel, Mexico; Gulf of Mexico. 

Cirolana parva Hansen, 1890 

Figures 59C-E, 60 

DIAGNOSIS 6 6.9 mm, 9 7.9 mm. Cephalon with furrow between eyes hav 

ing middorsal posterior deflection. Antenna reaching posteriorly to pereonite 

4. Pleopod 1 (5, endopod narrowed in distal third. Pleopod 2 6, copulatory 

stylet reaching by 'A its length beyond rami. Uropodal endopod with mesial 

margin convex, apically acute; exopod 2.5 times longer than wide, apically 

acute. Pleotelson evenly tapering to angled posterior margin, with seven or 

eight spines. 

RECORDS North and South Carolina; Turks and Caicos Islands; St. 

Thomas and St. Croix, U.S. Virgin Islands; Andros Island, Bahamas; Pu 

erto Rico; Jamaica; Florida Keys; Dry Tortugas; Barbados; Carrie Bow Cay, 

Belize; Cozumel, Mexico; Panama; Gulf of Mexico; intertidal to 55 m.

Figure 59. Cirolana obtruncata: A, $; B, uropod. Cirolana parva: C, 9; D, uropod; E, 

pleotelson and uropods.

Figure 60. Cirolana parva, 

anteroventral cephalon. 

C r e a s e r i e l l a R i o j a , 1 9 5 3 

Haptolana 137 

D I A G N O S I S A n t e n n u l a r p e d u n c l e o f t w o a r t i c l e s ( a r t i c l e s a n d f u s e d ) . 

A n t e n n a l p e d u n c l e o f f i v e a r t i c l e s . P e r e o p o d s a l l a m b u l a t o r y . P e n i a l r a m i 

f u s e d t o f o r m s h o r t s t o u t p r o c e s s . P l e o p o d 2 i n cT w i t h c o p u l a t o r y s t y l e t i n s e r - 

t e d a t b a s e o f e n d o p o d . P l e o p o d s 

P l e o n i t e 5 l a c k i n g f r e e l a t e r a l m a r g i n s . 

Creaseriella anops (Creaser, 1936) 

Figure 61A,B 

, e n d o p o d s l a c k i n g m a r g i n a l s e t a e . 

DIAGNOSIS 1 5 . 5 m m . A n i m a l a b l e t o c o n g l o b a t e . E y e s l a c k i n g . F r o n t a l 

l a m i n a p e n t a g o n a l , l o n g e r t h a n w i d e , w i t h t r a n s v e r s e r i d g e a t w i d e s t p o i n t . 

M a x i l l i p e d a l e n d i t e w i t h f o u r o r five c o u p l i n g h o o k s . P l e o t e l s o n w i d e r t h a n 

l o n g , p o s t e r i o r m a r g i n v e r y b r o a d l y r o u n d e d a n d f i n e l y c r e n u l a t e . B o t h 

u r o p o d a l r a m i d i s t a l l y r o u n d e d , m a r g i n s b e a r i n g s p i n e s a n d s e t a e , s y m p o d 

p r o d u c e d a l o n g m e s i a l m a r g i n o f e n d o p o d . 

RECORDS S e v e r a l c a v e s a n d c e n o t e s o n t h e Y u c a t a n P e n i n s u l a , M e x i c o . 

H a p t o l a n a B o w m a n , 1 9 6 6 

D I A G N O S I S M a n d i b u l a r p a l p d i r e c t e d p o s t e r i o r l y . A n t e n n u l a r p e d u n c l e o f 

t w o a r t i c l e s , b a s a l a r t i c l e e x p a n d e d . P e r e o p o d s a l l p r e h e n s i l e , w i t h d a c t y l i 

c l o s i n g i n p r o p o d a l g r o o v e . P l e o p o d s 3 - 5 , e x o p o d s w i t h p a r t i a l s u t u r e a n d 

m a r g i n a l s e t a e ; e n d o p o d s u n d i v i d e d , l a c k i n g m a r g i n a l s e t a e . P l e o n i t e 4 o v e r 

l a p p i n g p l e o n i t e 5 l a t e r a l l y .


Figure 61. Creaseriella anops: A, adult; B, frontal lamina. Haptolana trichostoma: C, 

adult; D, frontal lamina. Oncilorpheus stebbingi: £, juvenile (from Paul and Menzies, 

1971). 

Haptolana trichostoma Bowman, 1966 

Figure 61C,D 

DIAGNOSIS 6 13.8 mm. Eyes lacking. Frontal lamina broad, pentagonal, 

with anterior angle very broad. Pleotelson wider than long, roughly rec 

tangular, posterior margin faintly crenulate, with spine between and setae on 

crenulations. Uropodal endopod distally broad, exopod distally narrowly 

rounded, 2 fo width of endopod, both rami with marginal setae and spines. 

RECORDS Cave in Camaguey Province, Cuba. 

REMARKS A second species of Haptolana, H. somata Messana and Chelazzi, 

has been described from northern Somalia in Africa.

Oncilorpheus Paul and Menzies, 1971 

Natatolana 139 

DIAGNOSIS Frontal lamina projecting ventrally. Pleopod 1, exopod indu 

rate, opercular; endopod membranous, less than half width of exopod. 

Uropodal sympod longer than rami, slightly produced along mesial margin 

of endopod; rami inserted subapically. 

Oncilorpheus stebbingi Paul and Menzies, 1971 

Figure 61E 

DIAGNOSIS 9 11.0 mm. Body narrow, about five times longer than wide. 

Pleotelson triangular, bearing faint middorsal longitudinal ridge, apex nar 

rowly rounded. 


Subfamily Conilerinae, new name 

DIAGNOSIS Clypeus flattened. Frontal lamina flat, narrow. Antennal ped 

uncular articles 3 and 4 subequal. Pereopods 1-3, ischium and merus pro 

duced anterodistally. Pereopods lacking secondary unguis on dactyli. Nata 

tory setae present on pereopods 4—7. 

REMARKS In a discussion of the Cirolanidae, Botosaneanu, Bruce, and 

Notenboom (1986:412) refer to the subfamilies Eurydicinae and Cirolaninae 

but place the Conilera group under the heading "Unnamed Subfamily." For 

consistency, the Conilera group is here recognized as a subfamily. 

Key to genera of Conilerinae 

1. Uropodal endopod with distal notch; pereopods 4-7 natatory, ischium, 

merus, and carpus flattened Politolana 

Uropodal endopod lacking distal notch; pereopods 4-7 with basis 

flattened and expanded, bearing natatory setae Natatolana 

Natatolana Bruce, 1981 

DIAGNOSIS Frontal lamina narrow; clypeus flat. Pereopods 1-3 bearing 

long setae. Pleopod 2


Natatolana gracilis (Hansen, 1890) 

Figure 62 

DIAGNOSIS 8 8.0 mm. Antennule short, not reaching distal end of antcnnal 

peduncle. Antenna reaching posteriorly to pereonite 4. Pleopod 2, copulatory 

stylet of endopod cylindrical, bowed, distally rounded. Pleotelson with ob 

tuse apex, with slight transverse indentation near anterior margin. 

RECORDS Probably St. Thomas, U.S. Virgin Islands; off Sombrero Light, 

Florida, 100-120 m. 

Northern Brazil, 7-85 m. 

REMARKS Hansen (1890) indicated some uncertainty about the exact type 

locality, but thought it likely to have been St. Thomas. Koening (1972) re 

corded this species from several localities off northern Brazil, but did not 

illustrate her material. 

Politolana Bruce, 1981 

DIAGNOSIS Frontal lamina slender, flattened; clypeus flattened. Antennal 

peduncular articles 3—5 subequal. Pereopods 1—3 with ischium and merus 

anterodistally produced. Pereopods 4—7 with ischium, merus, and carpus 

flattened. 6 pleopod 2 with copulatory stylet inserted basally on endopod. 

Endopod of pleopod 5 lacking marginal setae. Pleonite 5 overlapped laterally 

by pleonite 4. Uropodal endopod with distal emargination; exopod slender, 

elongate; sympod produced along mesial margin of endopod. 

Key to species of Politolana 

1. Uropodal endopod broad distal to emargination, margin obliquely 

truncate; coxae of pereonites 2—6 with impressed line impressa 

Uropodal endopod distal to emargination somewhat narrowed, margin 

evenly convex; coxae of pereonites lacking impressed line polita 

Politolana impressa (Harger, 1883) 

Figure 63A,B 

DIAGNOSIS 6 and 9 up to 27 mm. Frontal lamina slightly expanded ante 

riorly. Coxae of pereonites 2-6 with impressed oblique line. Uropodal endo-

Figure 62. Natatolana gracilis: A, 9; B, frontal lamina; C, pleotelson; D> uropod; E> 

pereopod 7.


Figure 63. Politolana impressa: A, 6, lateral view; B, uropod, (setae and spines 

omitted). Politolana polita: Cy 6; D, pereopod 1; Ey uropod, setae and spines 

omitted; F, pereopod 1. 

pod broad distal to emargination, margin subtruncate. Pleotelson posteriorly 

broadly rounded. 

RECORDS Massachusetts to Palm Beach, Florida, 32-650 m.

Figure 64. Politolana polita, 

anteroventral cephalon. 

Politolana polita (Stimpson, 1853) 

Figures 63C-F, 64 

Politolana polita 143 

DIAGNOSIS 2 7 . 0 m m , 9 2 9 . 0 m m . A n t e n n u l e b a r e l y r e a c h i n g d i s t a l e n d 

o f a n t e n n a l p e d u n c l e . F r o n t a l l a m i n a b a s a l l y s l e n d e r , a n t e r i o r l y e x p a n d e d , 

j u s t v i s i b l e i n d o r s a l v i e w . C o x a e l a c k i n g i m p r e s s e d o b l i q u e l i n e . U r o p o d a l 

e n d o p o d d i s t a l t o e m a r g i n a t i o n n a r r o w e d , m a r g i n c o n v e x . P l e o t e l s o n p o s 

t e r i o r l y b r o a d l y r o u n d e d . 

RECORDS 

Mexico. 

Bay of Fundy, Canada, to Florida Keys, 2-600 m; Gulf of 

Subfamily Eurydicinae Stebbing, 1905 

D I A G N O S I S C l y p e u s p r o j e c t i n g . P l e o n i t e 5 w i t h f r e e l a t e r a l m a r g i n s ( e x c e p t 

i n X y l o l a n a ) . P e n e s p r o m i n e n t . P l e o p o d 2 o f 6 w i t h c o p u l a t o r y s t y l e t a r t i c 

u l a t i n g s u b b a s a l l y , m e d i a l l y , o r s u b a p i c a l l y . 

Key to genera of Eurydicinae 

Uropodal sympod not produced along mesial margin of endopod 

Eurydice 

Uropodal sympod produced along mesial margin of endopod 

Rostrum prominent, fused with frontal lamina, separating antennal 

bases 

Rostrum not prominent 

{continued)


Key to genera of Eurydicinae {Continued) 

3. Picon of five free plconites plus pleotelson 4 

Picon of three free plconites plus pleotelson Colopisthus 

4. Endopods of plcopods 3-5 lacking marginal setae, or with no more 

than three marginal setae; copulatory stylet of endopod of pleopod 2 

in 6 articulating subterminally Arubolana 

Marginal setae lacking only on endopod of pleopod 5; copulatory stylet 

of pleopod 2 endopod in 8 articulating basally Metacirolana 

5. Clypeus conical; uropodal endopod lacking notch in outer distal margin 

Xylolana 

Clypeus flattened; uropodal endopod with notch in outer distal margin 

Arubolana Botosaneanu and Stock, 1979 

Excirolana 

DIAGNOSIS Animal not able to conglobate. Blind, or with very small eyes. 

Anterior margin of frontal lamina broad. Antennular peduncle of three arti 

cles. Maxillipedal palp of four articles (articles 2 and 3 fused). Maxilla 2 

reduced, endite unarmed, exopod with few marginal setae. Pereopods 1 and 

2 and sometimes pereopod 3 prehensile; pereopods 4—7 ambulatory. 

Pleopods 1 and 2, rami undivided. Pleopod 2 6 with copulatory stylet artic 

ulating subterminally on endopod. Pleopods 3-5, exopods biarticulate; endo 

pods lacking marginal setae, or with few setae on endopods of 3 and 4; 

pleopod 5 exopod with marginal setae interrupted. 

Key to species of Arubolana 

1. Eyes present, small parvioculata 

Eyes absent 2 

2. Pleotelson posteriorly rounded; rostrum not distinct in dorsal view 

Pleotelson posteriorly subtruncate; rostrum distinct in dorsal view 

aruboides 

imula

Arubolana aruboides (Bowman and IlifTe, 1983) 

Figure 65A-C 

Arubolana parvioculata 145 

DIAGNOSIS 6 3.9 mm, 9 4.1 mm. Body about three times longer than wide. 

Eyes absent. Antennular peduncle article 3 longer than articles 1 and 2 to 

gether. Antenna reaching posteriorly to pereonite 6 or 7. Frontal lamina vis 

ible in dorsal view between antennal bases, anteriorly rounded and only 

slightly wider than proximally, with distally flared ridge on ventral (exposed) 

surface. Pleotelson as long as basal width, evenly narrowing to broadly 

rounded posterior margin, latter with few small serrations and setae. Uropo- 

dal exopod four times longer than wide; endopod length about twice basal 

width, distally obliquely truncate, with elongate setae on inner margin. 

RECORDS Church Cave and Wonderland Cave, Bermuda. 

Arubolana imula Botosaneanu and Stock, 1979 

Figure 65D 

DIAGNOSIS 6 and 9 6.25 mm. Body 2.3 times longer than wide. Eyes ab 

sent. Antenna reaching posteriorly to pereonite 4 or 5. Rostrum distinct, 

anteriorly truncate, separating antennal bases, fused with rectangular frontal 

lamina ventrally. Pleotelson basally slightly wider than long, posterior mar 

gin broadly rounded to subtruncate, with irregular crenuiations or faint 

teeth. Uropodal exopod apically acute, reaching to about midlength of endo 

pod; latter distally broad, with slight tooth at distolateral angle. 

RECORDS Mangel Cora Tunnel, Aruba. 

Arubolana parvioculata Notenboom, 1984 

Figure 65E,F 

DIAGNOSIS 6 2.8 mm, 9 2.9 mm. Body 3.3 times longer than wide. 

Cephalon with tiny pigmented eyes. Antenna reaching posteriorly to per 

eonite 5. Frontal lamina projecting, dorsally visible. Pleotelson basally wider 

than long, tapering to broadly rounded/subtruncate posterior margin bear 

ing about six low teeth. Uropodal exopod distally acute, almost three times 

longer than basal width; endopod distally serrate, apically acute. 

RECORDS Interstitial water near Discovery Bay, Jamaica.


Figure 65. Arubolana aruboides: A, 8; B, pereopod 1; C, pereopod 2. Arubolana imula 

D, 9. Arubolana parvioculata: Ey 6\ Fy pleopod 2, 8. 

Colopislhus Richardson, 1902 

DIAGNOSIS Cephalon broader than long, becoming triangular between an- 

tennal bases. Pleon consisting of three short free pleonites (often obscured 

beneath pereonite 7), plus triangular pleotelson. 

D

Colopisthus parvus Richardson, 1902 

Figure 66A 

Eurydice convex a 147 

DIAGNOSIS 9 3.6 mm. Eyes large, well pigmented. Frontal lamina prox- 

imally narrow, anteriorly widened to truncate distal margin. Antennules and 

antennae short, latter reaching posteriorly to pereonite 1. Pleotelson with 

strong middorsal ridge. 

RECORDS Bermuda; Puerto Rico, intertidal rocks and algae. 

Eurydice Leach, 1815 

DIAGNOSIS Antennular peduncle article 2 at right angle to article 1. Anten- 

nal peduncle of four articles. Frontal lamina usually slender; clypeus usually 

a ventrally directed triangular blade. Maxillipedal endite reduced, lacking 

coupling hooks. 6 pleopod 2 with copulatory stylet articulating at mid- 

length. Pleopod 5, endopod lacking marginal setae. Pleonite 5 with free lat 

eral margins, not overlapped by pleonite 4. Uropodal sympod not produced 

along medial margin of endopod. 

Key to species of Eurydice 

1. Frontal lamina distally truncate to faintly bilobed 2 

Frontal lamina lanceolate, distally acute personata 

2. Posterior margin of pleotelson between notches rounded, with four 

moderate spines convexa 

Posterior margin of pleotelson between notches almost straight, with 

four very short spines and several elongate setae piperata 

Eurydice convexa Richardson, 1900 

Figures 66B-E, 67A,B 

DIAGNOSIS 6 6.1 mm, 9 6.1 mm. Frontal lamina slender, anteriorly widen 

ing slightly and becoming truncate to faintly bilobed. Posterior margin of 

pleotelson between lateral notches convex, with four spines and few setae 

between serrations; spines between three and four times longer than wide. 6: 

Plicate process on antennal flagellar articles about l /s length of article. 

Pleopod 2 with copulatory stylet distally blunt, reaching well beyond rami.


Figure 66. Colopisthus parvus: A> 9. Eurydice convexa: B, 6; C, uropod; Dy antennule; 

E, pleotelsonic apex. Eurydice per sonata: F> 8, lateral view; Gy pleotelsonic apex. 

Eurydice piperata: Hy pleotelsonic apex. 

RECORDS South Carolina to Florida Keys; Bahamas; Gulf of Mexico and 

Caribbean. 

REMARKS Whether E. convexa and E. littoralis are conspecific needs further 

investigation. Differences can be detected in the male plicate organs of the 

antennae, and in the mandibular palp spination, but range of variation in

Excirolana 149 

these features is still unknown. It seems unlikely that the species recorded as 

E. littoralis by Moreira (1972) from Brazil is the same species. 

Eury dice per sonata Kensley, 1987a 

Figures 66F,G; 67C 

DIAGNOSIS 6 6.0 mm, ovigerous 9 5.1—6.4 mm. Frontal lamina slender, 

lanceolate, anteriorly acute. Posterior margin between notches faintly con 

vex, with four relatively elongate spines (inner pair five or six times longer 

than wide) and few setae between dentitions. 6: Plicate organ on antennal 

flagellar articles half length of article. Pleopod 2, copulatory stylet on endo- 

pod clavate, barely reaching beyond ramus. 

RECORDS Bermuda; off Georgia, 18-27 m; off South Carolina, 34 m; off 

Miami, Florida; Puerto Rico, 13—17 m; Bahamas, 1—2 m and surface 

plankton tow; Haiti; Cuba; Venezuela. 

REMARKS This recently discovered species has masqueraded under the 

names of E. convexa and E. littoralis for some time, which may explain some of 

the inconsistencies in the literature, especially in variation in the pleotelsonic 

apex. 

Eury dice piper at a Menzies and Frankenberg, 1966 

Figure 66H 

DIAGNOSIS 6 4.0 mm, 9 4.5 mm. Frontal lamina slender, widening ante 

riorly to become slightly bilobed. Posterior margin of pleotelson between 

notches straight to faintly convex, with four spines barely twice longer than 

wide, and several much longer setae between dentition. 6: Plicate organ on 

antennal flagellar articles about l /e length of article but situated subdistally. 

Pleopod 2, copulatory stylet clavate, reaching well beyond ramus. 

RECORDS Georgia to Florida, Gulf of Mexico, 37-150 m. 

Excirolana Richardson, 1912a 

DIAGNOSIS Cephalon with prominent rostrum separating antennular bases; 

fused with flattened frontal lamina. Clypeus with short, broadly triangular 

blade projecting anteroventrally. Antennal peduncle with four or five arti 

cles. Maxillipedal endite with single coupling hook. Pleopods 3—5, endopods


F i g u r e 6 7 . E u r y d i c e c o n v e x a : A , a n t e r o v e n t r a l c e p h a l o n ; B , f r o n t a l l a m i n a . E u r y d i c e 

per sonata: C, frontal lamina. 

l a c k i n g m a r g i n a l s e t a e . P l e o n i t e 5 w i t h f r e e l a t e r a l m a r g i n s , n o t o v e r l a p p e d 

b y p l e o n i t e 4 . U r o p o d a l s y m p o d p r o d u c e d a l o n g m e s i a l m a r g i n o f e n d o p o d . 

Key to species of Excirolana 

1 . P l e o t e l s o n w i t h t w o a n t e r i o r h o l l o w s c l e a r l y j o i n e d b y i m p r e s s e d l i n e ; 

u r o p o d a l e n d o p o d a b o u t h a l f l e n g t h o f e x o p o d braziliensis 

P l e o t e l s o n w i t h t w o a n t e r i o r h o l l o w s n o t c o n n e c t e d b y i m p r e s s e d l i n e ; 

u r o p o d a l e n d o p o d a b o u t t w o - t h i r d s l e n g t h o f e x o p o d mayana 

Excirolana braziliensis Richardson, 1912a 

F i g u r e s 6 8 A - C , 6 9 A - C 

D I A G N O S I S 6 6 . 0 m m , 9 7 . 5 m m . F r o n t a l l a m i n a v e r y s l e n d e r b e t w e e n 

a n t e n n a l b a s e s , w i d e n i n g a n t e r i o r l y i n t o r o u n d e d s t r u c t u r e b e t w e e n a n t e n -

Figure 68. Excirolana braiiliensis: Ay 6; B, pleopod 2 8; C, uropod, (setae and 

spines omitted). Excirolana mayana: D, 9; Ey pleopod 2 6\ F, uropod, setae and 

spines omitted.


400HM M'Hrl 

F i g u r e 6 9 . Excirolana b r a z i l i e n s i s : A , a n t e r o v e n t r a l c e p h a l o n ; B , p l e o t e l s o n ; 

u r o p o d a l e n d o p o d . Excirolana mayana: D , a n t e r o d o r s a l c e p h a l o n ; E , a n t e r c 

c e p h a l o n . 

n u l a r b a s e s , j o i n e d t o r o s t r u m b y v e r y s l i m i s t h m u s . C l y p e u s d i s t a l l y s u b 

a c u t e . U r o p o d a l e n d o p o d a b o u t h a l f l e n g t h o f e x o p o d . P l e o t e l s o n w i t h t w o 

l a t e r a l h o l l o w s d e f i n e d a n d c o n n e c t e d b y c l e a r i m p r e s s e d l i n e ; p o s t e r i o r m a r 

g i n e v e n l y c o n v e x , b e a r i n g n u m e r o u s p l u m o s e s e t a e .

Metacirolana 153 

RECORDS Caribbean to Brazil; common in the intertidal of sandy beaches; 

Gulf of Mexico. 

Gulf of California to Chile. 

REMARKS Glynn et al. (1975) produced a thorough study of the taxonomy, 

zonation, and distribution of this Pan-American species. 

Excirolana may ana (Ives, 1891) 

Figures 68D-F, 69D,E 

DIAGNOSIS 6 8.2 mm, 9 10.0 mm. Frontal lamina between antennal bases 

about half anterior width. Clypeus anteriorly rounded. Uropodal endopod 

about 2 /3 length of exopod. Pleotelson with two faint lateral hollows in ante 

rior half, not connected by impressed line. 

RECORDS Florida to Venezuela, intertidal. 

Metacirolana Nierstrasz, 1931 

DIAGNOSIS Frontal lamina anteriorly dilated, free, projecting; clypeus tri 

angular, projecting ventrally. Maxillipedal endite with one coupling hook. 

Pleon with five free segments, pleonite 5 not overlapped laterally by pleonite 

4. Eyes often larger, and antennular flagellum of more articles in 6 than in 

2. 

Key to species of Metacirolana 

1. Telson posteriorly truncate halia 

Telson posteriorly rounded or angulate 2 

2. Posterior margin of telson an obtuse angle agaricicola 

Telson posteriorly rounded 3 

3. Posterior margin of telson narrowly rounded; uropodal rami, margins 

strongly dentate menziesi 

Posterior margin of telson broadly rounded; uropodal rami, margins 

obscurely dentate sphaeromiformis


Metacirolana agaricicola Kensley, 1984 

Figure 70A-C 

DIAGNOSIS 6 2,6 mm, ovigerous 9 2.1 mm. Antennular flagellum of six or 

seven articles. Antennal flagellum of 10 articles. Posterior margin of telson 

finely dentate, with broadly obtuse median point. Uropodal exopod about 

half width of endopod, margins dentate, apically acute; endopod, margins 

dentate, distally angled, apically acute. 

RECORDS Carrie Bow Cay, Belize, 1-20 m, in coral on reef slope, and spur 

and groove zone. 

Metacirolana halia Kensley, 1984 

Figure 70D-F 

DIAGNOSIS 6 2.9 mm, ovigerous 9 2.7 mm. Antennular flagellum of 10 

articles in 9, 14 in 6. Antennal flagellum of 10 articles in 9, 11 in 6. Pos 

terior margin of telson truncate, bearing about eight sensory spines. Uropo 

dal exopod distally broadly rounded, more than half distal width of endopod, 

outer margin dentate, with about 11 sensory spines; endopod distally broad, 

margin straight, bearing about 12 sensory spines. 

RECORDS Carrie Bow Cay, Glover's Reef, Belize; intertidal to 23 m; Turks 

and Caicos Islands, 1 m; Bahamas; Jamaica; Cozumel, Mexico. 

Metacirolana menziesi Kensley, 1984 

Figure 71A,B 

DIAGNOSIS 8 2.3 mm, ovigerous 9 2.4 mm. Antennular flagellum of six 

articles in 9, eight in 6. Antennal flagellum of nine articles in 9, 10 in

Figure 70. Metacirolana agaricit 

halia: D, uropod; E, 8; F, 9. 

rou 

pair 

Metacirolana sphaeromijormis 155 

nded, obscurely dentate. Telson with low rounded middorsal ridge and 

of lateral ridges. Uropodal exopod more than half width ofendopod, 

margin dentate; exopod distally broadened, margin dentate, with few sen 

sory spines


Figure 71. Metacirolana menziesi: A, 9; B, uropod. Metacirolana sphaeromiformis: C, 

D, uropod. 

RECORDS Looe Key, Florida, intertidal reef crest; Turks and Caicos Is 

lands, 1 m; St. Thomas, U.S. Virgin Islands. 

Xylolana Kensley, 1987a 

DIAGNOSIS Frontal lamina and rostrum fused, broad, separating antennu- 

lar bases. Clypeus conical, projecting. Maxillipedai endite reduced, lacking

A Icirona 15 7 

coupling hooks; palp of five articles. Gopulatory stylet in 6 articulating in 

distal half of mesial margin of pleopod 2 endopod. Pleopods 3-5, exopods 

biarticulate; endopods lacking marginal setae. Pleonite 5 lacking free lateral 

margins, overlapped by pleonite 4. Uropodal sympod produced along mesial 

margin of endopod. 

Xylolana radicicola Kensley, 1987a 

Figure 72 

DIAGNOSIS 8 2.6 mm, 9 3.3 mm. Body about four times longer than great 

est width. Uropodal exopod about 2 fo width of endopod, bearing single short 

subapical spine; both uropodal rami distally rounded. Pleotelson with lateral 

margins subparallel, with poorly defined middorsal longitudinal ridge. 

RECORDS Twin Gays, Belize, in dead red mangrove roots, 1 m. 

Family Corallanidae Hansen, 1890 

DIAGNOSIS Outer ramus of maxilla 1 apex with single strong falcate spine, 

with single strong spine with one or more smaller hooked spines at base, or 

with two large recurved spines, occasionally with one to three smaller spines 

between them. Maxillipedal endite reduced or lacking. 

Key to genera of Corallanidae 

1. Maxilla 1 with single strong falcate spine 2 

Maxilla 1 with two large falcate spines Alcirona 

2. Maxilla 2, distal article slender-elongate; article 2 of maxillipedal palp 

longest Nalicora 

Maxilla 2 distally bluntly bilobed; article 3 of maxillipedal palp longest 

Alcirona Hansen, 1890 

Excorallana 

DIAGNOSIS Antennular peduncle of two articles. Mandible lacking molar. 

Maxilla 1 having two large recurved spines, with one or more smaller spines 

between these. Maxilla 2 a simple rounded lobe. Maxilliped lacking endite. 

Posterior half of body hirsute.

158 FLABELLIFERA • CORALLANIDAE 

Key to species of Alcirona 

1. Golden-brown setae starting dorsally on pereonite 3; pereopod 1, 

dactylus having several elongate spines insularis 

Golden-brown setae starting dorsally on pereonites 5 or 6; pereopod 1, 

dactylus having accessory spine only krebsi 

Alcirona insularis Hansen, 1890 

Figure 73A 

DIAGNOSIS 8 and 9, 5.0 mm. Posterior half of body, especially in (J, bear 

ing stiff golden-brown setae, these beginning as posterior row on pereonite 3, 

and becoming dense on posterior pereonites, pleonites and pleotelson. Per- 

eopods 1— 3, dactylus strongly falcate, having distal unguis equal in length to 

rest of article, and with three or four strong teeth on posterior margin. Apex 

of pleotelson rounded, bearing six short marginal spines in addition to setae. 

RECORDS Looe Key, Florida, 0.5-6 m; St. Thomas, U.S. Virgin Islands, 

40—46 m; Puerto Rico, from intertidal coral rubble and from gills of nurse 

shark Ginglymostoma sirratum; St. Lucia, from coral rubble. 

Alcirona krebsii Hansen, 1890 

Figure 73B-D 

DIAGNOSIS 8 10 mm, ovigerous 9 15.5 mm. Posterior half of body, espe 

cially in (J bearing stiff golden-brown setae, these beginning in posterior row 

on pereonites 5 or 6, becoming dense on posterior pereonites, pleonites, 

uropods, and pleotelson. Pereopod 1, dactylus strongly falcate, with unguis 

equal in length to rest of article, and with one strong tooth and several low 

tubercles on posterior margin. Apex of pleotelson rounded, bearing six short 

spines in addition to setae. 

RECORDS Bermuda, in sponges; Florida Keys; Quintana Roo, Yucatan 

Peninsula; St. Thomas, U.S. Virgin Islands; Venezuela. 

REMARKS A single 8-mm 8 specimen of Alcirona from Panama Bay (Pa 

cific) has the characteristic pereopod 1 of A. krebsii, but has the rows of stiff 

setae beginning on about pereonite 3. The possibility that A. krebsii is another 

amphi-Panamic species needs to be investigated. 

*

Excorallana 159 

Figure 72. Xylolana radicicola: A, By 6; C, maxilliped; D, pleopod 2 6; E, pleopod 3. 

Excorallana Stebbing,1904 

DIAGNOSIS Eyes well developed and pigmented, sometimes contiguous or 

nearly so. Maxilla 1, outer ramus a single falcate spine. Maxillipedal palp of 

five articles; endite reduced or absent. Pereopods 1—3 subprehensile or pre-


Figure 73. Alcirona insularis: A, pereopod 1. Alcirona krebsi: B3 d\ C, maxilla 1; D, 

pereopod 1. 

hensile, pereopods 4—7 ambulatory. All rami of pleopods bearing marginal 

setae. Pleotelson often with characteristic spination and tubcrculation; lat 

eral margins often with incision. 

REMARKS Excorallana subtilis (Hansen, 1890) was described from St. 

Thomas, U.S. Virgin Islands, based on a specimen undergoing ecdysis; the 

true identity of this species remains uncertain. 

In the key, two species have been included which are not illustrated. These 

are E. mexicana Richardson, 1905, from the Gulf of Mexico, and E. delaneyi

Excorallana 161 

Stone and Heard, 1989, from the northeastern Gulf of Mexico. The latter 

species, particularly, could conceivably be encountered in the Florida Keys. 

Delaney (1984) provides a useful review of the genus Excorallana, and of the 

distribution of the species. 

Key to species of Excorallana 

1. Eyes contiguous 2 

Eyes well separated 4 

2. Apex of pleotelson with deep slit Jissicauda 

Apex of pleotelson entire 3 

3. Pleotelson with lateral incision oculata 

Pleotelson lacking lateral incision warmingii 

4. Pleotelson with lateral incision 6 

Pleotelson lacking lateral incision 5 

5. Frontal lamina linguiform, anteriorly rounded berbicensis 

Frontal lamina posteriorly with faintly concave margins, anteriorly 

subacute delaneyi 

6. Frontal lamina strongly grooved for entire length 7 

Frontal lamina with ventral surface flat 8 

7. Pleotelson with medial row of small tubercles flanked by row of larger 

tubercles in posterior half mexicana 

Pleotelson lacking rows of tubercles in posterior half antillensis 

8. Frontal lamina distinctly bell shaped; 8 cephalon with two pairs of 

tubercles and antennular bases not tuberculate quadricornis 

Frontal lamina, and 6 cephalon and antennules not as above 9 

9. Frontal lamina anteriorly broadly rounded, length 1.5 or less times 

basal width; 6 cephalon with two pairs of tubercles and basal 

antennular article each with tubercle sexticornis 

Frontal lamina anteriorly narrowly rounded, length about twice basal 

width; 8 cephalon with three tubercles, tubercles lacking on 

antennular bases tricornis tricornis

162 FLABELL1FERA • G0RALLAN1DAE 

Excorallana antillensis (Hansen, 1890) - t. ar- '

Excorallana oculata 163 

Figure 74. Excorallana antillensis: A, 6; B, frontal lamina; C, maxilla 2; D, 

maxilliped. Excorallana berbicensis: E, 6\ F, frontal lamina. 

Excorallana oculata (Hansen, 1890) 

Figure 75A,B 

DIAGNOSIS 6 6.9 mm, 9 8.5 mm. Eyes contiguous. Cephalon unorna- 

mented. Frontal lamina slender, linguiform, widest posteriorly. Pleonites 3-5 

each with slightly hollowed middorsal area containing strong flattened tuber 

cle; pleonite 5 with two strong flanking tubercles; pleotelson basally with low 

median ridge and two strong submedian tubercles; short strong spines in two 

roughly triangular submedian patches; lateral incisions present; apex nar 

rowly rounded. 

RECORDS Bahamas; Cuba, Puerto Rico, 40 m; Barbados. 

Brazil.

J 

Figure 75. Excorallana oculata: A, d; B> frontal lamina. Excorallana quadricomis: C, 

d; Dy frontal lamina. Excorallana sexticomis: Ey d; F, frontal lamina. Excorallana 

tricomis tricomis: G, d; H, frontal lamina. Excorallana warmingi: I, d; J, frontal 

lamina. 

H

Excorallana quadricornis (Hansen, 1890) 

Figures 75C,D; 76A-C 

Excorallana tricornis tricornis 165 

DIAGNOSIS 6 13.2 mm, 9 12.1 mm. Eyes well separated. Cephalon in 6 

with two pairs of tubercles, anterior pair connected by low rounded ridge, 

posterior pair situated mesial to eyes. Frontal lamina bell shaped, broadest 

posteriorly. Pereonite 1 with submedian pair of low tubercles. Posterior mar 

gin of pereonite 7 and pleonites faintly tuberculate. Pleotelson with two sub- 

median raised areas bearing short spines; lateral margins with incision; few 

low tubercles basally. 

RECORDS Bermuda; St. Thomas, U.S. Virgin Islands; Jamaica, intertidal 

in grassflats and between mangrove roots; Martinique; Belize; Venezuela. 

Excorallana sexticornis (Richardson, 1901) 

Figures 75E,F; 76D-F 

DIAGNOSIS 6 7.9 mm, ovigerous 9 6.9—8.3 mm. Eyes well separated. Basal 

antennular peduncular article in 6 with short anterodorsally directed tuber 

cle. Cephalon with two pairs of prominent tubercles, anterior pair shorter 

than posterior pair, latter situated mesial to eyes. Frontal lamina, length less 

than twice width, sides parallel to faintly converging anteriorly, apically 

broadly rounded. Posterior margins of pleonites 2—5 with low rounded tuber 

cles, those near middorsal line largest. Pleotelson with two basal submedian 

tubercles, numerous scattered dorsal spines, lateral margins with incision, 

apex narrowly rounded. 

RECORDS Key West, Florida; Cuba; Puerto Rico; Twin Cays, Belize, shal 

low infratidal from dead mangrove wood. 

Excorallana tricornis tricornis (Hansen, 1890) 

Figures 75G,H; 77 

DIAGNOSIS 6 8.2 mm, $ 9.9 mm. Basal antennular article narrow, not 

dilated. Cephalon in 6 with one median and two dorsal "horns." Eyes large, 

well separated. Frontal lamina between two and three times longer than 

basal width, sides subparallel, anteriorly rounded to subacute. Pereon 

smooth. Margins of lateral incision of pleotelson separated by gap; anterior 

margin of incision lined with short spines; scattered short spines on dorsum 

of telson, but especially concentrated in two submedian patches. Uropodal 

exopod, length 2.3-2.5 times width; apical notch nearly symmetrical.


Figure 76. Excorallana quadricornis: A, cephalon and pereonite 1 6; B, ventral 

cephalon; C, frontal lamina enlarged. Excorallana sexticornis: D, cephalon and 

o 

pereonite 1 8; E, cephalon 9; F, ventral cephalon. 

RECORDS Turks and Caicos Islands, 1 m; St. Thomas and St. Croix, U.S. 

Virgin Islands, 48—55 m; Cuba; Puerto Rico, on gills of rays Aetobatus narinari 

and Dasyatis americana, and on squirrel fish; Belize, intertidal to 15.2 m, in 

tertid bble sediments in Syringodium and Thalassia 

g r a s s b e d s , o n b r o w n a l g a T u r b i n a r i a , o n M a d r a c i s s p . s p o n g e , o n A g a r i c i a s p 

c o r a l ; G u l f o f M e x i c o .

Excorallana warmingii 167 

F i g u r e 7 7 . Excorallana tricornis tricornis: A , c e p h a l o n a n d p e r e o n i t e 1 6 ; B , v e n t r a l 

c e p h a l o n ; C, p l e o t e l s o n a n d u r o p o d s . 

R E M A R K S T h e s u b s p e c i e s E x c o r a l l a n a t r i c o r n i s o c c i d e n t a l i s R i c h a r d s o n , 1 9 0 5 a , 

f r o m s o u t h e r n C a l i f o r n i a , d i f f e r s f r o m t h e G u l f a n d C a r i b b e a n s u b s p e c i e s i n 

l a c k i n g a g a p b e t w e e n t h e m a r g i n s o f t h e p l e o t e l s o n i c i n c i s i o n , a n d i n h a v i n g 

a r e l a t i v e l y w i d e r u r o p o d a l e x o p o d w h i c h s h o w s a d i s t i n c t l y a s y m m e t r i c a l 

a p i c a l n o t c h . 

E x c o r a l l a n a w a r m i n g i i ( H a n s e n , 1 8 9 0 ) 

F i g u r e 7 5 1 , J 

D I A G N O S I S ( J 9 . 7 m m , 9 1 2 . 0 m m . C e p h a l o n u n o r n a m e n t e d . E y e s c o n 

t i g u o u s , o c c u p y i n g m o s t o f d o r s a l s u r f a c e o f h e a d . P o s t e r i o r m a r g i n s o f 

p l e o n i t e s v e r y f a i n t l y t u b e r c u l a t e . F r o n t a l l a m i n a , l e n g t h s l i g h t l y m o r e t h a n 

t w i c e b a s a l w i d t h , t a p e r i n g a n t e r i o r l y t o r o u n d e d a p e x . P l e o t e l s o n u n o r n a 

m e n t e d e x c e p t f o r t w o f a i n t s u b m e d i a n t u b e r c l e s b a s a l l y ; l a t e r a l i n c i s i o n s 

l a c k i n g ; a p e x b r o a d l y r o u n d e d , w i t h f i v e l o w b u t d i s t i n c t m a r g i n a l t e e t h . 

*

168 FLABELLIFERA • CORALLAN1DAE 

Figure 78. Nalicora rapax: A, 9; B> maxilla 1; C> maxilla 2; D, maxilliped. 

RECORDS Bahamas; between Cuba and the Yucatan Pensinsula; Puerto 

Rico. 

Off Brazil near Rio de Janeiro. 

Nalicora Moore, 1901 

DIAGNOSIS Maxilla 1 exopod a single strongly falcate distal spine with 

knoblike mesial process, and basal caplike convex papilla-bearing structure. 

Maxilla 2 of four articles, distal article slender. Maxillipedal palp of five 

articles; endite lacking.

Nalicora rapax Moore, 1901 

Figure 78 

FLABELL1FERA • CYMOTHOIDAE 169 

DIAGNOSIS 6 6.9 mm, ovigerous 9 10.0 mm. Eyes well developed. Frontal 

lamina basally slender, widening anteriorly, apex subacute. Posterior half of 

body bearing numerous scattered stiff setae. Pereonites 4-7 with row of low 

rounded tubercles near posterior margin. Posterior margins of pleonites 3—5 

faintly tuberculate, more noticeable in 6. Pleotelson wider than long; lateral 

margins faintly sinuous; apex rounded. 

RECORDS Florida Keys, 55 m; Puerto Rico, 50-150 m; Gulf of Mexico off 

Florida, 37-73 m. 

Family Cymothoidae Leach, 1818 

DIAGNOSIS Antennules and antennae reduced, no clear distinction between 

peduncles and flagella. Mandibular palp of three articles. Maxilla 1 with four 

terminal spines. Maxilla 2 apically bilobed, armed with several spines. Max- 

illipedal palp of two articles, terminal article bearing hooks. All seven pairs of 

pereopods prehensile, ending in strongly hooked dactyli. Pleopods lacking 

marginal setae in adults. 

REMARKS The cymothoids are exclusively ectoparasites on marine, fresh 

water, and brackish-water fishes. Most cymothoids occur in shallow water, 

mainly in tropical and subtropical areas. The position of attachment on the 

host (externally, in the buccal cavity, or in the gill chamber) is usually genus- 

or species-specific. The body of gill parasites is often asymmetrical, being 

slightly twisted, perhaps an effect of the position on the host. The mouthparts 

are highly adapted for the parasitic mode of life, while all seven pairs of 

pereopods are strongly prehensile. The posterior pereopods of some genera 

have the basal article expanded and carinate, allowing for increased mus 

culature. The secretion of anticoagulants in the juvenile stages further aids 

the blood-feeding habit. The surface area of the pleopods is often increased 

by the development of lobes on the bases or the lamellae, providing an in 

creased respiratory ability. 

The post-mancal juvenile stages (sometimes referred to as the aegathoid 

stage) have large eyes, and highly setose pleopods for active swimming. The 

juveniles will attach themselves indiscriminantly to any convenient fish host, 

but eventually attach to the preferred host-species. The juvenile then de 

velops into a functional male, losing the swimming setae of the pleopods. 

Both juveniles and males feed actively, drawing blood from the host fish. The

170 FLABELLIFERA • GYMOTHOIDAE 

Key to genera of Cymothoidae 

1. Antennule broader and usually longer than antenna; cephalon very 

weakly sunk into pcreonite 1 2 

Antennule not broader or longer than antenna; cephalon distinctly 

immersed in, or not at all immersed in pereonite 1 4 

2. Bases of antennules widely separated 3 

Bases of antennules contiguous Glossobius 

3. Body curved to one side; pleonitc 1 extended laterally more on one side 

than on other Mothocya 

Body rarely curved to one side; pleonitc 1 extended equally on each 

side Renocila 

4. Pereonites and coxal plates 4-7 strongly expanded on one side only 

Agarna 

No pereonites or coxal plates strongly expanded 5 

5. Cephalon not immersed in pereonite 1; posterior margin of cephalon 

trisinuate 6 

Cephalon to some degree immersed in pereonite 1; posterior margin of 

cephalon not trisinuate 7 

6. Posterolateral angles of pereonites 2-6 not produced; coxal plates short, 

rarely reaching posterior margin of their pereonites Anilocra 

Posterolateral angles of pereonites 2-6 posteriorly increasingly 

produced; coxal plates usually reaching to posterior margin of their 

pereonites Nerocila 

7. Basal antennular articles expanded and contiguous Ceratothoa 

Basal antennular articles expanded but not contiguous, or basal 

antennular articles neither expanded nor contiguous 8 

8. Basal antennular articles expanded but not contiguous Kuna 

Basal antennular articles neither expanded nor contiguous 9 

9. Plconal margins continuous with pereonal margins, pleon not abruptly 

narrowed, only weakly immersed in pereonite 7 Lironeca 

Pleon to some degree narrower than pereon; pleon usually deeply 

immersed in pereonite 7 Cymothoa

FLABELLIFERA • CYMOTIIOIDAE 171 

male eventually becomes a female (all cymothoids are protandrous) should a 

female not already be present. In some species, the female is nonfeeding. In 

those species which settle either in the mouth cavity or gill chamber of the 

host, integumental pigment is frequently lost, and the eyes become reduced. 

Given the highly variable morphology of the cymothoids, in part imposed 

by the parasitic mode of life, and the existence of polymorphism and possible 

sibling species, the taxonomy of this family demands the examination of large 

numbers of specimens. As a further aid to identification, Table 3 is provided, 

giving host species, parasite, and site of attachment. 

TABLE 3. GYMOTHOID PARASITES FROM THE CARIBBEAN AREA, LISTED BY FISH 

HOST SPECIES 

Fish host Cymothoid parasite Site of attachment 

Abudefduf saxatilis 

Acanthurus bahianus 

Acanthurus chirurgus 

Alutera schoepji 

Anchoa lamprotaenia 

Apogon lachneri 

Apogon maculatus 

Apogon townsendi 

Ariusfelis 

Astrapogon stellatus 

Batrachoides surinamensis 

Caranx hippos 

Caranx latus 

Caranx ruber 

Caranx sp. 

Chaetodipterus faber 

Chaetodon capistratus 

Chaetodon ocellatus 

Chaetodon sedentarius 

Chaetodon striatus 

Anilocra abudefduji 

Kuna insularis 

Anilocra acanthuri 

Anilocra acanthuri 

Nerocila acuminata 

Lironeca tenuistylis 

Mothocya bohlkeorum 

Renocila colini 

Renocila colini 




Cymothoa oestrum 





Anilocra chaetodontis 




beneath eye 

gill chamber 

9 at base of pectoral fin; 

immature on or near 

pectoral or pelvic fin 

9 at base of pectoral fin; 

immature on or near 

pectoral or pelvic fin 

on or at base of fin 

posterior to pectoral fin 

in gill chamber 

next to dorsal fin 





inside mouth 

inside mouth 

inside mouth 

inside mouth 


beneath eye 

beneath eye 

beneath eye 

beneath eye 

(continued)

172 



Chilomycterus schoepfi 

Chromis cyaneus 

Chromis multilineatus 

Cynoscion nebulosus 

Cynoscion sp. 

Epinephelus cruentatus 

Epinephelus Julvus 

Epinephelus guttatus 

Epinephelus itajara 

Epinephelus sp. 

Exocoetus spp. 

Genes rhombeus 

Haemulon aurolineatum 

Haemulon carbonarium 

Haemulon chrysargyreum 

Haemulon Jlavolineatum 

Haemulon macrostomum 

Haemulon plumieri 

Haemulon sciurus 


Anilocra chromis 

Anilocra chromis 

Cymothoa excisa 


Anilocra haemuli 





Glossobius impressus 

Lironeca redmanni 








Hemirhamphus brasiliensis Glossobius hemirhamphi 

Hirundichthys speculifer 

Holacanthus tricolor 

Holocentrus ascensionis 

Glossobius impressus 

Anilocra holacanthi 

Anilocra holocentri 

Hyporhamphus unifasciatus Mothocya nana 

Leiostomus xanthurus 

Lepiosteus spatula 

Lutjanus analis 

Lutjanus mahogoni 

Lutjanus synagris 

Megalops atlanticus 

Monacanthus ciliatus 

Mugil cephalus 

Myripristis jacobus 

Ocyurus chrysurus 

Orthopristis chrysoptera 











Anilocra myripristi 




beneath eye 

beneath eye 

inside mouth 

inside mouth 

beneath eye 

beneath eye 

beneath eye 



inside mouth 


beneath eye 

beneath eye 

beneath eye 

beneath eye 

beneath eye 

beneath eye 

beneath eye 

inside mouth 

inside mouth 

beneath eye 

? between eyes, 8 and 

immature beneath 

eye 



inside mouth 


on at base of fin 

inside mouth 

inside mouth 

inside mouth 

inside mouth 



9 between eyes, imma 

ture beneath eye 

inside mouth 

inside mouth

Agarna cumulus 173 


Orthopristis ruber 

Paranthias Jurcifer 

Phaeoptyx conklini 

Phaeoptyx pigmentaria 

Pogonias cromis 

Pomacentrus partitas 

Priacanthus arenatus 

Scomberomorus cavalla 

Scomberomorus maculatus 

Scomberomorus regalis 

Selar crumenophthalmus 

Serranus tigrinus 

Sphoeroides maculatus 

Synodus foetens 






Anilocra partiti 






Renocila bowmani 

Renocila waldneri 



Agarna Schioedte and Meinert, 1883 

beneath eye 

beneath eye 




beneath eye 

inside nouth 




inside mouth 




inside mouth 

DIAGNOSIS Cephalon with posterior margin not trilobed; immersed in per- 

eonite 1. Antennular bases contiguous. Pereonites 4-7 on one side flattened 

and expanded; coxal plates of pereopods 4-7 also expanded and flattened 

but generally hidden by lateral expansion of pereonites. Bases of posterior 

three pereopods with well-formed carinae. Pleonites 1 and 2 immersed in 

pereonite 7; pleonites 2-5 with free fingerlike lateral margins. 

Agarna cumulus (Haller, 1880) 

Figure 79 

DIAGNOSIS 9 18 mm. Eyes present, indistinct. Pereon strongly "humped" 

dorsally. Uropod about { fo length of pleotelson; uropodal exopod slightly 

longer, and twice width of endopod. Pleotelson triangular, length 3 A basal 

width, apex rounded. 

RECORDS No host recorded: Key West, Florida.


Figure 79. Agarna cumulus: A, 9, dorsal view; By 9, ventral view, coxal plates 

stippled. 

Anilocra Leach, 1818 

DIAGNOSIS Cephalon usually narrowed anteriorly to triangular apex folded 

ventrally between bases of antennules; posterior margin trilobed; not im 

mersed, or only weakly immersed in pereonite 1. Coxal plates small, com 

pact, not reaching level of posterior margin of their respective pereonites. 

Pereopods increasing in length posteriorly, pereopod 7 often markedly longer 

than 6. Pleon not immersed or only slightly immersed in pereonite 7. 

Pleopods 3—5 often formed into deep pockets or pleats. Uropods often ex 

tending beyond pleotelsonic apex. 

REMARKS Williams and Williams (1981) have provided a comprehensive 

treatment of this genus and nine of its species in the West Indies. Table 1 in 

this latter paper provides characters for separating these nine species. This 

table also indicates that for each species, the site of attachment of the adult to 

the host fish is specific, with six species attaching under the eye of the host.

Key to species of Anilocra 

Anilocra abudefdufi 175 

1. Pereopods 2-4 with swelling on outer margin of dactylus 2 

Pereopods 2-4 lacking swelling on outer margin of dactylus 5 

2. Body axis distorted by more than 10° holacanthi 

Body axis distorted by less than 5° 3 

3. Dactylus of pereopod 7 longer than propodus partiti 

Dactylus of pereopod 7 shorter than propodus 4 

4. Posteroventral angle of pereonite 7 overlapping pleonite 1 only 

Posteroventral angle of pereonite 7 overlapping pleonites 1 and 2 

abudefdufi 

chaetodontis 

5. Posteroventral angle of pereonite 7 produced 6 

Posteroventral angle of pereonite 7 not produced 7 

6. Uropod reaching posterior margin of pleotelson myripristis 

Uropod not reaching posterior margin of pleotelson haemuli 

7. Posteroventral angle of pereonite 7 overlapping pleonite 1 . . . . holocentri 

Posteroventral angle of pereonite 7 not overlapping pleonite 1 8 

8. Uropod reaching posterior margin of pleotelson acanthuri 

Uropod not reaching posterior margin of pleotelson chromis 

Anilocra abudefdufi Williams and Williams, 1981 

Figure 80A-C 

DIAGNOSIS Ovigerous 9 19.0-31.0 mm, 6 7.0-8.5 mm. Pereopods 2-4 

with swelling on outer margin of dactylus. Posteroventral angle of pereonite 6 

slightly produced, of pereonite 7 more produced, overlapping pleonite 1. 

Uropodal endopod variable, not reaching, to extending well beyond, apex of 

exopod. Color: upper lateral half to three-fourths of dorsal surface of 9 when 

attached to host is dark brown; rest of dorsal surface light brown to yellow. 

Attaching beneath eye of host.


Figure 80. Anilocra abudefdufi: A, 9, lateral view; B, pereopod 3; C, pereopod 7. 

Anilocra acanthuri: D, pleotelson and uropods. Anilocra chaetodontis: E, 9, lateral 

view. Anilocra chromis: F, pleotelson and uropods. 

Anilocra acanthuri Williams and Williams, 1981 

Figure 80D 

DIAGNOSIS Ovigerous 9 29.0-40.0 mm, 6 4.0-8.0 mm. Pereopods 2-4 

without swelling on outer margin of dactylus. Posteroventral angles of per- 

eonites not produced. Uropod not reaching posterior margin of pleotelson. 

Endopod of uropod variable, not reaching, to extending well beyond, apex of 

exopod. Color: dorsal surface of 9 black to lead gray, ventral surface gray. 

Attaching under pectoral fin of host. 

RECORDS Doctorfish Acanthurus chirurgus: Florida Keys; Bahamas; Puerto 

Rico; U.S. Virgin Islands. Ocean surgeon Acanthurus bahianus: Florida Keys; 

Bahamas; Cuba; Jamaica; Dominican Republic; Puerto Rico; U.S. Virgin 

Islands.

Anilocra chaetodontis Williams and Williams, 1981 

Figure 80E 

Anilocra haemuli 177 

DIAGNOSIS Ovigerous 9 18-28 mm, 6 4-5 mm. Pereopods 2-4 with swell 

ing on outer margin of dactylus, Posteroventral angles of pereonites 4—7 be 

coming progressively produced, that of pereonite 7 overlapping pleonite 2. 

Uropod not reaching posterior margin of pleotelson; uropodal endopod ex 

tending beyond apex of exopod. Pleotelson as wide as long to slightly wider 

than long. Color: dorsal surface of 9 black to lead gray, ventral surface gray. 

Attaching beneath eye of host. 

RECORDS Foureye butterflyfish Chaetodon capistratus: Bahamas; Puerto Rico; 

British and U.S. Virgin Islands. Banded butterflyfish Chaetodon striatus: 

Bahamas; Puerto Rico; British Virgin Islands. Spotfin butterflyfish Chaetodon 

ocellatus: Bahamas; Puerto Rico; U.S. Virgin Islands. Reef butterflyfish 

Chaetodon sedentarius: Puerto Rico. 

Anilocra chromis Williams and Williams, 1981 

Figure 80F 

DIAGNOSIS Ovigerous 9 16-28 mm, 8 4-9 mm. Pereopods 2-4 lacking 

swelling on outer margin of dactylus. Posteroventral angles of pereonites not 

produced. Uropod extending beyond posterior margin of pleotelson; uropo 

dal endopod not reaching beyond exopod. Color: upper lateral one-fourth to 

two-thirds of dorsal surface of 9 when attached is dark gray, shading to off- 

white lower lateral area. Attaching beneath eye of host. 

RECORDS Brown chromis Chromis multilineatus: Puerto Rico; British and 

U.S. Virgin Islands. Blue chromis Chromis cyaneus: Bahamas; Dominican Re 

public. No host recorded: Anguilla. 

Anilocra haemuli Williams and Williams, 1981 

Figure 81A,B 

DIAGNOSIS Ovigerous 9 21-40 mm, 6 7 mm. Body axis distorted less than 

5°. Pereopods 2—4 lacking swelling on outer margin of dactylus. Postero 

ventral angle of pereonites 6 and 7 produced, latter overlapping pleonite 1. 

Uropod not reaching posterior margin of pleotelson; uropodal endopod 

reaching beyond apex of exopod. Color: dorsal surface of 9 yellow to light 

brown. Attaching beneath eye of host.


Figure 81. Anilocra haemuli: A, 9, dorsal view; B, 9, lateral view. Anilocra 

holacanthi: C, 9. Anilocra holocentri: D, 9. Anilocra myripristis: E, pleotelson and 

uropods. Anilocra partiti: F, 9; G, pereopod 7. 

RECORDS French grunt Haemulon Jlavolineatum: Florida Keys; Puerto Rico; 

British and U.S. Virgin Islands. Tomtate Haemulon aurolineatum: Jamaica; 

Puerto Rico. Smallmouth grunt Haemulon chrysargyreum: Puerto Rico; U.S. 

Virgin Islands. Caesar grunt Haemulon carbonarium: Puerto Rico; U.S. Virgin

Anilocra holocentri 179 

Islands. Spanish grunt Haemulon macrostomum: Puerto Rico. White grunt 

Haemulon plumieri: Florida Keys; Yucatan Peninsula. Bluestriped grunt 

Haemulon sciurus: Florida Keys. Cora cora Orthopristis ruber: Margarita Island, 

Venezuela. Coney Epinephelus Julvus: Bahamas; Dominican Republic; Puerto 

Rico; U.S. Virgin Islands; Guadeloupe. Red hind Epinephelus guttatus: Puerto 

Rico; British and U.S. Virgin Islands. Graysby Epinephelus cruentatus: 

Bahamas; Dominican Republic; U.S. Virgin Islands. Creole-fish Paranthias 

jurcifer: Dominican Republic; Puerto Rico; Colombia. No host recorded: 

Cuba; Jamaica; Dominica; Barbados; Venezuela; Brazil. 

Anilocra holacanthi Williams and Williams, 1981 

Figure 81C 

DIAGNOSIS Ovigerous 9 21-33 mm, 6 4-7 mm. Body axis distorted by 

more than 10°. Pereopods 2-4 with swelling on outer margin of dactylus. 

Posteroventral angles of pereonites 5—7 progressively more produced, that of 

pereonite 7 overlapping pleonite 1. Uropod not reaching posterior margin of 

pleotelson; uropodal endopod reaching beyond apex of exopod. Color: dorsal 

surface of 9 black to lead gray. Attaching beneath eye of host. 

RECORDS Rock beauty Holacanthus tricolor: Bahamas; Jamaica; Dominican 

Republic; Puerto Rico; British and U.S. Virgin Islands. 

Anilocra holocentri Williams and Williams, 1981 

Figure 81D 

DIAGNOSIS Ovigerous 9 32-46 mm, 6 5-9 mm. Body axis distorted less 

than 5°. Pereopods 2—4 lacking swelling on outer margin of dactylus. 

Posteroventral angle of pereonite 7 produced, overlapping pleonite 1. 

Uropod not reaching posterior margin of pleotelson; uropodal endopod 

reaching beyond apex of exopod. Color: dorsal surface of 9 dark brown, 

ventral surface light brown. 9 attaching between eyes of host; 6 or transi 

tional stage beneath eye. 

RECORDS Squirrelfish Holocentrus ascensionis: Puerto Rico; U.S. Virgin 

Islands. 

No host recorded: Patagonia, Straits of Magellan.

180 FLABELLIFERA • CYMOTHOIDAE 

Anilocra myripristis Williams and Williams, 1981 

Figure 81E 

DIAGNOSIS Ovigerous 9 29-40 mm, 6 6-7 mm. Body axis distorted less 

than 5°. Pereopods 2-4 lacking swellings on outer margin of dactylus. 

Posteroventral angle of pereonites 6 and 7 produced, latter overlapping 

pleonite 1. Uropod reaching beyond posterior margin of pleotelson; uropodal 

endopod reaching beyond apex of exopod. Color: dorsal surface of 9 light 

reddish brown, ventral surface yellow. 9 attaching between eyes of host; 

immature or transitional forms sometimes beneath eve. 

4 

RECORDS Blackbar soldierfish Myripristis jacobus: Bahamas; Dominican Re 

public; Puerto Rico. 

Anilocra partiti Williams and Williams, 1981 

Figure 81F,G 

DIAGNOSIS Ovigerous 9 12-16 mm, transitional 7.6-9.0 mm. Body axis 

distorted less than 5°. Pereopods 2-4 with swelling on outer margin of dac 

tylus. Pereopod 7 with dactylus longer than propodus. Posteroventral angle 

of pereonite 7 produced, overlapping pleonite 1. Uropod not reaching pos 

terior margin of pleotelson; uropodal endopod not reaching apex of exopod. 

Color: dorsal surface black to slate gray. Attaching beneath eye of host. 

RECORDS Bicolor damselfish Pomacentrus partitas: Jamaica. 

Ceratothoa Dana, 1852 

DIAGNOSIS Cephalon more or less immersed in pereonite 1, posterior mar 

gin not trisinuate. Bases of antennules expanded, contiguous. Coxal plates 

compact; anterior plates not extending beyond posterior margins of their 

respective pereonites; posterior coxal plates may or may not be produced 

beyond the posterior margins of the pereonites. Anterior pleonites narrowed, 

immersed in pereonite 7, Copulatory stylet lacking on pleopod 2 of 6 of some 

species. 

Ceratothoa deplanata Bovallius, 1885 

Figure 82A 

DIAGNOSIS 9 18 mm. Cephalon subtriangular, anterior margin rounded. 

Pereopods 4—7 with strongly carinate bases. Uropod reaching or extending

Ceratothoa deplanata 181 

Figure 82. A, Ceratothoa deplanata (from Bovallius, 1885); B, Cymothoa caraibica; C, 

Cymothoa excisa; D, Cymothoa oestrum. 

slightly beyond posterior margin of pleotelson; rami subequal in length and 

width. Pleotelson basally wider than long, posterior margin broadly 

rounded. Color: bright yellow. 

RECORDS Haiti, host not recorded.

182 FLABELUFERA • GYMOTHOIDAE 

Cymothoa Fabricius, 1793 

DIAGNOSIS Body usually not distorted. Cephalon with posterior margin not 

trilobed; more or less immersed in pereonite 1; latter with anterolateral cor 

ners produced to embrace cephalon. Bases of antennules not expanded, well 

separated. Anterior coxal plates not reaching posterior borders of their re 

spective pereonites, posterior coxal plates nearly reaching or extending be 

yond posterior borders of pereonites. Pleon narrower than, and immersed in 

pereonite 7. Pleonites increasing in length and width posteriorly. 

Key to species of Cymothoa 

1. Anterolateral angles of pereonite 1 reaching to half length of cephalon 

or less; eyes or traces of eyes present 2 

Anterolateral angles of pereonite 1 broad, reaching to anterior margin 

of cephalon; eyes absent oestrum 

2. Anterolateral angles of pereonite 1 narrow, subacute excisa 

Anterolateral angles of pereonite 1 broad, rounded caraibica 

Cymothoa caraibica Bovallius, 1885 

Figure 82B 

DIAGNOSIS 9 17 mm, 6 12-16 mm. Anterior margin of cephalon broadly 

rounded. Eyes large, distinct. Broadly rounded anterolateral angles of per 

eonite 1 reaching to about midlength of cephalon. Bases of pereopods 4—7 

with strong, rounded carina. Uropodal rami subequal in length, equal to 

peduncle in length. Pleotelson width about twice length, posterolateral mar 

gin broadly rounded. 

RECORDS Puerto Rico; Gulf of Mexico. 

Cymothoa excisa Perty, 1833 

Figure 82C 

DIAGNOSIS Ovigerous 9 20-24 mm. Anterior margin of cephalon in dorsal 

view truncate to slightly excavate; eyes small, indistinct. Anterolateral angles 

of pereonite 1 narrowly rounded to subacute, reaching anteriorly to about 

midlength of cephalon. Pereopods 4—7 with high rounded carina on basis.

Glossobius 183 

Uropods hardly reaching halfway along lateral margin of pleotelson; exopod 

slightly longer than endopod. Pleotelson about twice wider than long; 

broadly rounded and somewhat bilobed. 

RECORDS Yellowtail snapper Ocyurus chrysurus: Yucatan Peninsula, Mexico; 

Carrie Bow Cay, Belize; Margarita Island, Venezuela; Panama. Mutton 

snapper Lutjanus analis: Yucatan Peninsula, Mexico; Panama. Lane snapper 

Lutjanus synagris: Panama. Mahogany snapper Lutjanus mahogoni: Panama. 

Pigfish Orthopristis chrysoptera: Florida, Gulf of Mexico. Spot Leiostomus 

xanthurus: Texas, Gulf of Mexico. Spotted seatrout Cynoscion nebulosus: Texas, 

Gulf of Mexico. Inshore lizardfish Synodusfoetens: Texas, Gulf of Mexico. No 

host recorded: Massachusetts; South Carolina; Georgia; Florida Keys; 

Bahamas; Cuba; Trinidad; Brazil. 

Cymothoa oestrum (Linnaeus, 1793) 

Figure 82D 

DIAGNOSIS Ovigerous 9 38 mm. Cephalon in dorsal view with ante 

rolateral angles rounded, anterior margin slightly excavate; eyes absent. An 

terolateral angles of pereonite 1 expanded, broadly rounded, reaching to 

level of anterior margin of cephalon. Pereonites 4—7 with high rounded 

carina on basis. Uropod reaching posteriorly beyond midlength of 

pleotelson; exopod slightly longer than endopod. Pleotelson length slightly 

more than half basal width. 

RECORDS Bigeye scad Selar crumenophthalmus: Bermuda; U.S. Virgin Is 

lands. Bigeye Priacanthus arenatus: Bermuda. Bar jack Caranx ruber: Florida 

Keys; Carrie Bow Cay, Belize. Horse-eye jack Caranx latus: Bahamas; Bar 

bados. Crevalle jack Caranx hippos: Venezuela. Jack Caranx sp.: Jamaica; Cu 

rasao. Hind Epinephelus sp.: Grenada. Parrotfish: Jamaica. Seatrout Cynoscion 

sp.: Panama. Tarpon Megalops atlantica: Texas, Gulf of Mexico. No host re 

corded: Honduras; Haiti. 

Glossobius Schioedte and Meinert, 1883 

DIAGNOSIS Cephalon not immersed in pereonite 1; excavate on either side 

in anterior half, forming broad and anteriorly rounded median area; anten 

nae fitting into excavate areas. Bases of antennules contiguous, expanded. 

Antennules broader and longer than antennae. Bases of pereopods 4—7 with 

posterior margin expanded and flattened. Pleonites 1-3 immersed in per 

eonite 7.


Key to species of Glossobius 

1. Coxal plates of perconites 1 and 2 anteroventrally protruding impressus 

Coxal plates of pereonites 1 and 2 close to body, not protruding 

Glossobius hemiramphi Williams and Williams, 1985a 

Figure 83A 

hemirhamphi 

DIAGNOSIS Ovigerous 9 27 mm. Eyes small but distinct. Cephalon pointed 

anteriorly. Fused coxa of pereonite 1 and free coxa of pereonite 2 carinate but 

not protruding. Coxa of pereonite 7 semicircular in dorsal view. Pleotelson 

with middorsal length more than half basal width; lateral margins somewhat 

tapered; posterior margin variable, sinuate or excavate. Uropods reaching to 

or slightly beyond posterior pleotelsonic margin; rami subequal in length, 

exopod slightly broader than endopod. 

RECORDS Ballyhoo Hemiramphus brasiliensis; Puerto Rico. 

Glossobius impressus (Say, 1818) 

Figure 83B 

DIAGNOSIS Ovigerous 9 33 mm. Eyes small but distinct. Cephalon 

rounded anteriorly. Fused coxal plate of pereonite 1 and distinct coxal plate 

of pereonite 2 protruding strongly in oblique anteroventral direction. Uropod 

reaching to posterior half of pleotelson; exopod shorter and narrower than 

endopod. Pleotelson basal width twice length, posteriorly broadly bilobed. 

Attaching inside mouth of host. 

RECORDS Flyingfish Exocoetus spp.: Rio de Janeiro, Brazil; North Atlantic, 

especially in the Gulf Stream. 

Mirrorwing flyingfish Hirundichthys speculifer: North Atlantic. No host rec 

ord: Senegal, West Africa. 

Kuna Williams and Williams, 1986 

DIAGNOSIS Cephalon somewhat immersed in pereonite 1. Anterior margin 

of pereonite 1 not trisinuate. Number of articles in antennules and antennae

Kuna 185 

Figure 83. A, Glossobius hemiramphi; B, Glossobius impressus; C, Kuna insularis; D, 

Lironeca redmani; E, Lironeca tenuistylis. 

reduced. Antennule somewhat expanded; basal article expanded but not 

contiguous. Copulatory stylet present on pleopods 1-3 in 6. Pleonites dor- 

sally strongly convex, not immersed in pereonite 7.


Kuna insularis (Williams and Williams, 1985b) 

Figure 83C 

DIAGNOSIS Ovigerous 9 11.1-17.2 mm, 6 4.2-8.7 mm, transitional 9.6- 

9.8 mm. Antennules and antennae consisting of four articles each. Uropods 

short, not reaching posterior margin of pleotelson. Clavate eopulatory stylet 

present on pleopods 1—3 in 6. Pleotelson basally broader than long, pos 

terior margin broadly rounded. 

RECORDS Sergeant major Abudefdufsaxatilis: Carrie Bow Cay, Belize; Cura 

sao; Panama. 

Lironeca Leach, 1818 

DIAGNOSIS Cephalon weakly to deeply immersed in pereonite 1; posterior 

border rarely trisinuate. Bases of antennules not expanded, well separated. 

Posterior pereopods with carinae on bases in 6, carinae present or absent in 

9. Pleonites subequal in width; pleonites 1 and 2 rarely narrowed and 

weakly to moderately immersed in pereonite 7. Pleopods highly folded, and 

with lamellar or digitiform accessory gills in some species. 

Key to species of Lironeca 

1. Uropodal endopod about twice longer than wide; pleon somewhat 

immersed in pereon redmanni 

Uropodal endopod about three times longer than wide; pleon barely 

immersed in pereon tenuistylis 

Lironeca redmanni Leach, 1818 

Figure 83D 

DIAGNOSIS Ovigerous 9 19.5-25.0 mm. Cephalon barely immersed in per 

eonite 1. Pleon somewhat immersed in pereon, but lateral margins of pleonite 

1 free. Pleotelson basally wider than long. Uropodal rami reaching well be 

yond posterior margin of pleotelson; exopod longer than endopod, both rami 

somewhat broad, endopod about twice longer than wide. Attaching to gills of 

host.

Mothocya 187 

RECORDS New Jersey to Florida; gills of kingfish, Jamaica; Cuba; St. 

Christopher; Spanish mackerel Scomberomorus maculatus and cero Scomberomorus 

regalisy Puerto Rico; king mackerel Scomberomorus cavalla, Colombia; Genes rho- 

mbeus, Panama; spot Leiostomus xanthurus, Gulf of Mexico. 

Brazil. 

Lironeca tenuistylis (Richardson, 1912b) 

Figure 83E 

DIAGNOSIS 9 13 mm. Cephalon barely immersed in pereonite 1. Uropodal 

rami reaching beyond rounded posterior margin of pleotelson; exopod longer 

than endopod; endopod slender, about three times longer than wide. Pleonite 

1 barely immersed in pereonite 7. Pleotelson basally wider than long. Attach 

ing to host between pectoral and anal fin. 

RECORDS Longnose anchovy Anchoa lamprotaenia: Panama. 

Mothocya Costa, 1851 

DIAGNOSIS Cephalon more or less immersed in pereonite 1. Bases of anten- 

nules widely separated; antennules longer and more robust than antennae. 

Coxae nearly reaching or extending beyond posterior margin of respective 

pereonites. Pleon somewhat immersed in pereonite 7. Uropodal exopod 

longer than endopod. 

REMARKS Bruce (1986b) revised the genus Mothocya, The species of Moth 

ocya are almost entirely gill parasites on the fish families Hemiramphidae, 

Apogonidae, Belonidae, and Atherinidae. 

Key to species of Mothocya 

1. Cephalon anteriorly narrowed, slightly immersed in pereonite 1; 

pleotelson subrectangular bohlkeorum 

Cephalon anteriorly broad, deeply immersed in pereonite 1; pleotelson 

subtriangular nana


Mothocya bohlkeorum Williams and Williams, 1982 

Figure 84B 

DIAGNOSIS Ovigerous 9 7.6-8.5 mm, 6 3.7 mm. Cephalon anteriorly nar 

rowed in dorsal view, ventrally flexed, broadly rounded; slightly immersed in 

pereonite 1. Pleotelson subrectangular. Uropods extending slightly beyond 

posterior margin of pleotelson; exopod only slightly longer than endopod. 9 

lateral lobes of pleopodal peduncles not developed. Endopods of pleopods 3- 

5 with small proximomedial lobe. 

RECORDS Whitestar cardinalfish Apogon lachneri: Puerto Rico. Dusky car- 

dinalfish Phaeoptyx pigmentaria: Bahamas. Freckled cardinalfish Phaeoptyx con- 

klini: Florida Keys; Bahamas. Conchfish Astrapogon stellatus: Leeward 

Islands. 

Mothocya nana (Schioedte and Meinert, 1884) 

Figure 84A 

DIAGNOSIS Ovigerous 9 11.0-17.0 mm, 6 7.9-8.3 mm. Cephalon deeply 

immersed in pereonite 1; rostrum anteroventrally narrowly rounded. Uropo- 

dal exopod markedly longer than endopod. Pleotelson broad, with posterior 

margin rounded sufficiently to give appearance of being subtriangular. 

RECORDS Halfbeak Hyporhamphus unifasciatus: Chesapeake Bay, Maryland; 

Georgia; Florida; Colon, Panama. Halfbeak Hemiramphus bermudensis: 

Bermuda. 

Nerocila Leach, 1818 

DIAGNOSIS Body generally more depressed than in most cymothoid genera, 

rarely curved. Cephalon with anterior margin convex, narrowly rounded, or 

concave; not, or only slightly, immersed in pereonite 1. Pereonite 1 anterior 

margin trisinuate. Posterolateral angles of pereonites weakly to strongly pro 

duced, increasing in length posteriorly. Coxal plates prominent, usually al 

most reaching or extending to posterior margin of their respective pereonites. 

Juveniles and 6 usually with spines on posterior pereopods; 9 lacking these 

spines. Pleon not immersed in pereonite 7. Pleonites subequal in length; 

pleonites 1 and 2 usually produced posterolateral^. Pleopods typically with 

small lamellar accessory gills; pleopods 3-5 folded into deep pockets or 

pleats. Uropods usually extending beyond pleotelsonic apex.

Figure 84. A, Mothocya nana; By Mothocy 

acuminata; D, Nerocila acuminata f. aster. 

B


Nerocila acuminata Schioedte and Meinert, 1881 

DIAGNOSIS Ovigerous 9 16.2-19.0 mm. Cephalon with anterior margin 

convex. Posterolateral angles of all, or of posterior pereonites only, produced 

into acute or subacute angles. 

RECORDS Striped burrfish Chilomycterus schoepfi: Texas, Gulf of Mexico. 

Northern puffer Sphoeroides maculatus: New York. Striped mullet Mugil 

cephalus: Texas, Gulf of Mexico. Jewfish Epinephelus itajara: Texas, Gulf of 

Mexico. Hogfish: Bermuda. Alligator gar Lepisosteus spatula: Louisiana, Gulf 

of Mexico. Hardhead catfish Ariusfelis: Texas, Gulf of Mexico. Sawfish: Flor 

ida (Atlantic). Black drum Pogonias cromis: Texas, Gulf of Mexico. Orange 

filefish Alutera schoepfi: Texas, Gulf of Mexico. Toadfish Batrachoides sur- 

inamensis: Colon, Panama. Spot Leiostomus xanthurus: Florida, Gulf of Mexico. 

Spadefish Chaetodipterus faber: Florida, Gulf of Mexico; Virginia. Fringed fil 

efish Monacanthus ciliatus: Florida, Gulf of Mexico. No host recorded: Mas 

sachusetts; Florida Keys; Florida, Gulf of Mexico. Louisiana, Gulf of Mex 

ico. Texas, Gulf of Mexico. 

REMARKS Brusca (1981) has shown that this highly variable species occurs 

on both sides of the Isthmus of Panama, in two relatively distinct forms. 

Intergrades between the two forms do occur but are uncommon. Brusca 

(1981:159) also lists all the host-records for this species in the eastern Pacific. 

Nerocila acuminata Schioedte and Meinert, 1881, forma acuminata 

Figure 84C 

DIAGNOSIS Cephalon width equal to or greater than length; frontal margin 

narrowly rounded. Posterolateral angles of anterior pereonites weakly pro 

duced, rounded to subacute; of posterior pereonites more strongly produced, 

subacute to acute. Coxal plates 3—7, 4—7, or 5—7 with acute posterolateral 

angles; coxae rarely reaching beyond posterior margins of their respective 

pereonites. 

Nerocila acuminata Schioedte and Meinert, 1881, forma aster 

Figure 84D 

DIAGNOSIS Cephalon always wider than long; anterior margin broadly 

rounded. Posterolateral angles of all pereonites strongly produced, acute, all 

reaching well beyond posterior margins of their respective pereonites. Coxal 

plates 2—7 strongly produced with acute posterior angles.

Renocila Miers, 1880 

Renocila colini 191 

DIAGNOSIS Body rarely curved. Cephalon anteriorly weakly to distinctly 

truncate. Antennular bases well separated. Antennules and antennae some 

what flattened, antennules usually broader and longer than antennae. Per- 

eonites 5—7 with posterolateral corners more or less strongly produced. 

Pleonites not laterally incised. 

REMARKS Williams and Williams (1980) provide a key to nine species of 

Renocila. 

Key to species of Renocila 

1. Posteroventral angle of pereonite 7 reaching pleonite 1 colini 

Posteroventral angle of pereonite 7 reaching beyond pleonite 1 2 

2. Dorsal surface of body brown; posteroventral angle of pereonite 7 

reaching pleonite 2 waldneri 

Dorsal surface of body black; posteroventral angle of pereonite 7 

reaching pleonite 3 bowmani 

Renocila bowmani Williams and Williams, 1980 

Figure 85A 

DIAGNOSIS 9 18.0 mm, 6 11.5 mm. Posteroventral angles of pereonites 5— 

7 produced, that of pereonite 7 overlapping pleonites 1-3. Pereopods 1—3 

lacking swelling on dactylus. Pereopods 6-7 subequal in length. Uropodal 

exopod longer than endopod. Pleotelson length 3 A basal width. Color: dorsal 

surface of body and appendages uniform black. Attached to dorsum of body 

close to dorsal fin. 

RECORDS Harlequin bass Serranus tigrinus: Dominican Republic. 

Renocila colini Williams and Williams, 1980 

Figure 85B,C 

DIAGNOSIS Ovigerous 9 12.0-17.5 mm, 6 7.5-13.0 mm. Pereonites 5-7 

with posteroventral angle produced, that of pereonite 7 overlapping pleonite 

1 only. Pereopods 1—3 lacking swelling on dactyli; pereopods 6—7 subequal in 

length. Uropod reaching beyond pleotelson, endopod more than half length


c 

Figure 85. A, Renocila bowmani. Renocila colini: B, 9; C, d. D> Renocila waldneri. 

of exopod. Pleotelson l /i to 72 wider than long, with slight rounded apex 

Color: dorsal surface of body and appendages uniformly yellowish brown 

Attached to dorsum of body, close to dorsal fin.

FLABELLIFERA • L1MNOR11DAE 193 

RECORDS Flamefish Apogon maculatus: Puerto Rico. Belted cardinalfish Ap- 

ogon townsendi: Puerto Rico. 

Renocila waldneri Williams and Williams, 1980 

Figure 85D 

DIAGNOSIS Ovigerous 9 15.3-19.3 mm, 6 5.0-10.8 mm. Posteroventral 

angle of pereonite 5 moderately produced, of pereonites 6-7 more strongly 

produced, that of pereonite 7 overlapping pleonites 1 and 2. Pereopods 1-3 

without swelling on dactyli. Pereopods 6 and 7 subequal in length. Uropodal 

exopod slightly longer than endopod. Pleotelson basally wider than long; 

posterior margin broadly and evenly rounded. Color: dorsal surface of body 

uniform brown; appendages yellowish brown. Attached to dorsum of body 

close to dorsal fin. 

RECORDS Harlequin bass Serranus tigrinus: Dominican Republic. 

Family Limnoriidae Harger, 1879 

DIAGNOSIS Body ovate in cross section, often becoming more setose posteri 

orly. Cephalon subspherical, freely articulating with pereonite 1; eyes lateral. 

Antennules and antennae well separated at bases. Mandible with strong in 

cisor; lacking molar and well-defined lacinia mobilis, but with species- 

distinctive lacinioid bristle or seta; palp usually of three articles. Maxillipe- 

dal palp of five articles; endite well developed. Coxae present on pereonites 

2-7. Pleon consisting of five free pleonites plus pleotelson; latter subcircular, 

set obliquely to axis of body, usually with anterolateral crests. Uropod with 

strong protopod inserted ventrolaterally. 

Key to genera of Limnoriidae 

1. Uropodal rami very unequal 2 

Uropodal rami subequal Paralimnoria 

2. Mandibular incisors possessing rasp and file Limnoria 

Mandibular incisors lacking rasp and file Phycolimnoria

194 FLABELLIFERA • LIMNORIIDAE 

REMARKS This family includes a number of species that are of considerable 

economic importance. Given that species of Limnoria are wood borers, 

wooden structures such as wharf pilings that are immersed in sea water and 

even in water of reduced salinity are vulnerable to attack by these gribbles. 

Prolonged exposure can lead to weakening and eventual collapse of these 

structures (see Ray, 1959). Even creosote-treated wood is not fully protected; 

Limnoria tuberculata will bore into such wood to where the creosote has not 

penetrated. 

The isopods rasp at the wood fibres with the rasp and file structures of the 

mandibles, usually following the grain of the wood. With this boring activity, 

saprophytic fungi and bacteria invade the wood and assist in the breakdown 

process. Limnoria lack cellulase-secreting microflora in their gut, but proba 

bly secrete a cellulase themselves (Boyle and Mitchell, 1978). It is also prob 

able that the fungi and bacteria, the latter often densely aggregated on the 

setae of the isopod, form part of the animals' diet. In the natural environ 

ment, Limnoria perform an important role in the breakdown of dead wood, 

especially in mangrove areas. 

Sexual dimorphism of the pleotelson does occur in some species. This as 

pect of the morphology, however, has hardly been investigated. 

Limnoria Leach, 1814 

DIAGNOSIS Antennular flagellum of four articles. Antennal flagellum of 

three to five articles. Incisor of right mandible equipped with filelike struc 

ture on upper surface; incisor of left mandible with rasplike structure. Rami 

of pleopod 5 lacking marginal setae. Uropodal exopod much shorter than 

endopod, bearing terminal claw. Pleotelson smooth, or variously ornamented 

with tubercles and ridges. 

Limnoria indica Becker and Kampf, 1958 

Figure 86A,B 

DIAGNOSIS 6 3.0 mm, ovigerous 9 3.0 mm. Pleonite 5 with submedian pair 

of strong rounded ridges, converging slightly posteriorly. Pleotelson basally 

with two pairs of submedian tubercles and pair of lateral tubercles. 

RECORDS Cozumel, Mexico; Man o'War Cay, Belize. 

India; Hong Kong; Philippines; east coast of Australia.

Key to species of Limnoria 

Limnoria insulae 195 

1. Dorsal surface of pleotelson lacking prominent tubercles, ridges, or 

carinae (L. simulata may appear to lack ornamentation; in this 

species the tubercles are very small) 2 

Dorsal surface of pleotelson bearing tubercles, ridges, or carinae .... 3 

2. Pleotelson flat; pleonite 5 with broadly rounded middorsal ridge 

Pleotelson cup shaped; pleonite 5 with strong narrowly rounded 

platycauda 

middorsal ridge insulae 

3. Pleotelson with basal tubercles but lacking ridges 4 

Pleotelson with ridges but lacking freestanding tubercles 7 

4. 6 pleotelson with single strong middorsal tubercle unicornis 

Pleotelson with more than one basal tubercle 5 

5. Pleotelson with three basal tubercles tuberculata 

Pleotelson with more than three basal tubercles 6 

6. Pleotelson with four basal tubercles in line (difficult to detect) simulata 

6 pleotelson with six basal tubercles indica 

7. Pleotelson with single middorsal longitudinal ridge multipunctata 

Pleotelson with two rounded basal ridges 8 

8. Pleonite 5 with strong Y-shaped ridge Pfeffe 

Pleonite 5 with two posteriorly converging ridges saseboensis 

Limnoria insulae Menzies, 1957 

Figure 86C 

DIAGNOSIS 6 3.0 mm, ovigerous 9 3.4 mm. Pleonite 5 with strong middor 

sal ridge. Pleotelson cup shaped, lateral crests extended anteromesially, sep 

arated basally by distinct gap; posterior margin and lateral crests not 

tuberculate. 

RECORDS Twin Cays, Belize.


200HM 

Figure 86. Limnoria indica: A, pi 6;B, pi 9. Limnoria insulae: C 

pleotelson. Limnoria multipunctata: D, pleotelson in oblique-lateral view 

L i m n o r i a m u l t i p u n c t a t a M e n z i e s , 1 9 5 7 

F i g u r e s 8 6 D ; 8 7 A 

D I A G N O S I S 6 2 . 8 m m , o v i g e r o u s 9 3 . 0 m m . P l e o n i t e 5 d o r s a l l y s m o o t h . 

P l e o t e l s o n w i t h m i d d o r s a l l o n g i t u d i n a l r o u n d e d r i d g e b e a r i n g s e v e r a l

Limnoria multipunctata 197 

Figure 87. Limnoria multipunctata: A, pleotelson; Limnoria pfefferi: B} pleotelson 

Limnoria platycauda: C, pleotelson; Limnoria saseboensis: D> pleotelson; Limnoria 

simulata: E, pleotelson; Limnoria tuberculata: F, pleotelson. 

b u t t o n - s h a p e d t u b e r c l e s i n p o s t e r i o r h a l f ; p o s t e r i o r m a r g i n a n d l a t e r a l c r e s t s 

t u b e r c u l a t e . 

R E C O R D S P u e r t o R i c o ; J a m a i c a ; T w i n C a y s , B e l i z e 

J a p a n ; K a i I s l a n d s , S o u t h P a c i f i c .


Limnoria pfefferi Stebbing, 1904 

Figure 87B 

DIAGNOSIS 8 3.8 mm, ovigerous 9 4.0 mm. Pleonite 5 with conspicuous 

middorsal Y-shaped carina. Pleotelson basally with pair of submedian 

rounded ridges; lateral crests lacking tubercles. 

RECORDS Florida Keys; Bahamas; Puerto Rico; U.S. Virgin Islands; Twin 

Cays and Man o'War Cay, Belize; Yucatan Peninsula, Mexico. 

Minikoi Atoll and Aldabra Atoll, Indian Ocean; Philippines; New Guinea; 

Panama. 

Limnoria platycauda Menzies, 1957 

Figure 87C 

DIAGNOSIS 6 2.5 mm, ovigerous 9 2.6 mm. Pleonite 5 with broad middor 

sal longitudinal rounded ridge. Pleotelson lacking dorsal ornamentation; 

posterior margin and lateral crests bearing tubercles. 

RECORDS Cuba; Puerto Rico to Curasao; Cozumel, Mexico; Twin Cays 

and Man o'War Cay, Belize. 

Aldabra Atoll, Indian Ocean. 

Limnoria saseboensis Menzies, 1957 

Figure 87D 

DIAGNOSIS 6 3.5 mm. Pleonite 5 with submedian pair of ridges, converging 

slightly posteriorly. Pleotelson basally with submedian pair of anteriorly tu- 

berculate ridges; posterior margin and lateral crests tuberculate. 

RECORDS Miami, Florida. 

Japan; Fiji. 

Limnoria simulata Menzies, 1957 

Figure 87E 

DIAGNOSIS 6 3.8 mm, ovigerous 9 4.0 mm. Pleonite 5 with obscure me 

dian longitudinal groove. Pleotelson basally with submedian pair of tubercles 

and small lateral tubercles, latter often difficult to detect; lateral crests 

tuberculate. 

RECORDS Florida Keys; U.S. Virgin Islands; Gulf of Mexico.

Limnoria tuberculata Sowinsky, 1884 

Figure 87F 

Paralimnoria andrewsi 199 

DIAGNOSIS 6 2.8 mm, ovigerous 9 3.0 mm. Pleonite 5 with two anterior 

tubercles, one middorsal posterior tubercle, area between tubercles de 

pressed. Pleotelson basally with middorsal tubercle, followed by pair of sub- 

median tubercles, all three tubercles having short obscure carina; posterior 

margin and lateral crests tuberculate. 

RECORDS Rhode Island to Venezuela; Cuba; Man o'War Cay, Belize; Gulf 

of Mexico. 

Uruguay; West Africa; Mediterranean; Black Sea; India; Hong Kong; 

Hawaii; Australia; California. 

REMARKS This species has frequently been recorded under the name Lim 

noria tripunctata Menzies, 1951a. 

Limnoria unicornis Menzies, 1957 

Figure 88A,B 

DIAGNOSIS 6 2.6 mm, ovigerous 9 2.6 mm. Mandibular palp of one arti 

cle. Pleonite 5 with somewhat obscure Y-shaped ridge middorsally. 

Pleotelson in 6 with strong basal slightly curved middorsal tubercle; lateral 

crests lacking tubercles. 

RECORDS Bahamas; Man o'War Cay and Twin Cays, Belize. 

Caroline Islands; Palau; Society Islands. 

Paralimnoria Menzies, 1957 

DIAGNOSIS Antennular flagellum of five articles. Antennal flagellum of five 

or six articles. Mandibular incisor with rasp and file. Pleopod 5, rami bearing 

marginal setae. Uropodal rami subequal in length, each with clawlike apex. 

Paralimnoria andrewsi (Caiman, 1910) 

Figure 88C,D 

DIAGNOSIS 6 2.6 mm, 9 2.6 mm. Pleonite 5 with or without triangular 

middorsal depressed area. Pleotelson with basal submedian pair of tubercles 

either obscurely or strongly carinate; lateral crest tubercles of variable 

strength.


Figure 88. Limnoria unicornis: A, pleotelson, 8; B, pleon, 6*, in lateral view. 

Paralimnoria andrewsi: C, p l e o n i t e 5 a n d p l e o t e l s o n ; D , u r o p o d . Phycolimnoria clarkae: 

E , p l e o n i t e 5 a n d p l e o t e l s o n ; F , u r o p o d a n d p l e o t e l s o n i n l a t e r a l v i e w . 

R E C O R D S F l o r i d a K e y s ; P u e r t o R i c o ; T w i n C a y s , B e l i z e ; C u r a g a o 

C h r i s t m a s I s l a n d s , I n d i a n O c e a n ; S a m o a ; H a w a i i ; J a p a n . 

R E M A R K S M e n z i e s ( 1 9 5 7 ) d i s c u s s e s t h r e e f o r m s o f t h i s s p e c i e s : F o r m a typ- 

i c a , w h i c h l a c k s a c e n t r a l d e p r e s s e d a r e a d o r s a l l y o n p l e o n i t e 5 a n d h a s a p a i r 

o f s u b m e d i a n o b s c u r e l y c a r i n a t e t u b e r c l e s o n t h e p l e o t e l s o n ; F o r m a A , w h i c h 

h a s a t r i a n g u l a r d e p r e s s e d a r e a d o r s a l l y o n p l e o n i t e 5 a n d a p a i r o f s u b m e -

FLABELLIFERA • SEROLIDAE 201 

dian tubercles supported by strong carinae on the pleotelson; Forma B5 hav 

ing a triangular depressed area dorsally on pleonite 5 and an obscurely cari- 

nate pair of tubercles on the pleotelson. Given that at least two of these forms 

have been recorded occurring together, it would seem that this is merely a 

highly variable species. 

Phycolimnoria Menzies, 1957 

DIAGNOSIS Mandibular incisor lacking rasp and file. Uropodal rami une 

qual, exopod longer than endopod, latter usually with clawlike apex. 

REMARKS Most species of Phycolimnoria are algal borers, frequently encoun 

tered in the holdfasts of brown algae such as Macrocystis, Laminaria, and 

Sargassum, The one species recorded from the Caribbean, P. clarkae, however, 

has only been taken from decaying wood. 

Phycolimnoria clarkae Kensley and Schotte, 1987 

Figure 88E,F 

DIAGNOSIS 6 4.3 mm, ovigerous 9 3.3-4.4 mm. Uropodal exopod less 

than half length of endopod, straight, tipped with short squat claw. Pleonite 

5 with broad raised middorsal region having irregular bumps. Pleotelson 

wider than long, with two rounded submedian ridges basally, becoming ob 

solete posteriorly. 

RECORDS Bahamas; Twin Cays, Belize. 

Aldabra Atoll, Indian Ocean. 

Family Serolidae Dana, 1852 

DIAGNOSIS Body dorsoventrally depressed. Eyes present or absent. 

Cephalon fused with pereonite 1 dorsally. Mandible bearing palp. Maxillipe- 

dal palp of one to four articles. Pereonites 2—4 with coxae demarked; per- 

eonites 5 and 6 with coxae not demarked; pereonite 7 narrow, lacking free 

lateral margins. Pereopod 1 in 6 and 9 subchelate, pereopod 2 subchelate or 

ambulatory in 6, ambulatory in 9. Pleonites 1 and 2 free, articulated, re 

mainder of pleonites fused with telson. Pleopods 1—3 small, natatory; 

pleopods 4 and 5 large, operculate. Uropods lateral, biramous. 

REMARKS The serolids reach their greatest diversity (and their greatest size 

of up to 80 mm in length) in the southern oceans, with few species extending

202 FLABELLIFERA • SPHAEROMATIDAE 

into the subtropics and tropics. The deep- and abyssal-dwelling species usu 

ally lack eyes. The animals are epibenthic, living in the upper few centime 

ters of the bottom sediment, where they are scavengers and carnivores. 

Serolis Leach, 1818 

DIAGNOSIS Body markedly dorsoventrally flattened. Coxal plates produced 

laterally. Mandible having lacinia mobilis and single spine. Maxillipedal 

palp of three articles (rarely two to four). Pereopod 2 exhibiting sexual di 

morphism, subchelate in cJ, ambulatory in 9. Pleopods 1-3, peduncles elon 

gate, rami subelliptical. Pleopod 3, exopod uniarticulate. 

Serolis mgrayi Menzies and Frankenberg, 1966 

Figure 89 

DIAGNOSIS S 4.5 mm, ovigerous 9 4.7 mm. Eyes present. Cephalon with 

two middorsal tubercles. Pereonites 2—4 each with faint rounded tubercle 

just mesial to coxal suture. Pereon and pleon with faint middorsal longitudi 

nal carina bearing small blunt tubercle on posterior margin of each segment. 

Pleonites 1 and 2 with lateral margins not contributing to body outline, over 

lapped by pereonite 6. Pleotelson broadly triangular, with lateral carina in 

anterior half; apex truncate. Uropodal rami reaching to or slightly beyond 

pleotelsonic apex. 

RECORDS Off North Carolina, 18-34 m; off South Carolina, 22 m; off 

Georgia, 18-47 m; Florida Keys, 18-88 m; Trinidad; Venezuela, 95 m; Flor 

ida, Gulf of Mexico, 11-88 m. 

Family Sphaeromatidae H. Milne Edwards, 1840 

DIAGNOSIS Antennular peduncle of three articles, antennal peduncle of five 

articles. Mandible stout, lacinia mobilis and molar usually well developed, 

palp of three articles. Maxillipedal palp of five articles. Mouthparts in some 

genera metamorphosed and somewhat reduced in ovigerous 9 . Pleon of five 

partially or completely fused pleonites, often indicated by lateral sutures, 

plus dorsally convex and sometimes inflated pleotelson. Uropods lateral, ex 

opod free if present, endopod fused with sympod. Sexual dimorphism often 

marked, especially in pleotelsonal structure. Animal often capable of con 

globating or folding over. Young brooded in internal pouches or anterior or 

posterior pockets; oostegites variable in number, if present.

FLABELLIFERA • SPHAEROMATIDAE 203 

Figure 89. Serolis mgrayi: A, 6; B, pereopod 1; C, pereopod 2, 8. 

REMARKS Right into the 1980s this family was routinely divided into three 

groups, based on the structure of the two posterior pairs of pleopods: 

Platybranchiatae—pleopods 4 and 5 with both rami membranous and lack 

ing branchial pleats; Hemibranchiatae—pleopods 4 and 5 with branchial 

pleats on endopods only; Eubranchiatae—pleopods 4 and 5 with branchial 

pleats on both rami. These three "groups" were recognized formally as sub 

families by Hurley and Jansen (1977) but the names were not based on con-

204 FLABELL1FERA • SPHAEROMATIDAE 

tained genera and were replaced with current subfamily names by Bowman 

(1981) and Iverson (1982), the latter providing diagnoses for all five sub 

families. Four of these are represented in the Caribbean area; the fifth, the 

Tecticipitinac, contains only the single primarily Pacific genus Tecticeps. 

While the subfamilial status now appears to be resolved, many of the gen 

era still require unambiguous diagnoses. The work of Harrison (1984) on the 

structure of the female broodpouch, with its various components of 

oostegites, internal pouches, and anterior and posterior pockets (Figure 90), 

along with the metamorphosis of the female mouthparts (see Figure 96) has 

helped enormously to standardize the genera. Nevertheless, these features of 

the female remain unknown in several genera. Further, with this stabilization 

based on females, many problems of incorrect generic designation have been 

uncovered. In this work, Harrison's generic diagnoses are followed as far as 

possible. Where uncertainty exists, this is indicated. In some cases, we may 

still be unaware of existing problems: future work will without doubt result in 

the shifting of species to different genera, as well as in the creation of new 

genera. 

Key to subfamilies of Sphaeromatidae 

1. Pereopod 1 prehensile in both sexes; pereopod 2 prehensile only in 6 

Ancininae 

Pereopods 1 and 2 ambulatory 2 

2. Pleopods 4 and 5 lacking branchial pleats Cassidininae 

Pleopods 4 and 5 with branchial pleats on endopods 3 

3. Pleopods 4 and 5 with branchial pleats on both rami . . . Dynameninae 

Pleopods 4 and 5 with branchial pleats on endopods only 

Subfamily Ancininae Tattersall, 1905 

Sphaeromatinae 

DIAGNOSIS Body markedly dorsoventrally depressed. Cephalon fused me 

dially with pereonite 1. Pereopod 1 prehensile in 8 and 9. Pereopod 2 pre 

hensile in 6 only. Pleopods 4 and 5 similar, lacking branchial pleats. 

Uropods uniramous.

internal pouch 

Ancinus belizensis 205 

Figure 90. Diagrammatic representation of 9 sphaeromatid, showing marsupial 

structures (adapted from Harrison, 1984). 

Ancinus H. Milne Edwards, 1840 

DIAGNOSIS Eyes dorsal. 9 mouthparts not metamorphosed. Mandibular 

molar absent; palp of three articles. Maxilla 1 of single ramus, endite rudi 

mentary. Maxilla 2 of two rami. 9 with oostegites absent; brood held in two 

opposing pockets, opening as narrow ventral slit between pereopods 4. Pleon 

consisting of short anterior pleonite with free lateral margin, plus broadly 

triangular pleotelson. Pleopod 1 uniramous, endopod absent. Pleopod 2 op- 

erculiform. Pleopod 3, exopod of single article. Uropod lacking exopod, sym- 

pod not laterally expanded. 

Key to species of Ancinus 

1. Pleotelson as long as basal width, apex narrowly rounded . . . brasiliensis 

Pleotelson with basal width greater than length, apex subtruncate 

Ancinus belizensis Kensley and Schotte, 1987 

Figure 91A-C 

belizensis 

DIAGNOSIS 8 4.1 mm, 9 2.8 mm. Body oval, about twice longer than wide. 

Dorsal integument strongly pitted. Antennular flagellum of 12 articles; an-


Figure 91. Ancinus belizensis: A, 9; B, pereopod 1 cJ; C, pereopod 2 8. Ancinus 

braziliensis: D, adult (from Glynn and Glynn, 1974). 

tennal flagellum of 10 articles. 8 pereopod 2, dactylus strongly curved, 

reaching to proximal lobe of propodus. Pleopod 2 about 2.5 times longer than 

basal width. 

RECORDS Carlson Point, Belize, in seagrass flats, 0.5 m. 

Ancinus brasiliensis Lemos de Castro, 1959 

Figure 91D 

DIAGNOSIS 8 7.0 mm, 9 6.0 mm. Body about twice longer than wide. Dor 

sal integument smooth. Antennular flagellum of 17 articles; antennal 

flagellum of 10 articles. 8 pereopod 2, dactylus strongly curved, reaching to 

midlength of posterior margin of carpus. Pleopod 2 almost three times longer 

than basal width.

Cass idin idea 207 

RECORDS Brazilian coast from Rio de Janeiro northward, 1.5 m; Costa 

Rica, Panama; shallow infratidal below sandy beaches. 

REMARKS Glynn and Glynn (1974) discussed color polymorphism in this 

species. 

Subfamily Cassidininae Iverson, 1982 

DIAGNOSIS Cephalon not medially fused with pereonite 1. Pereopod 1 am 

bulatory. Pleopods 4 and 5, both rami lacking transverse pleats, outer rami 

unsegmented. Pleopod 5, outer ramus with low subapical squamiferous pro 

tuberances. Pleotelsonic apex entire. Uropods with exopods reduced. 

REMARKS The genus Dies has twice been recorded from the Caribbean: D. 

arndti Ortiz and Lalana, 1980, from Cuba, and D. barnardi Carvacho, 1977, 

from Guadeloupe. This genus is distinguished from Cassidinidea solely on the 

basis of the penial structure: biramous in Cassidinidea, uniramous in Dies. 

Harrison (1984) has pointed out that the separation of these two genera has 

not been satisfactorily resolved. The penis of neither the Cuban nor the 

Guadeloupan species has been illustrated, but the whole-animal illustrations 

of both look suspiciously like Cassidinidea ovalis. Examination of material of D. 

barnardi from the Paris Museum supports the view that this species was based 

on immature material of C. ovalis. Neither of the so-called species oi Dies are 

dealt with in this work, both being regarded as junior synonyms of C. ovalis. 

Key to genera of Cassidininae 

1. Frontal lamina visible dorsally between antennular bases; two basal 

articles of antennular peduncle not expanded Cassinidinea 

Frontal lamina not visible between antennular bases; two basal articles 

of antennular peduncle broadly expanded Paraleptosphaeroma 

Cassidinidea Hansen, 1905b 

DIAGNOSIS Body strongly dorsoventrally depressed. Eyes dorsal, situated 

at posterolateral corners of cephalon. Latter somewhat sunken into pereonite 

1. Frontal lamina expanded, visible dorsally between antennular bases. An 

tenna directed laterally. Pleon consisting of one free pleonite having short 

free lateral margin, plus broadly triangular pleotelson. Uropodal endopod


well developed, fused with sympod; exopod markedly reduced. Penial rami 

elongate, separate. 9 mouthparts not metamorphosed. Oostegites absent. 

Brood housed in pouch formed by opposing pockets overhanging ventrum, 

opening by slit between fourth pereopods. 

Key to species of Cassidinidea 

1. Posterior margin of pleotelson truncate ovalis 

Posterior margin of pleotelson rounded mosaica 

Cassidinidea mosaica Kensley and Schotte, 1987 

Figure 92A 

DIAGNOSIS 6 1.8 mm, ovigerous 9 1.6 mm. Body twice longer than wide. 

Dorsal integument bearing close-packed flattened tubercles. Pleotelson tri 

angular, with posterior margin narrowly rounded, dorsally convex, basally 

inflated. 

RECORDS Carrie Bow Cay, Belize, 1.5-10 m; in silty sand and rubble be 

tween patch reefs and coral buttresses. 

Cassidinidea ovalis (Say, 1818) 

Figure 92B-E 

DIAGNOSIS 6 and 9 3.6 mm. Body width slightly less than half length. 

Dorsal integument smooth. Pleotelson with raised anteromesial area, but 

lacking sculpture; posterior margin truncate. 

RECORDS New Jersey to Florida, in marsh mud and among dead leaves, 0— 

1 m; Trinidad; Belize; Panama; Dominica; Louisiana and Vera Cruz, Gulf of 

Mexico. Known from waters of less than l%o to 35%o. 

Paraleptosphaeroma Buss and Iverson, 1981 

DIAGNOSIS Body oval in outline, entire circumference with transparent 

flange of fused setae on two expanded basal articles of antennule, on per- 

conites, pleonite 1, and uropods. Expanded basal articles of antennules con-

Paraleptosphaeroma 209 

Figure 92. Cassidinidea mosaica: A, 6. Cassidinidea ovalis: B, 6] C, pereopod 1; Dy 

pleopod 4; E, pleopod 5. Paraleptosphaeroma glynni: Fy 6. 

tiguous in midline. Single articulated pleonite with short free lateral margin. 

Uropodal sympod and endopod fused; exopod articulated, much shorter 

than fused endopod.


Paraleptosphaeroma glynni Buss and Iverson, 1981 

Figure 92F 

DIAGNOSIS 6 2.58 mm, ovigerous 9 2.38 mm. Pleotelson basally broad, 

tapering to notched posterior margin. Fused uropodal endopod and sympod 

of each side almost touching posterior to pleotelsonic apex. 

RECORDS Portsmouth, Dominica, intertidal rock pools. 

Punta Paitilla, Pacific Panama. 

REMARKS Buss and Iverson (1981) demonstrated that this species displays 

sequential protogynous hermaphroditism, and that the change from female 

to male seems to be mediated by social conditions, especially the proportion 

of males to females. The principal food source for this species was shown to 

be abascan bryozoans. 

Key to genera of Dynameninae 

1. Pleotelson very similar in both sexes 2 

Pleotelson showing marked sexual dimorphism 3 

2. Cephalon and pleotelson smooth, lacking ridges Ischyromene 

Pleotelson and cephalon with ridges Cerceis 

3. Uropods lamellar in both sexes 4 

Uropods lamellar in 9, endopod reduced, exopod elongate-cylindrical 

in 6 5 

4. Ovigerous 9 lacking oostegites;

Subfamily Dynameninae Bowman, 1981 

Discerceis 211 

DIAGNOSIS Cephalon not fused with pereonite 1. Pereopods 1 and 2 am 

bulatory. Pleopods 4 and 5, both rami having branchial pleats. Pleopod 4, 

exopod unjointed, usually lacking setae, endopod with few setae at most. 

Pleotelsonic apex often with terminal notch or foramen, especially in


Figure 93. "Cerceis" carinata: A, 6. Discerceis linguicauda: B, d. Dynamenella 

acutitelson: C, 9; D, pleon 8. Dynamenella angulata: E, 9. 

internal pouches (number unknown). Pockets absent. 6 with uropodal endo- 

pod and sympod fused, very short; exopod elongate, cylindrical.

Discerceis linguicauda (Richardson, 1901) 

Figure 93B 

Dynamenella 213 

DIAGNOSIS 6 7.2 mm. Dorsal integument, especially of posterior half, with 

numerous scattered granular tubercles. Uropodal endopod and sympod 

fused, very short, exopod elongate, subcylindrical and slightly bowed. Ante 

rior half of pleotelson inflated, with three elongate rounded ridges (each com 

posed of two contiguous tubercles) ending posteriorly in subacute tubercle; 

posterior margin trilobed, median lobe broadly rounded, with subacute tu 

bercle at base, lateral lobe truncate, well separated from median lobe. Head 

and pereonite 1 not fused. Frontal lamina visible dorsally between antennal 

bases. Penes short, separate. Copulatory stylet basally relatively broad, dis- 

tally broadly rounded. 

RECORDS Cape Catoche, Yucatan, Mexico, 48-50 m. 

REMARKS This species is known only from the four male syntypes. 

Dynamenella Hansen, 1905b 

DIAGNOSIS Species exhibiting obvious sexual dimorphism. Both sexes lack 

ing processes on pereon and pleon, Uropodal rami lamellar. Exopod of 

pleopod 3 with or without articulation. 9: Mouthparts not metamorphosed. 

Broodpouch lacking oostegites, but formed by two opposing ventral pockets 

opening in midline between fourth pereopods. Apex of pleotelson with notch, 

Key to species of Dynamenella 

1. 6 with pleotelsonic foramen 2 

6 lacking foramen but with notch, or appearing entire; 9 pleotelson 

with faint notch visible acutitelson 

2. 6 with four strong pleotelsonic ridges; 9 with subcircular pleotelsonic 

foramen quadrilirata 

6 lacking pleotelsonic ridges; 9 with posterior margin of pleotelson 

entire perforata


groove, or foramen. 6: Penes basally fused, rami long, tapering. Copulatory 

stylet proximally broad, tapering to acute tip, reaching to or just beyond 

apex of endopod. Uropods broader than in $. Posterior pleotelson with dor- 

sally directed foramen connected to apex by narrow slit. 

REMARKS The species described by Richardson (1901) as Dynamene angulata 

from No Name Key, Florida, and referred to by some authors as a Dyna- 

menella, while figured here (Figure 93E), is not included in the present key. 

The species is known only from immature females; correct generic placement 

is thus not possible. 

Dynamenella acutitelson Menzies and Glynn, 1968 

Figure 93C,D 

DIAGNOSIS 6 3.5 mm, 9 2.3 mm. 6: Pereonites 4-6 with transverse ridge 

over dorsum, ridge interrupted to form short median section. Pleotelson with 

two submedian and two lateral rounded tubercles basally, two submedian, 

poorly defined ridges in central area; posterior margin tapering in dorsal 

view, with slit eitherjust visible or appearing entire. In lateral view, posterior 

pleotelson seen to be laterally compressed, forming narrow groove. 

RECORDS Puerto Rico, intertidal rocks and algae. 

REMARKS Menzies and Glynn (1968) described this species with two vari 

eties, the holotype as D. acutitelson var. typica, and 11 paratypes as D. acu 

titelson var. glabrothorax. The major difference between these varieties lay in 

the presence of transverse ridges on pereonites 4—6 in typica and their absence 

xnglabrothorax. The holotype, however, at 3.5 mm, would seem to be a mature 

male, while all the paratypes are smaller. The differences described by Men 

zies and Glynn (1968) may thus be due to immaturity. As further compara 

tive material is lacking, these varieties (or whatever their true status) are not 

recognized here. 

Menzies and Glynn (1968, fig. 30a) illustrate D. acutitelson var. glabrothorax 

as having scattered tiny granules over the dorsal integument. These were not 

seen when the type material was reexamined. 

Harrison and Holdich (1982) placed this species in Paradella, based on the 

literature. However, the penes for both varieties are shown as short and sepa 

rate, as in Ischyromene. Again, until further mature males and ovigerous 

females are seen, the generic placement of this species must remain in doubt.

Dynamenella perforata (Moore, 1901) 

Figure 94A,B 

Geocerceis barbarae 215 

DIAGNOSIS 6 3.2 mm, 9 2.6 mm. 6: Pleon bearing two low rounded sub- 

median "mounds." Pleotelson with strongly convex anterior two-thirds, with 

T-shaped foramen. Pleon and pleotelson with numerous scattered small tu 

bercles. Uropodal rami broadly ovate, outer margins crenulate. 9: 

Pleotelson broadly rounded in dorsal view, posterior margin entire. Inner 

uropodal ramus distally subacute. 

RECORDS Bermuda to Puerto Rico, intertidal coral rubble and algae, and 

under chiton Acanthopleura granulate; Dominican Republic; Cuba. 

Dynamenella quadrilirata Kensley, 1984 

Figure 94C-H 

DIAGNOSIS 6 2.6 mm, 9 2.5 mm. 6: Two low rounded submedian tuber 

cles on last pleonite. Anterior half of pleotelson inflated, with four rounded 

longitudinal ridges; posterior half tapered, somewhat dorsally flexed, with 

cordate foramen. Uropodal rami distally rounded, outer margins crenulate 

to dentate. 9: Lacking pleonal tubercles. Pleotelson inflated, unornamented, 

posterior margin forming subcircular foramen. 

RECORDS Carrie Bow Cay, and Twin Cays, Belize; intertidal to 3 m. 

Geocerceis Menzies and Glynn, 1968 

DIAGNOSIS Ovigerous 9 with mouthparts metamorphosed. Broodpouch 

with three pairs of oostegites, on pereonites 2—4, just overlapping in midline. 

Brood held in internal pouches (number unknown). Pockets absent. Uropo 

dal rami lamellar, shorter than pleotelson. 6 uropodal endopod fused with 

sympod, very short; exopod elongate, club shaped. Pleopod 2 with copula- 

tory stylet articulating distally on endopod. 

Geocerceis barbarae Menzies and Glynn, 1968 

Figure 95A-C 

DIAGNOSIS 8 3.3 mm, 9 2.5 mm. Pleopod 3 exopod of single article. Pleon 

with two elongate sutures reaching lateral pleon margin. 6: Frontal lamina 

expanded into ventrally directed beaklike process. Penes separate, relatively


Figure 94. Dynamenella perforata: A, 6; B> pleon 9. Dynamenella quadrilirata: C, 6; 

D, pleon 9; E, pleon

Ischyromene 217 

Figure 95. Geocerceis barbarae: A, 8; B, pleon 9; C,


nounced. Uropodal rami lamellar. 8 pleopod 2 with copulatory stylet 

basally narrow, reaching to or just beyond distal margin of endopod. 

Ischyromene barnardi (Menzies and Glynn, 1968) 

Figure 95D 

DIAGNOSIS 6 4.5 mm, 9 3.7 mm. Both sexes lacking processes on pereon 

and pleon. Accessory unguis of pereopods often bifid. Pleopod 3, exopod of 

single article. Uropodal rami lamellar. 6: Pereonite 7, posterior margin 

bilobed. Penes short, separate to base. 

RECORDS Puerto Rico, intertidal. 

Paracerceis Hansen, 1905b 

DIAGNOSIS Pleopod 3 exopod with transverse suture in distal half. Pleon 

with two long sutures reaching to posterolateral margin. 6: Penial rami 

short, separate. Pleotelson with basal area strongly vaulted; deep posterior 

notch sometimes having denticles on inner margins, and/or median tooth at 

base of notch. Uropodal endopod short, fused with sympod; exopod elongate, 

club shaped. 9: Mouthparts metamorphosed. Mandible fused with 

cephalon. Broodpouch of four pairs of oostegites, three posterior pairs over 

lapping. Brood retained in internal pouches. Uropodal rami subequal, lamel 

lar. Pleon usually less ornamented than in

Paracerceis caudata 219 

3. 6, pleotelsonic notch deep, margins usually with two teeth on each 

side; strong median tubercle on anterior pleotelson bluntly bifid; 9, 

pleotelson with one or two rounded median tubercles and 2 smaller 

tubercles on each side caudata 

6, pleotelsonic notch shallow, with tiny lateral denticles; median 

tubercle of pleotelson conical, acute; 9, pleotelson with three large 

conical acute tubercles and several smaller scattered tubercles in 

anterior half cohenae 

Paracerceis caudata (Say, 1818) 

Figure 96 

DIAGNOSIS 6 8.1 mm, 9 6.4 mm. 8: Pleotelson with blunt median bifid 

tubercle, with two smaller tubercles on each side. Pleotelsonic notch usually 

with two strong denticles on each margin, basal median tooth lacking. 

Uropodal exopod reaching well beyond pleotelson, slightly bowed, with 2—4 

setose bumps on outer margin. 9: Pleonite 5 with three low tubercles. 

Pleotelsonic apex broadly rounded in dorsal view, with two rounded median 

tubercles and two smaller tubercles on each side. Uropodal rami subequal, 

lamellar, outer distal angle of each acute. 

RECORDS Bermuda; New Jersey to Florida Keys; Yucatan to Venezuela; 

Turks and Caicos Islands; Cuba; Puerto Rico; Bahamas; Jamaica; Haiti; St. 

Maartens, 0.2-127 m; St. Lucia; Gulf of Mexico. Found in the following 

algae: Caulerpa, Halimeda, Turbinaria> Amphiroa, Laurencia, Dictyota; between 

sponges and tunicates on red mangrove roots; in coral rubble; in spur and 

groove zone of reefs, lagoon, back reef, seagrass flats, and fringing 

mangroves. 

REMARKS Menzies and Glynn (1968:55, fig. 22f) named and figured P. 

caudata var. brevipes from Puerto Rico. This variant was characterized as hav 

ing the margins of the pleotelsonic notch lacking denticles. Given the con 

siderable variation in ornamentation in this species, we feel that no validity 

can be given to the name "brevipes." 

This is the commonest sphaeromatid in the Caribbean, and it has very 

broad ecological requirements, being found in a wide range of habitats and 

depths.


Figure 96. Paracerceis caudata: A, 8,B, pleon 9; C, mandible 6] D, maxilla 1 6; E, 

maxilla 2 6; F, maxilliped 6; G> mandible 9; Hy maxilla 19;/, maxilla 1 9; J, 

maxilliped 9. 

Paracerceis cohenae Kensley, 1984 

Figure 97A,B 

DIAGNOSIS 6 10.0 mm, 9 7.9 mm. 6: Pereonites each with median tuber 

cle and several smaller lateral tubercles near posterior margin of somite. 

Pleonite 5 with large median conical tubercle. Anterior two-thirds of

Paracerceis glynni 221 

pleotelson inflated, faintly tripartite, with strong median conical tubercle; 

notch in posterior margin shallow, with low median tooth and tiny lateral 

denticles; posterolateral margins finely dentate. Uropodal exopod cylindri 

cal, distally denticulate, six to seven times longer than basal width. 9: Per- 

eon and pleon much as in 6 y but pleotelsonic notch shallower and 

posterolateral margins not denticulate. Uropodal rami subequal, lamellar, 

exopod with distolateral angle acute. 

RECORDS Carrie Bow Cay, Belize, 15-16 m. Only known from sponge 

Callispongia plicifera growing on outer reef slope. 

Paracerceis edithae Boone, 1930 

Figure 97C-E 

DIAGNOSIS 6 4.0 mm, 9 3.1 mm. 6: Posterior three pereonites and 

pleonites each with irregular row of small tubercles near posterior margin, 

densely setulose tubercles becoming spinose more posteriorly. Pleotelson 

with strong median conical tooth in anterior half, flanked by convex spinose 

mound. Pleotelsonic notch deep, with elongate median basal tooth bearing 

strong acute tooth at its base. Lobes of posterior pleotelsonic margin broad, 

flattened, margins denticulate. Uropodal exopod tuberculate, tapering, api- 

cally acute. 9: Integument much less tuberculate-spinose than in 6. Imma 

ture 9, posterior margin of pleotelson with faintly rounded median lobe. In 

mature 9, posterior margin distinctly trilobed. Uropodal rami subequal, 

lamellar, distally rounded, with tiny distolateral spine on exopod. 

RECORDS Bahamas, 60-66 m, in vase sponge; Haiti; Puerto Rico, 20—25 m. 

Paracerceis glynni Kensley, 1984 

Figure 97F,G 

DIAGNOSIS 6 6.4 mm, 9 5.2 mm. 6: Integument becoming strongly setose 

and tuberculate posteriorly from about pereonite 5. Posterior margin of inf 

lated anterior area of pleotelson bearing strong median conical tubercle and 

smaller acute lateral tubercle, with low swelling beneath each lateral tuber 

cle. Posterior notch deep, narrow, with small basal median tooth, lobes form 

ing notch tricuspid, outer cusps recurved dorsally. Uropodal exopod fairly 

straight, cylindrical, apically acute. 9: Body far less setose and tuberculate 

than 6. Pleotelson with strongly inflated anterior area having very faint mid- 

dorsal tubercle; notch well marked, formed by triangular lobes of

222 

FLABELLIFERA • SPHAEROMATIDAE 

E 

Figure 97. Paracerceis cohenae: A, 6\ B, pleotelson, 9. Paracerceis edithae: C, 

pleotelson, 9; D, mature 6] E, immature S. Paracerceis glynni: F, 8\ G} pleotelson, 

9. Paracerceis nuttingi: H, 9. 

pleotelsonic margin. Uropodal rami subequal, flattened, endopod with distal 

margin faintly trituberculate; exopod with few distal tubercles. 

RECORDS Alligator Light, Florida, 11 m; Carrie Bow Cay, Belize, 11-15.2 

B 

G

Paradella 223 

m, from green alga Halimeda sp. on forereef, and from sponge Aphysina 

jistularis. 

Paracerceis nuttingi (Boone, 1921) 

Figure 97H 

RECORDS Barbados; Puerto Rico, 1.5 m, from Cymodocea seagrass, and coral 

rubble and sponges. 

REMARKS The types of this species from Barbados consist only of females 

(total length 4.1 mm). Menzies and Glynn (1968) record an immature male 

i 

from Puerto Rico with an incipient pleotelsonic notch. This specimen, how 

ever, still has the subequal lamellar uropodal rami. The mature male, with 

the characteristically reduced uropodal endopod and cylindrical exopod, is 

unknown. The possibility exists that this is not a true Paracerceis. 

Paradella Harrison and Holdich, 1982 

DIAGNOSIS Marked 

ual dimorphism. Accessory unguis of pereopods simple, not bifid. Pleopod 3 

Key to species of Paradella 

1. Pereonite 7 with projecting bilobed flange; pleon and pleotelson finely 

but distinctly granulate 2 

Pleon and pleotelson smooth 3 

2. 8 with pleotelsonic foramen distinctly heart shaped, with median 

point; four submedian tubercles of pleotelson in 8 and 9 somewhat 

elongate; 9 pleotelson posteriorly narrowed, slit visible dorsally 

8 with pleotelsonic foramen wider than long, but lacking median 

point; four submedian tubercles of pleotelson in 8 small, rounded, 

obscure in 9; 9 pleotelson posteriorly truncate, slit not visible 

dianae 

dorsally plicatura 

3. Tubercles on pleotelson in 8 and 9 small to obscure; 8 with 

pleotelsonic foramen subcircular quadripunctata 

Tubercles on pleotelson broadly rounded mounds in 8 and 9; 8 with 

pleotelsonic foramen wider than long tumidicauda

224 FLABELL1FERA • SPHAEROMAT1DAE 

exopod with articulation. Uropodal rami lamellar. 9: Mouthparts not meta 

morphosed. One pair of oostegites arising from pereonite 4, short, not reach 

ing midline. Brood held in pouch formed by two opposing pockets covering 

entire ventrum, opening by transverse slit between 4th pereopods.

Paradella quadripunctata 225 

Figure 98. Paradella dianae: A, 8; B, pleopod 2 8; C, pleon 9. Paradella plicatura: 

Dy 8; E, pleon 9. Paradella quadripunctata: F, 8; G, pleon 9. Paradella tumidicauda; 

H, pleon 9 (from Glynn, 1970); /,


Paradella tumidicauda (Glynn, 1970) 

Figure 98H,I 

DIAGNOSIS 8 6.7 mm, 9 6.5 mm. 6: Last pleonite with two submedian 

swellings. Pleotelson with four submedian broadly rounded swellinglike tu 

bercles and two pairs of lateral tubercles. Foramen wider than long, posterior 

contiguous borders of foramen each bearing rounded swelling. 9: Pleotelson 

with four submedian swellinglike tubercles, sometimes with two obscure lat 

eral tubercles; posterior slit not visible dorsally, area surrounding slit 

swollen, horseshoe shaped. 

RECORDS Margarita Island, Venezuela, from among intertidal barnacles. 

Subfamily Sphaeromatinae H. Milne Edwards, 1840 

DIAGNOSIS Cephalon not fused with pereonite 1. Pereopods 1 and 2 am 

bulatory. Pleopods 4 and 5, endopods having branchial pleats, exopods un- 

pleated, membranous, of two articles. Uropods biramous. 

Key to genera of Sphaeromatinae 

1. Uropodal exopod with outer margin serrate Sphaeroma 

Uropodal exopod with outer margin entire or faintly crenulate 2 

2.

Cymodoce ruetzleri 227 

Pleopod 5, exopod of two articles, distal article with apex and internal mar 

gin covered with fine teeth, anterior surface with long distally toothed boss; 

proximal article with two small toothed bosses at internodistal angle. 6: 

Maxillipedal palp articles 2—4 bearing setigerous lobes. Penial rami elongate, 

separate. Pleon usually more tuberculate than in 9. Uropodal exopod lamel 

lar, shorter than endopod. 9: Mouthparts metamorphosed. Broodpouch 

formed by four pairs of oostegites arising from pereonites 1-4, overlapping in 

midline. Brood housed in five pairs of internal pouches. 

"Cymodoce" barrerae (Boone, 1918) 

Figure 99A,B 

DIAGNOSIS 9 7.5 mm. 9: Body dorsally strongly vaulted, unornamented. 

Frontal lamina distally broadly rounded, lateral shoulders rounded. 

Mouthparts not metamorphosed. Pleotelson anteriorly strongly inflated with 

barest indication of two submedian swellings; posterior margin trilobed, with 

median lobe strong, narrowly rounded, outer lobes much smaller and ventral 

to median lobe. Uropodal endopod distally obliquely truncate; exopod dis 

tally acute. 

RECORDS Cabanas, Cuba. 

REMARKS This species is known only from the nonovigerous female 

holotype. Loyola e Silva (1960) placed the species in Cymodoce, based on a 

female specimen from Brazil. As the mouthparts are not metamorphosed, 

this does not agree with the present concept of Cymodoce, but with neither 

ovigerous females nor males available, the correct generic placement cannot 

be determined. 

Cymodoce ruetzleri Kensley, 1984 

Figure 99C-G 

DIAGNOSIS 6 5.0 mm, 9 4.2 mm. 6: Integument with numerous small 

tubercles, becoming densely setose posteriorly. Pleonite 4, posterior margin 

broadly bilobed. Pleotelson bearing pair of strong conical tubercles with 

acute tips, each tubercle flanked by low rounded tubercle; apex trilobed, 

outer lobes triangular, acute, sharp spine at base of incision, median lobe 

apically blunt. Uropodal exopod apically acute, oval in cross section, endo 

pod and sympod fused, somewhat flattened, apex triangular with strong 

tooth. 9: Pleotelson with two conical apically acute tubercles, apex barely 

notched, with short rounded lobe slightly offset from posterior margin. Both


A 

Figure 99. "Qymodoce" barrerae: A, $; B, frontal lamina. Cymodoce ruetzleri: C, pleopod 4; G, pleopod 5. 

uropodal rami flattened; exopod with tiny apical tooth, endopod distally 

truncate-rounded, with small mediodistal tooth. 

RECORDS Carrie Bow Cay, Belize, 0.5-13 m; in algal clumps, reef crest 

rubble, and seagrass flats. 

E

Exosphaeroma Stebbing, 1900 

Exosphaeroma alba 229 

DIAGNOSIS Maxillipedal palp articles 2-4 produced medially into lobes. 

Pereonites 6 and 7 dorsally unarmed. Pleopod 3, exopod biarticulate. 6: 

Penes short, separate. Copulatory stylet of pleopod 2 elongate, slender. 

Pleotclson lacking strong apical notch. 9: Mouthparts not metamorphosed. 

Broodpouch of three pairs of oostegites on pereonites 2-4; oostegites short, 

not reaching midline. Brood held in four pairs of internal pouches. 

Key to species of Exosphaeroma 

1. Pleotelson with posterior margin entire, evenly convex 2 

Pleotelson with posterior margin faintly notched or trilobed 3 

2. Frontal lamina with length less than 1.5 times greatest width diminuta 

Frontal lamina with length almost two times greatest width 

productatelson 

3. Pleotelson with posterior margin faintly trilobed, and with three low 

rounded tubercles anteriorly yucatanum 

Pleotelson with posterior margin faintly notched 4 

4. Pleotelson posteriorly broadly notched; two rounded submedian 

tubercles on inflated midregion antillense 

Pleotelson with faint narrow notch posteriorly; lacking dorsal tubercles 

Exosphaeroma alba Menzies and Glynn, 1968 

Figure 100A-C 

DIAGNOSIS 8 2.0 mm, 9 2.3 mm. Frontal lamina anteriorly broadly 

rounded, basally slightly wider than midlength. Pleotelson similar in 8 and 

9; anterodorsally inflated and unornamented, posteriorly tapering to slight 

median notch, seen in dorsal view. Uropodal rami distally shallowly serrate, 

exopod 2.5 times longer than wide. 

RECORDS Puerto Rico, intertidal to 0.5 m; in algae on rocks, and under 

Chiton tuberculatus and C. marmoratus. 

alba


Figure 100. Exosphaeroma alba: A; B, frontal lamina; Cy uropod. Exosphaeroma 

antillense: D; E, frontal lamina; F, uropod. Exosphaeroma diminuta: G; H, frontal 

lamina; /, uropod. Exosphaeroma productatelson: J; K> frontal lamina; L, uropod. 

Exosphaeroma yucatanum: My pleon (from Richardson, 1905). 

Exosphaeroma antillense Richardson, 1912d 

Figure 100D,F 

DIAGNOSIS 9 5.0 mm. Frontal lamina anteriorly tapering to subacute apex. 

Pleotelson with two broadly subconical submedian tubercles on inflated an-

Exosphaeroma yucatanum 231 

terior area; posterior margin subtruncate to very faintly emarginate. Uropo 

dal exopod distally crenulate, length slightly more than twice greatest width; 

endopod with faint distal notch. 

RECORDS Montego Bay, Jamaica. 

REMARKS The single ovigerous female holotype is the only known specimen 

of this species. The overlapping oostegites suggest that this may not be an 

Exosphaeroma. 

Exosphaeroma diminuta Menzies and Frankenberg, 1966 

Figure 100G-I 

DIAGNOSIS 8 2.2 mm. Frontal lamina widest at midlength, anteriorly 

truncate-rounded. Pleotelson with posterior margin broadly rounded. 

Uropodal rami not quite reaching pleotelsonic apex; exopod margin distally 

crenulate. 

RECORDS Chesapeake Bay to Florida; Venezuela; sand dwelling, intertidal 

and shallow subtidal. 

Exosphaeroma productatelson Menzies and Glynn, 1968 

Figure 100J-L 

DIAGNOSIS 6 2.5 mm, 9 1.5 mm. Sexes essentially similar. Frontal lamina 

widest at midlength, where slight shoulder apparent, anteriorly broadly 

rounded, 1.6 times longer than wide. Pleotelson unornamented, anteriorly 

inflated, posterior margin entire, evenly convex. Uropodal exopod distally 

shallowly serrate, almost four times longer than wide; endopod wider than 

exopod. Broad lateral patches of pigment on pleotelson in both sexes. 

RECORDS Puerto Rico, intertidal to 0,5 m, in algae on rocks; Texas, Gulf of 

Mexico. 

Exosphaeroma yucatanum (Richardson, 1901) 

Figure 100M 

DIAGNOSIS Frontal lamina anteriorly tapering from widest point to sub 

acute apex, proximally narrower than at midlength. Pleotelson posteriorly 

obscurely trilobed, median lobe narrowly rounded, longest; three low 

rounded tubercles on pleotelson in anterior region.


RECORDS Cape Catoche, Yucatan, Mexico, 48 m. 

REMARKS This species was described from a single specimen which has 

since been lost. The true generic placement of this species is thus undeter 

mined and full description awaits the finding of more material. 

Harrieta Kensley, 1987c 

DIAGNOSIS 9 with mouthparts metamorphosed. Broodpouch of three pairs 

of oostegites on pereonites 2—4, overlapping in midline; brood held in five 

pairs of internal pouches. Uropodal rami subequal, lamellar in 9, exopod 

twice length of endopod and oval in cross section in 8. Pleopod 2 in 6 with 

copulatory stylet articulating basally on endopod, curved, barely reaching 

apex of endopod. Penes basally fused, rami slender, elongate, tapering. 

Harrieta faxoni (Richardson, 1905) 

Figure 101 A,B 

DIAGNOSIS 6 6.0 mm, 9 6.5 mm. 6: Frontal lamina with broad slightly 

convex anterior margin. Two low rounded submedian tubercles on cephalon 

near posterior margin. Two rounded submedian tubercles on last pleonite. 

Pleotelson anteriorly inflated with two submedian tubercles; posterior mar 

gin trilobed. 9: Essentially similar to 69 but posterior margin of pleotelson 

less markedly trilobed, with median lobe longer, and uropodal rami subequal 

in length. 

RECORDS Florida to Texas, Gulf of Mexico, intertidal and subtidal in 

Thalassia, Halodule, and Syringodium seagrass beds, in salinities of 7%o to 36%o. 

Sphaeroma Bosc, 1802 

DIAGNOSIS Maxillipedal palp with three distal articles poorly developed, 

lacking lobes; fringe of robust plumose setae with swollen bases on internal 

margin of endite; distal margin of endite with simple setae. Pereopods 1—3 

with plumose setae on ischium and merus. Posterior margin of pleotelson 

entire, similar in 6 and 9. Pleopod 3, exopod uniarticulate. Uropodal ex 

opod with outer margin serrate. Able to conglobate. 6: Penes short, 

rounded, separate. Pleopod 2, copulatory stylet articulating basally on endo 

pod, slender, reaching well beyond rami. 9: Mouthparts not meta-

Figure 101. Harrieta faxoni: A, 6; B, pie 

Sphaeroma terebrans; E, Sphaeroma walkeri. 

B 

E


morphosed. Three pairs of overlapping oostegites arising from pereonites 2- 

4 (but S. terebrans has anterior pair rudimentary). 

REMARKS The genus Sphaeroma is one of the few sphaeromatids in which the 

number of oostegites varies, from the diagnostic three pairs, through two 

normal pairs (as in S. terebrans), to having the oostegites completely absent 

(as in S. annandalei). 

Jacobs (1987) has provided a useful reevaluation of the European, Medi 

terranean, and northwest African species of Sphaeroma and related genera. 

Key to species of Sphaeroma 

1. Pleotelson posteriorly bluntly triangular, with 4 strong anterior 

tubercles 

terebrans 

Pleotelson posteriorly broadly rounded 2 

2. Pleotelson dorsally smooth or with few low tubercles quadridentata 

Pleotelson dorsally with numerous strong tubercules walkeri 

Sphaeroma quadridentata Say, 1818 

Figure 101C 

DIAGNOSIS 6 11.0 mm, 9 8.0 mm. Pleotelson anteriorly inflated, some 

times with few low rounded tubercles, posteriorly flattened to concave; pos 

terior margin entire, broadly rounded. 

RECORDS New England to Florida; Gulf of Mexico, intertidal to 1 m, often 

in pilings and partially submerged dead tree trunks, and commonly associ 

ated with barnacles. 

Sphaeroma terebrans Bate, 1866 

Figure 10 ID 

DIAGNOSIS 6 10.0 mm, 9 11.5 mm. Pereonite 7 with pair of submedian 

and pair of lateral tubercles. Dorsal pleon densely tuberculate. Posterior 

pleonite with pair of submedian acute tubercles. Pleotelson anteriorly with 

submedian pair and lateral pair of tubercles, posteriorly rounded-triangular.

FLABELLIFERA • TRIDENTELLIDAE 235 

RECORDS Virginia to Florida; Belize; Cuba; Venezuela to Brazil; Gulf of 

Mexico. 

Nigeria, east coast of southern Africa, India, Sri Lanka, Thailand, Indo 

nesia, Philippines, Australia. 

REMARKS There is no agreement on whether this species is synonymous 

with S. destructor Richardson, 1897. This latter (if distinct) bores into wood 

pilings in estuarine waters, while S. terebrans is found in the prop roots of the 

red mangrove tree, Rhizophora mangle. In this habitat, the isopods are inter 

preted either as being destructive agents (e.g., Rehm and Humm, 1973) or as 

promoting increased root growth (Simberloff et al., 1978). It is unlikely that 

the bored wood itself is a source of food for the isopods; rather, as with the 

genus Limnoria, the food is probably detritus or fungi and bacteria growing on 

the wood fragments in the burrows or on the setae of the appendages. 

Sphaeroma walkeri Stebbing, 1905 

Figure 101E 

DIAGNOSIS 6 9.5 mm, 9 10.0 mm. Pereonites 3—7 with transverse row of 

large rounded tubercles. Last pleonite with row of prominent tubercles and 

smaller scattered tubercles laterally. Pleotelson anteriorly inflated, posteri 

orly concave and cuplike, with four irregular longitudinal rows of large tu 

bercles plus many small scattered tubercles. Posterior margin rounded, en 

tire to irregularly crenulate. Uropodal endopod with several rounded 

tubercles on dorsal surface; exopod with row of smaller tubercles on ventral 

surface. 

RECORDS Probably pan-tropical. Florida to Puerto Rico, intertidal. 

Family Tridentellidae Bruce, 1984 

DIAGNOSIS Eyes well developed. Pereonites 2—7 with distinct coxae. Pleon 

consisting of five free pleonites plus pleotelson. Mandible with acute incisor; 

lacinia mobilis absent; molar present; palp of three articles. Maxilla 1, outer 

ramus styliform with three to five strong terminal spines, and several short 

recurved subapical spines. Maxilla 2 uniramous, biarticulate, bearing small 

sometimes tridentate spines or scales distally. Maxillipedal palp of five arti 

cles; endite slender, lamellar, usually lacking coupling hooks.

236 FLABELLIFERA • TRIDENTELLIDAE 

Tridentella Richardson, 1905 

DIAGNOSIS Body dorsally often bearing spines, tubercles, or carinae, more 

developed in 8 than in 9. Frontal lamina narrow, pentagonal. Antcnnular 

peduncle of three articles; antennal peduncle of five articles. Mandibular 

molar weakly sclerotized. Pereopods 1-3 weakly prehensile; pereopods 4-7 

ambulatory. Copulatory stylet of pleopod 2 rodlike, arising proximally on 

mesial margin of endopod. Pleopod 5 endopod lacking marginal setae. 

REMARKS Delaney and Brusca (1985) provide useful taxonomic and dis 

tributional comments on the family Tridentellidae. 

Tridentella virginiana (Richardson, 1900b) 

Figure 102 

DIAGNOSIS 8 9.5 mm, ovigerous 9 9.5-11.0 mm. Cephalon and pereon 

dorsally smooth, pleon minutely granular. Uropodal rami with distal mar 

gins faintly dentate, apically narrowly rounded, endopod wider and slightly 

longer than exopod. Pleotelson basally wider than middorsal length; pos 

terior margin broadly rounded to subtruncate. 

RECORDS Nova Scotia to Florida; off Georgia, 550 m; Gulf Stream off Key 

West, 220 m. 

Suborder Gnathiidea Leach, 1814 

DIAGNOSIS Eyes usually well developed, rarely on short lateral processes, 

occasionally absent. Mandibles in 8 greatly enlarged, projecting anteriorly 

from cephalon, not used in feeding. Mandibles lacking in 9. Mouthparts of 

praniza larva styliform, with acute mandibles projecting anteriorly (see Fig 

ure 103D). Pereopod 1 modified, forming second pair of broad opercular 

maxillipeds covering mouthparts, referred to as pylopods. Pereopods 2—6 

ambulatory. Pereonite 7 reduced, lacking pereopod. Pleonites separate, nar 

rower than pereon. Uropods lateral, rami lamellar, forming tailfan with 

telson. Praniza larva with pereonites 4-6 enlarged, sometimes inflated. 9 

with pereonites 4-6 greatly inflated, forming broodpouch for internally 

brooded eggs (see Figure 103E). 

REMARKS The gnathiideans are entirely marine, most described species 

being from shallow waters. The males and females are frequently found in 

association with sponges and do not feed. The praniza larva is an efficient 

swimmer and has been recorded from shallow-water plankton, but is more

GNATHIIDEA 237 

Figure 102. Tridentella virginiana: A, 9; B, pereopod 1; C, maxilla 1; D, mandible; 

Ey maxilliped; F, maxilla 2. 

frequently encountered as a fish parasite, the favored site for sucking the 

host's blood being in the nares. Upton (1987a, 1987b) has shed light on the 

unusual life history of at least one gnathiid genus, Paragnathia. 

The taxonomy of the Gnathiidae is based almost entirely on males, the 

praniza and females of most species being remarkably similar.

238 GNATHIIDEA • GNATHIIDAE 

Family Gnathiidae Harger, 1879 

DIAGNOSIS As for the suborder Gnathiidea. 

Gnathia Leach, 1814 

DIAGNOSIS In addition to features mentioned in diagnosis of suborder: Eyes 

present in most species. Pylopod with two small articles distal to broad oper 

cular article 2, terminal article minute. 

Key to species of Gnathia (6 only) 

1. Anterior margin of cephalon with medial process or slightly convex . . 2 

Anterior margin of cephalon concave or lacking medial process 9 

2. Anterior margin of cephalon broadly triangular, projecting, with small 

lateral teeth triospathiona 

Anterior margin of cephalon not triangular and projecting 3 

3. Cephalon and two free anterior pereonites dorsally granular 4 

Cephalon and two free anterior pereonites smooth 5 

4. Lobe of outer margin of mandible notched; pereonite 5 twice wider 

than middorsal length velosa 

Lobe of outer margin of mandible rounded; pereonite 5 1.5 times wider 

than middorsal length 6 

5. Anterior margin of cephalon with distinct medial process virginalis 

Anterior margin of cephalon barely convex, lacking medial process 

rat hi 

6. Inner proximal lobe of mandible distinct 7 

Inner proximal lobe of mandible indistinct samariensis 

7. Inner proximal lobe of mandible entire 8 

Inner proximal lobe of mandible with rounded toothlike marginal 

structures Johanna 

8. Pereonites 3-5 poorly defined puertoricensis 

Pereonites 3-5 clearly defined magdalenensis 

9. Anterior margin of cephalon concave, lacking projections gonzalezi 

Anterior margin of cephalon with four projections beethoveni

Gnathia beethoveni Paul and Menzies, 1971 

Figure 103 A 

Gnathia magdalenensis 239 

DIAGNOSIS 6 3.0 mm. Anterior margin of cephalon with two low tubercles 

flanking shallow medial notch plus slightly larger pair of lateral tubercles. 

Cephalon lacking dorsal tubercles. Pereonite 5 1.5 times wider than middor- 

sal length. Uropodal endopods reaching beyond telsonic apex. 

RECORDS Off Venezuela, 95 m. Colombia. 

Gnathia gonzalezi Miiller, 1988 

Figure 103B 

DIAGNOSIS 6 2.0 mm. Body smooth. Anterior margin of cephalon concave. 

Pereonites 3—5 distinct; pereonite 5 2.5 times wider than middorsal length. 

Cutting margin of mandible with four or five low rounded teeth. 

RECORDS Colombia, 30 m. 

Gnathia Johanna Monod, 1926 

Figure 103C 

DIAGNOSIS 8 2.1 mm. Anterior margin of cephalon medially convex be 

tween pair of submedian tubercles. Pereonites 4 and 5 poorly separated. 

Proximomedial lobe of mandible having four or five rounded crenulations, 

with tiny seta between adjacent crenulations. 

RECORDS U.S. Virgin Islands, 29-46 m; Colombia. 

Gnathia magdalenensis Miiller, 1988 

Figure 103D 

DIAGNOSIS 6 3.0 mm. Anterior margin of cephalon with three tubercles, 

median tubercle slightly shorter than submedian pair. Cephalon with few 

scattered low granulations dorsally. Pereonite 5 about 1.5 times wider than 

middorsal length. Proximomedial lobe of mandible entire. 

RECORDS Carrie Bow Cay, Belize, intertidal; Colombia, 18 m.

Figure 103. Gnathia beethhoveni: A, 6. Gnathia gonzalezi: B, 6 (after Miiller, 1988). 

Gnathia Johanna: C,

Gnathia puertoricensis Menzies and Glynn, 1968 

Figure 103E-G 

Gnathia triospathiona 241 

DIAGNOSIS 6 3.0 mm, ovigerous $ 1.8 mm. Anterior margin of cephalon 

having three tubercles between mandibular bases, medial tubercle narrower 

than submedian pair. Dorsal integument finely granular, with coarser gran 

ules mediodorsal to eye. Pereonites 4 and 5 indistinctly separated. Mandible 

lacking proximomedial lobe. 

RECORDS Carrie Bow Cay, Belize, intertidal to 2 m; Puerto Rico, intertidal; 

Cuba. 

Gnathia rathi Kensley, 1984 

Figure 104 A 

DIAGNOSIS 6 1.9 mm, ovigerous 9 2.2 mm. Frontal margin faintly convex 

to straight between single low lateral tubercle mesial to mandibular bases. 

Dorsal integument of cephalon and anterior two free pereonites coarsely 

granular. Lateral margins of telson faintly denticulate. Pereonites 4 and 5 

poorly separated. 

RECORDS Carrie Bow Cay, Belize, 1-36 m. 

Gnathia samariensis Miiller, 1988 

Figure 104B 


median tubercle slightly shorter than submedian pair; dorsal integument 

smooth. Pereonites 4 and 5 well differentiated; pereonite 5 about 2.2 times 

wider than middorsal length. Mandible lacking proximomedial lobe. 

RECORDS Colombia. 

Gnathia triospathiona Boone, 1918 

Figure 104C 

DIAGNOSIS 6 8.8 mm. Anterior margin of cephalon with broad-based tri 

angular projection bearing three low teeth; deep V-shaped depression pos 

terior to anterior margin, with low flanking granulations. 

RECORDS Off Key West, in Gulf Stream, 218 m.

Figure 104. Gnathia rathi: A, 6; Gnathia samariensis: B, 6 (after Miiller, 1988); 

Gnathia triospathiona; C, 6; Gnathia virginalis: D, 6; Gnathia velosa: E, 6 (after 

Miiller, 1988). 

E

Gnathia velosa Muller, 1988 

Figure 104E 

MIGROGERBERIDEA • MIGROGERBERIDAE 243 


median tubercles slightly shorter and narrower than submedian pair. Dorsal 

integument of cephalon and anterior three pereonites granular. Pereonite 5 

about 2.5 times wider than middorsal length. Lateral lobe of mandible 

notched. 

RECORDS Colombia. 

Gnathia virginalis Monod, 1926 

Figure 104D 


median tubercles slightly longer than submedian pair. Dorsal integument of 

cephalon and anterior three pereonites granular. Pereonite 5 about 1.7 times 

wider than middorsal length. Lateral lobe of mandible rounded. 

RECORDS U.S. Virgin Islands, 29 m; Colombia. 

Suborder Microcerberidea Lang, 1961 

DIAGNOSIS Cephalon free. Mandibles with reduced palp, or lacking palp. 

Maxillipedal palp of five articles. Pereon of seven free segments. Pereopod 1 

subchelate; pereopods 2—7 ambulatory. Pleon of two free pleonites plus 

pleotelson. Pleopod 1 in 6 usually absent. Pleopod 2 modified for copulation. 

Pleopod 3 uniramous, opercular. Pleopod 4 biramous. Pleopod 5 reduced. 

Uropods usually uniramous or biramous. 

Family Microcerberidae Karaman, 1933b 

DIAGNOSIS Eyes absent. Body elongate, slender. Antennular peduncle of 

three articles; antennal peduncle of six to eight articles. Mandibular palp of 

single article; molar reduced to single stout fringed spine. Maxilla 2 reduced 

to single ramus bearing two distal fringed lobes. Pereopods 2—7 ambulatory, 

dactyli biunguiculate. 

REMARKS The species of the Microcerberidae are all very small (less than 2 

mm total length) and are most often found in interstitial habitats. They have 

been recorded from marine, brackish, and freshwater environments.

244 MICROCERBERIDEA • MICROCERBERIDAE 

The microcerbcrideans were often classified with the Anthuridea, mainly 

because of the similarity in body shape. Wagele (1983) however, has con 

vincingly demonstrated the asellotan affinities of the group. 

Key to genera of Microcerberidae 

1. MaxilHpedal palp articles 2 and 3 enlarged; basis of pereopods lacking 

spinous process Yvesia 

MaxilHpedal palp articles 2 and 3 not markedly enlarged; basis of 

pereopods with spinous process Microcerberus 

Microcerberus Karaman, 1933b 

DIAGNOSIS MaxilHpedal palp articles 2 and 3 not enlarged. Articles 2 and 3 

of antennal peduncle with spinous process. Basis of pereopods with spinous 

process. Propodus of pereopod 2 with two denticulate proximal spines. 

Microcerberus syrticus Kensley, 1984 

Figure 105A-E 

DIAGNOSIS (5 1.1 mm, 9 1.1 mm. Tergal lobes of pereonites 2—4 rounded. 

Apical lobe of 6 pleopod 2 acute. 

RECORDS Carrie Bow Cay, Belize, interstitial in intertidal sand bar. 

REMARKS In addition to M. syrticus, six species of Microcerberus have been 

recorded from the Caribbean area: M. littoralis Chappuis and Delamare De- 

boutteville, 1956, from the Bahamas; M. minutus Coineau and Botosaneanu, 

1973, from Cuba; M. mirabilis Chappuis and Delamare Deboutteville, 1956, 

from the Bahamas; M. nunezi Coineau and Botosaneanu, 1973, from Cuba; 

M. renaudi Chappuis and Delamare Deboutteville, 1956, from the Bahamas; 

M. simplex Coineau and Botosaneanu, 1973, from Cuba. The reader is re 

ferred to the original descriptions for separation of the species.

\ 

Figure 105. Microcerberus syrticus: A, 8; B, pereopod 1; C, maxilliped; D, pereopod 

2; Ey pleopod 2 8. Yvesia striata (from Coineau and Botosaneanu, 1973): F9 

maxilliped; G, pereopod 1; H, pereopod 2. 

G

246 MICROCERBER1DEA • ONISCIDEA 

Yvesia Coineau and Botosaneanu, 1973 

Yvesia striata Coineau and Botosaneanu, 1973 

Figure 105F-H 

DIAGNOSIS 9 1.6 mm. Antennal peduncular articles 2 and 3 smooth, lack 

ing spinous processes. Maxillipedal palp articles 2 and 3 enlarged. Bases of 

pereopods unarmed, lacking spinous processes. Propodus of percopod 1 with 

single smooth proximal spine. Body having longitudinal ventrolateral striae. 

RECORDS Oriente, Cuba, interstitial on beach. 

Suborder Oniscidea Latreille, 1803 

DIAGNOSIS Compound eyes usually present. Antennules usually very short. 

Antennae with 4- or 5-articulate peduncle; flagellum varying from few arti 

cles to multiarticulate. Mandibular palp present. Distal articles of maxillipe 

dal palp often reduced. Coxae of pereonites 1—7 usually distinct, expanded. 

Pleopods respiratory, often with pseudotrachea; 6 with pleopod 2, and 

sometimes pleopod 1 as well, modified for copulation. Uropods terminal or 

subterminal with terete rami, or ventral and opercular, with reduced rami. 

REMARKS The Oniscidea includes all the isopods that have successfully in 

vaded the terrestrial environment. While still in some degree reliant on exter 

nal moisture, their morphological and behavioral adaptations have allowed 

them to live in almost all terrestrial habitats, from hot, dry deserts, through 

tropical rainforests and grasslands, to cold-temperate niches. Several forms 

have successfully inveigled themselves into termite or ant colonies, where 

with varying degrees of morphological adaptations they take advantage of 

the security of these habitats. A small number of species have evolved to live 

in more constantly wet habitats. Several species may be found in the marine 

intertidal, either living in and under piles of decomposing litter along the 

high-tide line, digging into beach sand, or sheltering in the damp cracks and 

crevices of rocky shores. A few may also be found in mangrove swamps. 

A breakdown of families, genera, and species is not provided for this sub 

order, but those few species that are commonly encountered in intertidal 

habitats are dealt with individually. Schultz (1974, 1984) records several 

oniscidean isopods from the Caribbean area.

Key to genera and species of littoral Oniscidea 

ONISCIDEA 247 

1. At least one uropodal ramus reaching well beyond outline of body ... 5 

Uropodal rami very short, not reaching beyond outline of body 2 

2. Uropods ventral, not visible in dorsal view 3 (Tylos) 

Uropods visible in dorsal view Armadilloniscus ninae 

3. Ventral extensions of pleonite 5 meeting in midline Tylos niveus 

Ventral extensions of pleonite 5 not meeting in midline 4 

4. Ventral extensions of pleonite 5 very short, obsolete Tylos wegeneri 

Ventral extensions of pleonite 5 well separated Tylos marcuzzii 

Ventral extensions of pleonite 5 just falling short of meeting in midline 

Tylos latreillei 

5. Uropodal rami both elongate, subequal 6 (Ligia) 

Uropodal rami very unequal in length 8 

6. Propodus of 6 pereopod 1 with distal rounded lobe Ligia exotica 

Propodus of 6 pereopod 1 lacking rounded lobe 7 

7. Apex of 6 pleopod 2 club shaped Ligia olfersii 

Apex of 6 pleopod 2 bifid Ligia baudiniana 

8. Antennal flagellum of two articles Rhyscotus texensis 

Antennal flagellum of three articles 9 (Vandeloscia) 

9. Endopod of 6 pleopod 1 with large scalelike subapical process 

Endopod of 6 pleopod 1 with small scalelike subapical process 

Armadilloniscus Ul'yanin, 1875 

Armadilloniscus ninae Schultz, 1984 

Figure 106A 

Vandeloscia riedli 

Vandeloscia culebrae 

- - - - - , 

DIAGNOSIS 8 3.2 mm, 9 4.1 mm. Uropodal sympod expanded to form part 

of body outline; rami set mesial to expanded base, with exopod half length of 

endopod. 

RECORDS Ambergris Cay, Belize; under damp objects along beach drift 

line.

Figure 106. A, Armadilloniscus ninae. Ligia baudiniana: B; C} 6 pleopod 2 endopod. 

Ligia exotica: D, dactylus and propodus of pereopod 1; E, 6 pleopod 2 endopod. 

Ligia olfersii: F, 6 pleopod 2 endopod. G, Rhyscotus texensis. Tylos latreillei: H, 

ventral pleon. Tylos marcuzzi: I, ventral pleon (from Schultz, 1984). Tylos niveus: J, 

lateral view; K} ventral pleon. Tylos wegeneri: L} ventral pleon. Vandeloscia culebrae: 

M, apex of pleopod 1 endopod. N, Vandeloscia riedli.

Ligia Fabricius, 1798 

Ligia baudiniana H. Milne Edwards, 1840 

Figure 106B,C 

Rhyscotus texensis 249 

DIAGNOSIS 6 and 9 up to 22 mm. Antennal flagellum elongate, multiar- 

ticulate. Apex of 6 pleopod 2 bifid, with lateral lobe longer and more slender 

than mesial lobe. Uropods inserted terminally on pleotelson; sympods 

elongate-cylindrical; rami slender, elongate, subequal. 

RECORDS Bermuda; Bahamas; U.S. Virgin Islands; Antigua; Carrie Bow 

Cay, Belize; Bonaire; Aruba; Trinidad; Tobago; Gulf of Mexico. 

REMARKS As is typical in the genus Ligia, this species may be seen on rocks 

i 

and sea walls, as well as piles of drift debris at low tide. When disturbed, they 

run rapidly, to shelter in damp crevices and hollows. 

Ligia exotica Roux, 1828 

Figure 106D,E 

DIAGNOSIS 6 28.5 mm, ovigerous 9 32.0 mm. Propodus of 8 pereopod 1 

with rounded lobe on inner distal surface. Apex of 6 pleopod 2 club shaped, 

convoluted. 

RECORDS New Jersey to Uruguay; Indo-Pacific. 

Ligia olfersii Brandt, 1833 

Figure 106F 

DIAGNOSIS 6 20.0 mm, ovigerous 9 24.0 mm. Apex of 6 pleopod 2 simple, 

club shaped. 

RECORDS South Florida to Rio de Janeiro, Brazil; Texas, Gulf of Mexico. 

Rhyscotus Budde-Lund, 1885 

Rhyscotus texensis (Richardson, 1905) 

Figure 106G 

DIAGNOSIS 6 and 9 6.0 mm. Antennal flagellum of two unequal articles. 

Uropodal endopod at least twice length of exopod, inserted distally on base, 

exopod inserted distally on base. Pleotelson broadly triangular. 

RECORDS Carrie Bow Cay, Belize; Texas, Gulf of Mexico.

250 ONISCIDEA 

Tylos Latreille, 1826 

Tylos latreillei Audouin, 1826 

Figure 106H 

DIAGNOSIS 6 12.8 mm, 9 13.0 mm. Ventral extensions of pleonite 5 not 

meeting in midline. 

RECORDS Bermuda; Cuba; Puerto Rico; Honduras. 

Mediterranean. 

Tylos marcuzzii Soika, 1954 

Figure 1061 

DIAGNOSIS 6 6.6 mm. Antennal flagellum of four articles. Ventral exten 

sions of pleonite 5 well separated. 

RECORDS Florida Keys; Bahamas; Leeward Islands; Ambergris Cay, Be 

lize; under debris on sand beach drift line. 

Tylos niveus Budde-Lund, 1885 

Figure 106J,K 

DIAGNOSIS 8 11.0 mm., 9 12.0 mm. Antennal flagellum of four articles. 

Ventral extensions of pleonite 5 expanded, medially contiguous. 

RECORDS Bahamas; Florida Keys; Cuba; Dominica; Lesser Antilles; Bo 

naire; Curasao, under piles of decaying mangrove leaves at beach drift line; 

Carrie Bow Cay, Ambergris Cay, Belize, under deep piles of dead plant ma 

terial on beach drift line; Tobago; Panama. 

Rio de Janeiro, Brazil. 

Tylos wegeneri Vandel, 1952 

Figure 106L 

DIAGNOSIS 6 10.5 mm, 9 15 mm. Antennal flagellum of three articles. 

Ventral extensions of pleonites short or nearly absent. Pleonite 5 lacking free 

lateral margins. 

RECORDS Tobago; Venezuela, under decaying beach debris on drift line; 

Trinidad.

Vandeloscia Roman, 1977 

Vandeloscia culebrae (Moore, 1901) 

Figure 106M 

VALVIFERA 251 

DIAGNOSIS 8 5.0 mm, 9 6.1 mm. Tiny lateral tubercles present on per 

eonites. Endopod of pleopod 1 in 8 with small scalelike subapical process on 

laterally folded tip. 

RECORDS Florida Keys; U.S. Virgin Islands; Puerto Rico; under decaying 

plant material, especially Thalassia testudinea accumulated along beach drift 

line, 

Vandeloscia riedli (Strouhal, 1966) 

Figure 106N 

DIAGNOSIS 8 5.9 mm, 9 6.0 mm. Tiny obsolete tubercles present on all 

pereonites. Endopod of 8 pleopod 1 with large scalelike subapical process on 

laterally folded tip. 

RECORDS Yucatan Peninsula, Mexico; Ambergris Cay, Belize; Barbuda; 

Venezuela; Brazil. 

Gulf of Aqaba; Red Sea; northeastern coast of Africa; Madagascar; Bay of 

Bengal; St. Helena Is. 

Suborder Valvifera Sars, 1882 

DIAGNOSIS Pereopodal coxae, in addition to usual dorsal coxal plates, ex 

panded ventrally to form plates. Penes situated ventrally on articulation be 

tween pereon and pleon, or on pleonite 1. Pleonites and pleotelson variously 

fused. Uropods forming operculum covering over pleopods. 

Key to families of Valvifera 

1. Body often geniculate, flexed between pereonites 4 and 5; anterior 

pereopods setose for feeding, posterior pereopods ambulatory 

Arcturidae 

Body never geniculate; all pereopods ambulatory Idoteidae

252 VALV1FERA • ARGTUR1DAE 

REMARKS Of the six families in the suborder, only two have been recorded 

in the Caribbean area, the Idoteidae and the Arcturidae. 

Family Arcturidae Sars, 1897 

DIAGNOSIS Pereonite 1 either distinct, or completely or imcompletely fused 

with cephalon. Anterior four pairs of pereopods directed anteriorly, usually 

strongly setose; posterior three pairs of pereopods ambulatory, used for cling 

ing to substrate. Body often bent between pereonites 4 and 5. Uropods usu 

ally biramous, with minute endopod concealed by larger exopod. Pleonites 

variously fused with pleotelson. Sexual dimorphism often marked. 

REMARKS Menzies and Kruczynski (1983) described three species of 

arcturids from the west coast of Florida, in depths of 55—73 m: Arcturella 

spinata, Arcturella bispinata, and Edwinjoycea horologium. These species are not 

covered here. 

Key to genera of Arcturidae 

1. Pereonite 1 not fused with cephalon; at least one free pleonite 

Pereonite 1 fused with cephalon; pleonites fused with pleotelson 

Astacilla Cordiner, 1793 

Thermarcturus 

Astacilla 

DIAGNOSIS Antennae at least half length of body. Pereopod 1 with strong 

terminal claw on dactylus. Pereopods 2—4 lacking dactyli. Endopod of 

Key to species of Astacilla 

1. Body integument lacking ornamentation cymodocea 

Body integument with spines or tubercles 2 

2. Pereonite 4 in 6 and 9 with strong middorsal tubercle; pairs of spines 

lacking on pereonites regina 

Pereonite 4 lacking strong middorsal tubercle; pairs of spines on all 

pereonites lasallae

Astacilla regina 253 

pleopod 1 6 with median notch and three specialized setae; pleopod 2 copu- 

latory stylet apically trifid. Pereonite 4 considerably longer than preceeding 

or following pereonite. 

Astacilla cymodocea Menzies and Glynn, 1968 

Figure 107A,B 

DIAGNOSIS 6 6.4 mm, ovigerous 9 9.0 mm. Body cylindrical, ovigerous 9 

with pereonite 4 somewhat bulged, 6 with pereonite 4 elongate-cylindrical. 

Shallow groove marking fusion between cephalon and pereonite 1. Pleonites 

fused with pleotelson, with two incomplete shallow dorsal grooves marking 

lines of fusion anteriorly. Pleotelson lacking any shoulders, posteriorly tape 

red to narrowly rounded apex. 

RECORDS Florida Keys; Puerto Rico, 1.5 m, on Cymodocea sp. seagrass; Car 

rie Bow Cay, Belize, 1—2 m, on Syringodium filiforme seagrass. 

REMARKS In life, A. cymodocea is bright green, blending in with its preferred 

substrate of seagrasses. 

Astacilla lasallae Paul and Menzies, 1971 

Figure 107C 

DIAGNOSIS 9 3.5 mm. Cephalon with large rounded area bearing pair of 

spines; all pereonites and two anterior fused pleonites bearing pair of short 

submedian spines. Pleotelson with strong anterior shoulder, posteriorly tri 

angular, tapering sharply to narrowly rounded apex. 

RECORD Off Venezuela, 95 m. 

REMARKS This species is known only from the small female holotype, and 

until a mature male and ovigerous female are found, it cannot be confidently 

diagnosed. 

Astacilla regina Kensley, 1984 

Figure 107D-G 

DIAGNOSIS 6 6.5 mm, ovigerous 9 7.1 mm. Body strongly tuberculate, 

many tubercles acute. Cephalon with two submedian pairs of acute tuber 

cles; fused pereonite 1 and pereonites 2 and 3 each with single middorsal 

acute tubercle. Pereonite 4 with strong middorsal tubercle situated in ante-

254 VALVIFERA • ARGTURIDAE 

Figure 107. Astacilla cymodocea: A, 8; By 9. Astacilla lasallae: C, 6. Astacilla regina: 

D, 6\ E, 9; F, pereopod 4; G, pereopod 1. Thermarcturus venezuelensis: H, 9 (from 

Paul and Menzies, 1971). 

rior half. Pleotelson with strong lateral shoulder in anterior half, second 

shoulder in posterior half, then tapering to rounded apex. 

RECORDS Carrie Bow Cay, Belize, on forereef slope, 27—36 m; Barbados, 

100—400 m; St. Lucia, 2—3 m, associated with crinoids.

Thermarcturus Paul and Menzies, 1971 

VALVIFERA • IDOTEIDAE 255 

DIAGNOSIS Pereonite 1 not fused with cephalon. Pereonite 4 subequal in 

length to pereonite 3, not markedly elongate. Pereopods 2—4 having dactyli 

but lacking elongate setae. Body cylindrical, flexed between pereonites 4 and 

5. Pleon consisting of two free pleonites plus pleotelson. 

Thermarcturus venezuelensis Paul and Menzies, 1971 

Figure 107H 

DIAGNOSIS 9 4.5 mm. Cephalon, all pereonites, and anterior two pleonites 

each with submedian pair of dorsal tubercles, those on pereonites 2 and 3 

broad and expanded. Pleonite 2 with pair of bulbous lateral swellings, pos 

terior margin triangular. Pleotelson with lateral shoulder anteriorly, posteri 

orly triangular. 


REMARKS Only the holotype (which seems to be lost) is known of this spe 

cies. Considerable uncertainty exists regarding some of the features. 

Family Idoteidae Fabricius, 1798 

Subfamily Idoteinae Dana, 1852 

DIAGNOSIS Flagellum of antenna either multiarticulate; clavate, i.e., with 

large basal articles and with or without one to four reduced distal articles; or 

Key to genera of Idoteinae 

1. Antennal flagellum multiarticulate Idotea 

Antennal flagellum clavate 2 

2. Pereopod 4 reduced, considerably smaller than pereopods 3 or 5 .... 3 

Pereopod 4 not reduced, of similar size to pereopods 3 and 5 

Erichsonella 

3. Pleon consisting of three complete and one incomplete pleonites plus 

pleotelson Cleantioides 

Pleon consisting of two complete and two incomplete pleonites plus 

pleotelson Miratidotea

256 VALVIFERA • IDOTEIDAE 

vestigial. Maxillipedal palp consisting of five or fewer articles. Uropods uni- 

ramous or biramous, rami usually much smaller than sympod. Pleonites 

variously fused with pleotelson; number of fused pleonites often indicated by 

lateral sutures or furrows. 

REMARKS Brusca (1984) has reviewed the phylogeny, evolution, and bio- 

geography of the subfamily Idoteinae, the only one of the five subfamilies 

recorded from the Caribbean. 

Cleantioides Kensley and Kaufman, 1978 

DIAGNOSIS Antennal flagellum a single clavate article. Maxillipedal palp of 

four or five articles. Pereopod 4 somewhat reduced. Uropod uniramous. 

Pleon consisting of three complete and one incomplete pleonites plus 

pleotelson. 

Cleantioides planicauda (Benedict, 1899) 

Figure 108A 

DIAGNOSIS Ovigerous 9 5.5 mm. Body parallel sided. Maxillipedal palp of 

five articles. Pleotelson posteriorly broadly rounded, with obliquely truncate 

subcircular dorsal area in posterior half. 

RECORDS Maryland to Florida; Puerto Rico; Panama; Louisiana, Gulf of 

Mexico, intertidal to 44 m; often in hollow stems and roots of seagrasses, and 

tubes of the polychaete Diopatra cuprea. 

Oaxaca, Pacific Mexico. 

REMARKS Cleantioides planicauda has been recorded only once in the eastern 

Pacific, where it occurs with the more common C. occidentalis (Richardson, 

1899). 

Key to species of Erichsonella 

1. Pereonites with dorsal spines 2 

Pereonites lacking dorsal spines attenuata 

2. Pereonites 1—4 with middorsal and lateral spines floridana 

Pereonites 1-4 with middorsal spines only Jiliformis

Erichsonella 257 

Figure 108. Ay Cleantioides planicauda; B, Erichsonella attenuata 8\ C, Erichsonella 

filiformis 6\ Dy Erichsonella Jloridana 9; E, Idotea balthica 6] F, Idotea metallica $; G, 

Miratidotea bruscai $. 

Erichsonella Richardson, 1901 

DIAGNOSIS Antennal flagellum clavate. Maxillipedal palp of four articles. 

Uropod uniramous. Pleonites completely fused with pleotelson. 

REMARKS Pires (1984) reviewed the genus Erichsonella and did not recog 

nize the subspecies E. filiformis tropicalis Menzies and Glynn, 1968.


Erichsonella attenuata (Harger, 1873) 

Figure 108B 

DIAGNOSIS 6 11.4 mm, ovigerous 9 12.0 mm. Body dorsally smooth. 

Cephalon lacking middorsal elevation. Antcnnulc reaching only slightly be 

yond antennal peduncular article 2. Pleotelson with slight marginal bulge in 

anterior half, indicating ventrolateral articulation of uropod. 

RECORDS Connecticut to Miami; Florida, Mississippi, Texas, Gulf of Mex 

ico; intertidal to 2 m, usually associated with submerged seagrass and algal 

beds. 

REMARKS While not recorded in the Florida Keys, this species does reach 

Miami, and continues into the Gulf of Mexico. 

Erichsonella filifor mis (Say, 1818) 

Figure 108C 

DIAGNOSIS 6 10.5 mm, ovigerous 9 8.2 mm. Body dorsally with bifid tu 

bercle on cephalon, and low rounded middorsal tubercle on pereonites. An- 

tennule reaching midlength of antennal peduncular article 3. Basis of per- 

eopods 2—7 with larges tubercles. Pleotelson with distinct lateral shoulder in 

anterior half. 

RECORDS Connecticut to Florida, shallow infratidal to 55 m; Bahamas; 

Turks and Caicos Islands; Puerto Rico; Quintana Roo, Yucatan Peninsula, 

Mexico, 60-109 m; Florida and Texas, Gulf of Mexico. 

Brazil. 

Erichsonella jloridana Richardson, 1901 

Figure 108D 

DIAGNOSIS Ovigerous 9 10.0 mm. Antennule reaching distal end of anten 

nal peduncular article 3. Cephalon with strong trifid tubercle. Pereonites 1—7 

each with posteriorly directed spine near posterior margin; pereonites 1—4 

each with lateral spine. Basis of pereopods 2-7 smooth. 

RECORDS Florida Keys, intertidal to 2 m; Florida, Gulf of Mexico, interti 

dal mud flats.

Idotea Fabricius, 1798 

Miratidotea 259 

DIAGNOSIS Antennal flagellum multiarticulate. Maxillipedal palp of four or 

five articles. Uropod uniramous. Pleon consisting of two complete and one 

incomplete pleonites plus pleotelson. 

Key to species of Idotea 

1. Posterior margin of pleotelson truncate metallica 

Posterior margin of pleotelson with distinct median lobe balthica 

Idotea balthica (Pallas, 1772) 

Figure 108E 

DIAGNOSIS 6 24.5 mm, ovigerous 9 13.2-23.5 mm. Anterior margin of 

cephalon concave. Cephalon dorsally smooth. Pereonites evenly convex, 

smooth. Posterior margin of pleotelson with rounded median lobe. 

RECORDS Worldwide in tropical to cold-temperate waters, often on floating 

seaweed, from surface to 357 m. 

Idotea metallica Bosc, 1802 

Figure 108F 

DIAGNOSIS 6 30.0 mm, ovigerous 9 22.2 mm. Cephalon with sinuous fur 

row in posterior half. Pereonites 2—4 laterally with rounded convex area close 

to coxae. Posterior margin of pleotelson truncate. 

RECORDS Worldwide in tropical to cold-temperate waters, often on floating 

seaweed, from surface to 200 m. 

Miratidotea Kensley, 1987a 

DIAGNOSIS Antennal flagellum of single clavate article. Maxillipedal palp 

of four articles. Uropod uniramous. Pleon consisting of two complete and two 

incomplete pleonites plus pleotelson.


Miratidotea bruscai Kensley, 1987a 

Figure 108G 

DIAGNOSIS Ovigerous 9 13.0 mm. Body parallel sided. Maxillipedal palp 

of four articles, terminal article very short. Pereopods 1—3 increasing in 

length posteriorly, pereopod 4 reduced, shorter than pereopod 5, and with 

dactylus spinelike, pereopods 5—7 increasing in length. Pleotelson consisting 

of two complete and two incomplete pleonites plus pleotelson; latter with 

broadly rounded posterior margin, and with bifid median process situated 

dorsal to posterior oblique-concave area. 

RECORDS Carrie Bow Cay, Belize, 1.5 m, in hollow root-internodes of sea- 

grass Syringodium filiforme.

Zoogeography 

FAUNAL PROVINCES 

The area under discussion has been divided into several faunal regions or 

provinces, of which the Caribbean, West Indian, and Brazilian are the major 

ones (Briggs, 1974). The extent and boundaries of the provinces have been 

variously defined depending on the group of organisms under discussion. 

Inevitably, zones of overlap exist, but for the purposes of this discussion, the 

following rough limits have been used. 

Brazilian Province: This province stretches from Cape Frio near Rio de 

Janeiro in Brazil to the mouth of the Orinoco River in Venezuela. The out 

flow of freshwater from the major rivers of this region has probably contrib 

uted to the isolation of the Brazilian coral reefs and their associated fauna 

from those of the Caribbean. This isolation is demonstrated by the consider 

able endemism of the Brazilian reef fauna and that of the Caribbean reef 

fauna, with very few species being common to both. 

Caribbean Province: This province has two components, a northern part 

in peninsular Florida, that stretches from around Cape Kennedy on the east 

coast to Tampa or Sanibel Island on the west coast, and a southern compo 

nent that runs from the mouth of the Orinoco River to around Cabo Rojo or 

Tampico on the gulf coast of Mexico. The northern Gulf of Mexico is ex 

cluded from this province and is characterized as being warm-temperate, 

rather than subtropical (Briggs, 1974:66). 

West Indian Province: This includes all the islands of the West Indian 

chain, the Bahamas, and the isolated outrider, Bermuda. The West Indian 

Province closely approaches the Caribbean Province in the Yucatan Penin 

sula to the north, and between Grenada and Trinidad in the south. There is 

also some indication of the isolating effect on the Bahamas of the Florida 

Current through the Straits of Florida. 

It has been suggested, on the basis of the molluscan fauna, that a relict of 

the Neogene Gatunian Province exists around northern Venezuela and Col 

ombia (Petuch, 1982). While several isopod species have been recorded only 

from this area, these are all described in a single paper that covers a very 

small part of this region (Paul and Menzies, 1971). There is as yet too little 

evidence to explore the idea of this relict fauna further. 

261

262 ZOOGEOGRAPHY 

ANALYSIS OF THE ISOPOD FAUNA 

In the following discussion, the West Indian and Caribbean provinces are 

treated as one, the isopod faunas offering little evidence to warrant a separate 

treatment of each. 

It is a truism that for any discussion of the zoogeography of an area to have 

meaning, the true extent of the fauna must be known. With the area under 

review, collecting effort has been uneven, and the true faunal composition of 

many regions is still incompletely known. Obviously, any conclusions based 

on such incomplete data are approximate and subject to revision. Neverthe 

less, certain general patterns or trends emerge when the present isopod fauna 

is broken down into its components. 

The deepwater isopod fauna of the Caribbean (i.e., from deeper than 200 

m) has barely been explored, and little is to be gained from discussing the 

relatively few species known. A list of these deeper dwelling species is in 

cluded (Table 4). 

Although about 280 shallow-water species are covered by this work, cer 

tain categories of species must be excluded, for various reasons, before anal 

ysis can be attempted. Such excluded groups include the species of Oniscidea 

(being essentially terrestrial forms and not part of the marine regime); the 

cymothoid species and the species of Aegidae (being fish parasites for at least 

part of their life history, and whose distribution is complicated by the dis 

tribution and mobility of the hosts); the limnoriids (being wood-borers whose 

distribution is more a function of the distribution of floating wood); and the 

true cave species (which have a history more reflective of the geological his 

tory of the area than of the marine regime). The epicarideans have a distribu 

tion somewhat complicated by the distribution of their crustacean hosts and 

their pelagic epicaridean and cryptoniscan larvae. Nevertheless, the decapod 

hosts of the great majority of species covered here are Caribbean endemics, 

and inclusion of the epicarideans changes very little the overall patterns of 

distribution, as demonstrated by the two figures provided (Figure 109). After 

making these exclusions there remain about 166 species (218 with the epi 

carideans) that can be broken down into the following components (figures 

in brackets include epicarideans): 

1. True Caribbean/Bahamian species—124 species, 74.8% [147, 67.5%]. 

These are the species recorded only from the Caribbean and the Bahamas. 

The term endemic is avoided, as too little is known of the actual distribution 

of many species. Of these species, 86 [87] have been recorded from a single 

locality. 

2. Species occurring south of the discussion area, and extending into 

Brazil—5 species, 3.0% [9, 4.1%]. These low numbers indicate that the

TABLE 4. CARIBBEAN ISOPODS RECORDED FROM DEPTHS GREATER THAN 200 M 

SUBORDER ANTHURIDEA 

Family Paranthuridae 

Neoanthura coeca Menzies, 1956b. South of Jamaica, 1244 m 

SUBORDER ASELLOTA 

Family Dendrotiidae 

Dendrotion hanseni Menzies, 1956b. South of Jamaica, 1244 m 

Family Desmosomatidae 

Desmosoma magnispina Menzies, 1962a. Bay of Panama, 1906 m 

Family Echinothambematidae 

Echinothambema ophiuroides Menzies, 1956a. North of Puerto Rico Trench, 

5104-5122 m 

Family Eurycopidae 

Acanthocope spinosissima Menzies, 1956b. South of Jamaica, 1224 m 

Storthyngura pulchra caribbea (Benedict, 1901). Off Windward Islands, 1256 m 

Storthyngura snanoi Menzies, 1962a. Colombia abyssal plain, 4071 m 

Family Haploniscidae 

Antennuloniscus dirneroceras (Barnard, 1920). North of Puerto Rico Trench, 

5440-5410 m; South Atlantic off South and West Africa, 1400-3921 m; 

off Argentina, 5843 m 

Haploniscus unicornis Menzies, 1956a. North of Puerto Rico Trench, 5104- 

5122 m 

Hydroniscus quadrifrons Menzies, 1962a. North of Puerto Rico Trench, 5271- 

5684 m 

Family Ischnomesidae 

Haplomesus tropicalis Menzies, 1962a. Colombia abyssal plain, 4071 m; off 

South Africa, 2526 m; Mediterranean 

Heteromesus bifurcatus Menzies, 1962a. Colombia abyssal plain, 4071 m 

Ischnomesus armatus Hansen, 1916. North of Puerto Rico Trench, 5494-5477 

m; Davis Straits, 2702 m 

Ischnomesus caribbicus Menzies, 1962a. Off Panama, 1714 m 

Ischnomesus multispinis Menzies, 1962a. Off Panama, 975 m 

Family Janiridae 

Abyssianira dentifrons Menzies, 1956a. North of Puerto Rico Trench, 5104- 

5122 m; off Argentina, 5024-5293 m; off southwest Africa, 4588 m 

Ianirella caribbica Menzies, 1956b. South of Jamaica, 1244 m 

Ianirella vemae Menzies, 1956a. Near Puerto Rico Trench, 5104-5122 m 

Spinianirella serrata Kensley and Heard, 1985. Off Puerto Rico, 350 m 

Family Macrostylidae 

Macrostylis caribbicus Menzies, 1962a. Off Colombia, 2875-2941 m 

Macrostylis minutus Menzies, 1962a. North of Puerto Rico Trench, 5163- 

5494 

{continued)



Macrostylis setifer Menzies, 1962a. North of Puerto Rico Trench, 5477- 

5494 m 

Macrostylis vemae Menzies, 1962a. North of Puerto Rico Trench, 5410— 

5684 m 

Family Mesosignidae 

Mesosignum kohleri Menzies, 1962a. Colombia abyssal plain, 2868-4076 m 

Family Nannoniscidae 

Nannoniscus camayae Menzies, 1962a. Off Panama, 1714 m 

SUBORDER GNATH11DEA 

Family Gnathiidae 

Akidognathia poteriophora Monod, 1926. OffU. S. Virgin Islands, 914 m. 

SUBORDER VALVIFERA 

Family Arcturidae 

Antarcturus annaoides Menzies, 1956b. South of Jamaica, 1244 m 

Arcturus caribbaeus Richardson, 1901. OfTAves Island, 1360 m 

Arcturus purpureus Beddard, 1886. Off Leeward Islands, 900 m 

Note: Records from deep water around Bermuda are not included. 

great area of mixed-salinity waters resulting from the outflow of the Orinoco, 

Amazon, Tocantins, and Parnaiba rivers form an effective barrier to the 

movement of shallow-water isopod species. 

3. Species having an amphi-Panamic distribution—7 species, 4.2% [8, 

3.7%] (Table 5). In spite of the history of immergence and emergence of the 

Isthmus of Panama, this very small amphi-Panamic component in the Carib 

bean isopod fauna suggests that most of this fauna has evolved since the last 

emergence of the late Pliocene. Given the limited mobility of most isopod 

species, the Panama Canal seems to have played a minimal role in contribut 

ing to this component. 

4. Species occurring outside of the western Atlantic (but excluding the 

amphi-Panamic species)—3 species, 1.8% [7, 3.2%]. 

5. The role of the Gulf of Mexico isopod fauna (see Clark and Robertson, 

1982) in the composition of the Caribbean/Bahamian is complex and diffi 

cult to analyze. One hundred and thirteen species of shallow-water isopods 

have been recorded from the Gulf of Mexico (Table 6). This number would 

indicate that many species remain to be recorded in this region. Of these 113 

species, 61 (54%) have also been reported from the Caribbean region. It is 

therefore possible that there exists a true Gulf of Mexico fauna, whose evolu 

tion was perhaps spurred by the relative isolation and reduction of the Gulf

Car 

Car 

Sout 

South 3.0% % 

Panam 4.2% 

Epicaridea excluded 

Panam 3.7% 

Out 3.2% 

North 

Epicaridea included 

5.5% 

ZOOGEOGRAPHY 265 

GoM/N/Car 7.2% 

oM/Car 5.4% 

3.6% 

GoM/N/Car 9.6% 

/Car 6.4% 

Figure 109. Relative proportions of the zoogeographic components of the 

Caribbean isopod fauna, with and without the parasitic Epicaridea. Car, 

Caribbean; Out, extra-western Atlantic; GoM/Car, Gulf of Mexico-Caribbean; 

GoM/N/Car, Gulf of Mexico-Northern-Caribbean; North, northern; Panam, amphi- 

Panamic; South, southern. 

during a low-water stand (100 m below present sea level) during the 

Pleistocene. A significant proportion (about 27 species, 28%) of the Gulf of 

Mexico isopods are known from the eastern coast of the United States north


TABLE 5. SPECIES OF ISOPODS OCCURRING ON BOTH SIDES OF THE ISTHMUS 

OF PANAMA 

*Aega deshaysiana (H. Milne Edwards, 

1840) 

Anopsilana browni (Van Name, 1936) 

Cleantioides planicauda (Benedict, 

1899) 

Excirolana braiiliensis Richardson, 

1912 

Excorallana tricornis (Hansen, 1890) 

*Nerocila acuminata Schioedte and 

Meinert, 1881 

* fish parasite or fish predator 

of Cape Kennedy, which would indicate a significant cooler-water compo 

nent. What proportion of originally Gulf species have spread into the Carib 

bean, and what proportion of Caribbean and temperate east coast species 

have entered the Gulf, cannot yet be assessed, given our incomplete knowl 

edge of the Gulf fauna. Because of this unresolved situation, three categories 

of species have been separated: species ranging from north of Cape Kennedy 

into the Caribbean—6, 3.6% [12, 5.5%]; species occurring in the Gulf of 

Mexico and the Caribbean—9, 5.4% [14, 6.4%]; species occurring north of 

Cape Kennedy, in the Gulf, and in the Caribbean—12, 7.2% [21, 9.6%] The 

conclusion that the fauna of the Gulf of Mexico contains an endemic compo 

nent, a Caribbean component, and a warm-temperate component was also 

reached by Topp and HofT (1972), in an analysis of the pleuronectiform 

fishes of the Gulf. 

THE BAHAMAS 

The Florida Current flowing through the Straits of Florida has been sug 

gested as a factor in reducing the movement of shallow-water fauna between 

peninsular Florida and the Florida Keys on the west and the Bahamas on the 

east (Briggs, 1974). Comparison of the number of isopod species on either 

side of the Straits of Florida (13 from the Bahamas, 50 from southern penin 

sular Florida and the Florida Keys) supports this view. Of the 13 species 

from the Bahamas, only four arc "endemic," three of these being interstitial 

microcerberideans. 

Paradella dianae (Menzies, 1962b) 

Paraleptosphaeroma glynni Buss and 

Iverson, 1981 

Probopyrus pandalicola (Packard, 

1879) 

*Rocinela oculata Harger, 1883 

*Rocinela signata Schioedte and 

Meinert, 1879 

Uromunna reynoldsi Frankenberg and 

Menzies, 1966

TABLE 6. ISOPOD SPECIES OCCURRING IN THE GULF OF MEXICO 

SUBORDER ANTHURIDEA 

*Accalathura crenulata (Richardson, 

1901) 

*Amakusantkura magnified (Menzies 

and Frankenberg, 1966) 

Cyathura polita (Stimpson, 1855) 

Horoloanthura irpex Menzies and 

Frankenberg, 1966 

Kupellonura formosa (Menzies and 

Frankenberg, 1966) 

*Mesanthura Jloridensis Menzies and 

Kruczynski, 1983 

Mesanthura hopkinsi Hooker, 1985 

Mesanthura pulchra Barnard, 

1925 

Paranthura Jloridensis Menzies and 


Ptilanthura tricarina Menzies and 


Skuphonura lindae Menzies and 


*Xenanthura brevitelson Barnard, 

1925 

SUBORDER ASELLOTA 

Carpias Jloridensis Menzies and 


Gnathostenetrioides pugio Hooker, 

1985 

*Joeropsis coralicola Schultz and 

McCloskey, 1967 

*Joeropsis rathbunae Richardson, 

1902 

Mexicope kensleyi Hooker, 1985 

Munnogonium wilsoni Hooker, 1985 

*Pleurocope Jloridensis Hooker, 

1985 

*Santia milleri (Menzies and Glynn, 

1968) 

*Stenetrium stebbingi Richardson, 

1902 


Uromunna hayesi Robertson, 1978 

*Uromunna reynoldsi Frankenberg 

and Menzies, 1966 

SUBORDER EPICARIDEA 

Allodiplophryxus Jloridanus Markham, 

1985 

*Aporobopyrina anomala Markham, 

1973 

*Azygopleon schmitti (Pearse, 1932) 

*Bopyrina abbreviata Richardson, 

1904 

*Bopyrione synalphei Bourdon and 

Markham, 1980 

*Cancricepon choprae (Nierstrasz and 


Dactylokepon sulcipes Adkison, 1982 

Eophryxus subcaudalis (Hay, 1917) 

*Gigantione mortenseni Adkison, 

1984b 

Gigantione uberlackerae Adkison, 

1984b 

*Hemiarthrus synalphei (Pearse, 

1950) 

Hyperphrixus castrensis Markham, 

1985 

*Munidion longipedis Markham, 

1975a 

Ovobopyrus alphezemiotes Markham, 

1985 

Parabopyrella mortenseni (Nierstrasz 


*Parabopyrella richardsonae 

(Nierstrasz and Brender a 

Brandis, 1929) 

Parabopyriscus stellatus Markham, 

1985 

*Probopyria alphei (Richardson, 

1900b) 

Probopyrinella heardi Adkison, 

1984a 

{continued)

268 

TABLE 6. (Continued) 

*Probopyrinella latreuticola (Gissler, 

1882) 

Prodajus cf. bigelowiensis Schultz and 

Allen, 1982 

Pseudione cognata Markham, 1985 

Pseudione upogebiae Hay, 1917 

*Schizobopyrina urocaridis (Richardson, 

1904) 

*Stegophryxus hyptius Thompson, 1902 

*Synsynella choprae (Pearse, 1932) 

*Synsynella deformans Hay, 1917 

Synsynella Integra Bourdon, 1981 

* Urobopyrus processae Richardson, 1904 

SUBORDER FLABELLIFERA 

*Aega deshaysiana (H. Milne Edwards, 

1840) 

*Aega ecarinata Richardson, 1898 

Aega incisa Schioedte and Meinert, 

1879 

Alcirona krebsii Hansen, 1890 

Ancinus depressus (Say, 1818) 

Anilocra acuta Richardson, 1910 

Anilocra laticauda H. Milne Edwards, 

1840 

*Bathynomus giganteus A. Milne 

Edwards, 1879 

*Cassidinidea ovalis (Say, 1818) 

Ceratothoa transversa (Richardson, 

1900b) 

Cirolana borealis Lilljeborg, 1851 

*Cirolana obtruncata Richardson, 1901 

* Cirolana parva Hansen, 1890 

Conilera cylindracea (Montagu, 1804) 

*Cymothoa caraibica Bovallius, 1885 

*Cymothoa excisa Perty, 1833 

*Cymothoa oestrum (Linnaeus, 1793) 

*Cerceis carinata Glynn, 1970 

*Eurydice convexa Richardson, 1900b 

Eurydice littoralis (Moore, 1901) 

*Eurydice piperata Menzies and 


*Excirolana braziliensis Richardson, 

1912a 

*Excirolana mayana (Ives, 1891) 

Êxcorallana antillensis (Hansen, 

1890) 

Excorallana mexicana Richardson, 

1905a 

*Excorallana tricornis (Hansen, 

1890) 

*Harrieta faxoni (Richardson, 

1905a) 

*Limnoria tuberculata Sowinsky, 

1884 

Lironeca ovalis (Say, 1818) 

* Lironeca redmanni Leach, 1818 

Lironeca texana Pearse, 1952 

Lironeca tropicalis Menzies and 


*Nalicora rapax Moore, 1901 

*Nerocila acuminata Schioedte and 

Meinert, 1881 

Olencira praegustator (Latrobe, 1802) 

*Paracerceis caudata (Say, 1818) 

*Paradella dianae (Menzies, 1962b) 

Paradynamene benjamensis 

Richardson, 1905 

*Politolana polita (Stimpson, 1853) 

*Rocinela insularis Schioedte and 

Meinert, 1879 

*Rocinela oculata Harger, 1883 

*Rocinela signata Schioedte and 

Meinert, 1879 

*Serolis mgrayi Menzies and 


*Sphaeroma quadridentata Say, 1818 

*Sphaeroma terebrans Bate, 1866 

SUBORDER GNATHIIDEA 

Gnathia Jloridensis Menzies and 


SUBORDER MICROCERBERIDEA 

Microcerberus mexicanus Pennak, 1958

SUBORDER VALVIFERA 

Antarcturus jloridanus (Richardson, 

1900b) 

Arcturella bispinata Menzies and 


Arcturella spinata Menzies and 


Astacilla lauffi Menzies and 


Chiridotea excavata Harper, 

1974 

Cleantioides planicauda (Benedict, 

1899) 

species also occurring in the Caribbean 


Edotea lyonsi (Menzies and 

Kruczynski, 1983) 

Edotea montosa (Stimpson, 1853) 

Edwinjoycea horologium Menzies and 


*Erichsonella attenuata (Harger, 

1873) 

* Erichsonella filiformis (Say, 1818) 

* Erichsonella Jloridana Benedict, 

1901 

Erichsonella isabelensis Menzies, 

1951b 

*Idotea metallica Bosc, 1802 

Note: Records for the Gulf of Mexico have been assembled from published 

BERMUDA 

literature; in most cases, actual material has not been examined. 

Twenty-nine species of isopods have been recorded from Bermuda (Table 7). 

Of these, nine are endemics (three being cave forms). The remaining 20 

species have all been recorded from the Caribbean region, indicating a strong 

subtropical connection, in spite of the relatively high latitude (32°15'N). Al 

though Bermuda is of Eocene or Oligocene age, the tropical fauna was prob 

ably decimated by the low temperatures of the last Pleistocene glaciation 

(Briggs, 1974:76). 

GAVE ISOPODS 

With the expanding efforts of cave divers, more and more true stygobiont 

forms are being found. Concurrently, discussion of the origin of cave fauna 

has spurred several theories, all invoking the geological history of the Carib 

bean area. 

Among the isopods, cave forms have been found in four suborders, the 

Asellota, Anthuridea, Flabellifera, and Microcerberidea. Two valuable dis 

cussions on the origin of cave crustaceans may be found in Stock (1986) and 

Wagele (1985). 

The only true cave asellote, Atlantasellus cavernicolus Sket, was collected 

from Bermuda.


TABLE 7. ISOPOD SPECIES OCCURRING AT BERMUDA 

Alcirona krebsi Hansen, 1890 

*Anthomuda stenotelson Schultz, 1979 

*Apanthura harringtortiensis Wagele, 

1981 

*Arubolana aruboides (Bowman and 

Iliffc, 1983) 

*Atlantasellus cavernicolus Sket, 1979 

Bopyrissa wolffi Markham, 1978 

Cancricepon choprae (Nierstrasz and 


Carpias bermudensis Richardson, 1902 

*Carpias minutus (Richardson, 1902) 

*Colanthura tenuis Richardson, 1902 

Colopisthus parvus Richardson, 1902 

* Curassanthura bermudensis Wagele, 

1985 

Dynamenella perforata (Moore, 1901) 

Eurydice per sonata Kensley, 1987b 

* recorded only from Bermuda 

Excorallana quadricornis (Hansen, 1890) 

Joeropsis rathbunae Richardson, 1902 

Leidya bimini Pearse, 1951 

Paracerceis caudata (Say, 1818) 

Paranthura infundibulata Richardson, 

1902 

Pendanthura tanaiformis Menzies and 

Glynn, 1968 

Para theIges piriformis Markham, 1972b 

Parathelges tumidipes Markham, 1972b 

Probopyrinella latreuticola (Gissler, 

1882) 

Pseudione affinis (Sars, 1882) 

*Stegias clibanarii Richardson, 1904 

Stenetrium stebbingi Richardson, 1902 

Stenobermuda acutirostrata Schultz, 1979 

Synsynella choprae (Pearse, 1932) 

Synsynella deformans Hay, 1917 

The anthuridean cave representatives are found in two families: the genus 

Curassanthura Kensley in the Paranthuridae, and the genus Cyathura subgenus 

Stygocyathura Botosaneanu and Stock in the Anthuridae (see Figure 110). 

Three species of Curassanthura are known, one each from Curagao, Ber 

muda, and Lanzarote in the Canary Islands. Curassanthura halma Kensley, 

from Curagao, is an interstitial form found in hypersaline waters. Curas 

santhura bermudensis Wagele was found in water of about 26%o salinity. The 

Lanzarote species, C. canadensis Wagele, came from seawater in a lava cave. 

Wagele (1985) suggests that this amphi-Atlantic distribution of Curassanthura 

is the result of plate tectonics separating an ancestral hypogean progenitor 

that had a Tethyan distribution. 

The genus Cyathura has representatives in the sea, in estuarine-brackish 

habitats, and in freshwater caves, and is found in the Atlantic, Indian, and 

Pacific oceans. This widespread distribution suggests a very long history for 

the genus. Using the morphology of the male copulatory stylet, Wagele 

(1985) suggests that marine ancestors, having a Tethyan distribution, en 

tered freshwater interstitial habitats. The series of regressions of sea level 

forms. 

serv

Figure 110. Map showing distribution of cave anthurideans.

* cave cirolanids 

Figure 111. Map showing distrib


The flabelliferan family Cirolanidae contains five stygobiont genera in the 

Caribbean: Anopsilana, Arubolana, Bahalana, Creaseriella, and Haptolana (Figure 

111). Six other genera are known from the North American continent: 

Antrolana, Cirolanidesy Mexilana, Speocirolana, Sphaerolana, and Troglocirolana, all 

of which, except Antrolana from the Appalachian Valley of Virginia, occur in 

Mexico and Texas (see Notenboom, 1981). A few of these forms occur in 

brackish water, but most are found in freshwater of caves. Cave cirolanids 

are also known from Palau, North and East Africa, Madagascar, Bulgaria, 

Greece, Jugoslavia, Israel, France, and Spain. This widespread distribution 

again suggests a Tethyan marine origin, with dispersal and isolation due to 

sea regressions. 

The suborder Microcerberidea and the asellotan family Microparasellidae 

contain almost entirely interstitial forms, although few occur in caves. At 

least two genera, Microcerberus and Angliera, have very widespread distribu 

tions and are known from marine, brackish-water, and freshwater habitats, 

and may well have a history similar to that of Cyathura.

Appendix 

Since the manuscript of this work was completed and sent to press, a few 

papers have appeared either describing new species, mentioning new 

records for the Caribbean and associated areas, or instituting a major 

new taxon. It was thought useful to include these, if only in an appendix, 

to make the work as current as possible. The relevant taxa are listed 

alphabetically, with the full citation given below. 

Antheluridae Poore and Lew Ton, 1988 

Poore, G. C. B., and H. M. Lew Ton. 1988. Antheluridae, a new 

family of Crustacea (Isopoda: Anthuridea) with new species from 

Australia. Journal of Natural History 22:489—506. 

Within the geographical area covered by this work, only Anthomuda 

belongs to this new family. 

Aporobopyrus collardi Adkison, 1988 

Adkison, D. L. 1988. Pseudione parviramus and Aporobopyrus collardi, 

two new species of Bopyridae (Isopoda: Epicaridea) from the Gulf 

of Mexico. Proceedings of the Biological Society of Washington 

101(3):576-584. 

Booralana tricarinata Camp and Heard, 1988 

Camp, D. K., and R. W. Heard. 1988. Booralana tricarinata, a new 

species of isopod from the western Atlantic Ocean (Crustacea: Iso 

poda: Cirolanidae). Proceedings of the Biological Society of Washington 

101 (3):603-613. 

Originally recorded from the outer shelf and upper slope off the 

Little Bahama Bank and the Antilles Islands in 110-610 m, this 

species has since been recorded off Haiti in 620 m. 

Bythognathia yucatanensis Camp, 1988 

Camp, D. K. 1988. Bythognathia yucatanensis, new genus, new species, 

from abyssal depths in the Caribbean Sea, with a list of gnathiid 

275

276 APPENDIX 

species described since 1926 (Isopoda: Gnathiidae). Journal of Crust 

acean Biology 8(4):668-678. 

Edotea samariensis Muller, 1988 

Muller, H. G. 1988. Idoteidae aus N-Kolumbien mit Beschreibung 

von Edotea samariensis n. sp. (Crustacea: Isopoda: Valvifera). 

Senckenbergiana biologica 68(4/6):407-412. 

Gnathia Johanna Monod, 1926 

Miiller, H. G. 1988. Redescription of Gnathia Johanna, 1926 (Iso 

poda) from St. John, Virgin Islands. Bulletin Zoologisch Museum, Uni- 

versiteit van Amsterdam 11(15): 129—133. 

Phycolimnoria bacescui Ortiz and Lalana, 1988 

Ortiz, M., and R. Lalana. 1988. Una nueva especie del genero Phy 

colimnoria (Isopoda, Limnoriidae) de aguas cubanas. Revista de Inves 

tigations Marinas, La Habana 9(2): 37—42. 

Pseudione parviramus Adkison, 1988 

Adkison, D. L. 1988. Pseudione parviramus and Aporobopyrus collardi, 

two new species of Bopyridae (Isopoda: Epicaridea) from the Gulf 

of Mexico. Proceedings of the Biological Society of Washington 

101(3):576-584.

Literature Cited 

Adkison, D. L. 1982. Description of Dactylokepon sulcipes n. sp. (Crustacea: Isopoda: 

Bopyridae) and notes on D. caribaeus. Proceedings of the Biological Society of Washington 

95(4):702-708. 

. 1984a. Probopyrinella heardi n. sp. (Isopoda: Bopyridae) a branchial parasite of the 

hippolytid shrimp Latreutes parvulus (Decapoda: Caridea). Proceedings of the Biological Society of 

Washington 97(3):550-554. 

. 1984b. Two new species of Gigantione Kossmann (Isopoda: Epicaridea: Bopyridae) 

from the western North Atlantic. Proceedings of the Biological Society of Washington 97(4):761- 

772. 

Adkison, D. L., and Heard, R. W. 1978. Description of a new genus and species of 

Pseudioninae (Isopoda: Bopyridae) parasite of the hermit crab Pagurus annulipes (Stimpson) 

from North Carolina. Proceedings of the Biological Society of Washington 91 (2):408—417- 

Amar, R. 1957. Gnathostenetroides laodicense nov. gen. nov. sp. Type nouveau d'Asellota et 

classification des isopodes asellotes. Bulletin de I'lnstitut Oceanographique 1100:1 — 10. 

Argano, R. 1971. Cyathura sbordonii, nuova specie cavernicola del Messico sudorientale. 

Diagnosi preliminare (Crustacea, Isopoda, Anthuridae). Fragmenta Entomologica 7(4):303- 

304. 

Audouin, V. 1826. Explication Sommaire des Planches de Crustaces de PEgypte et de la 

Syrie. In J.-C. Savigny, Description de VEgypte ou Recueil des Observations et des Recherches qui ont 

etefaites en Egypte pendant {'Expedition de sa Majeste VEmpereur Napoleon le Grand. Histoire 

Naturelle, vol. 1, pp. 77-98. Paris: L'Imprimerie Imperiale. 

Barnard, K. H. 1914. Contributions to the crustacean fauna of South Africa. 3. Additions to 

the marine Isopoda, with notes on some previously incompletely known species. Annals of 

the South African Museum 10(1 l):325a-442. 

. 1920. Contributions to the crustacean fauna of South Africa. 6. Further additions to 

the list of marine Isopoda. Annals of the South African Museum 17(5):319—438. 

. 1925. A revision of the family Anthuridae (Crustacea Isopoda), with remarks on 

certain morphological peculiarities. Journal of the Linnaean Society of London, Zoology 36:109— 

160. 

Bate, C. S. 1866. Carcinological gleanings, 2. Annals and Magazine of Natural History (3)17:24- 

31. 

Bate, C. S., and J. O. Westwood. 1868. A History of the British Sessile-eyed Crustacea. Ivi + 536 

pp. London: John van Voorst. 

Becker, G., and W.-D. Kampf. 1958. Funde der holzzerstorenden Isopodengattung Limnoria 

an der Festland Indiens und Neubgeschreibung von Limnoria indica. Zeitschrifi fir Angewandte 

Zoologie 45:1—9. 

277

278 LITERATURE CITED 

Beddard, F. E. 1886. Report on the Isopoda collected by H. M. S. Challenger during the 

years 1873-76. Part 2. Reports of the Voyage of the Challenger 17:1-178. 

Benedict, J. E. 1899. [Cleantis planicauda Benedict, new species.] In Richardson, H. 1899. Key 

to the isopods of the Pacific coast of North America, with descriptions of twenty-two new 

species. Proceedings of the United States National Museum 21:815-869. 

. 1901. In Richardson, H. 1901. Key to the isopods of the Atlantic coast of North 

America with descriptions of new and little known species. Proceedings of the United States 

National Museum 23:493-579. 

Bliss, D. E., editor-in-chief. 1982-1985. The Biology of the Crustacea, vols. 1-10. New York: 

Academic Press. 

Bonnier, J. 1900. Contribution a l'etude des Epicarides. Les Bopyridae. Travaux de la Station 

Zoologique de Wimereux 8:1—396. 

Boone, P. L. 1918. Description often new isopods. Proceedings of the United States National 

Museum 54:591-604. 

. 1921. Report on the Tanidacea and Isopoda, collected by the Barbados-Antigua 

Expedition from the University of Iowa in 1918. University of Iowa Studies in Natural History 

9(5):91-98. 

Boone, L. 1927. Crustacea from tropical east American seas. Bulletin of the Bingham 

Oceanographic Collection 1(2): 1-147. 

. 1930. New decapod and isopod crustaceans from Gonave Bay, Haiti. Zoologica, New 

York 12(4):41-53. 

Bosc, L. A. G. 1802. Histoire Naturelle des Crustaces. In G. L. L. de BufTon, Histoire Naturelle 

de Buffon classee...d''apres le system de Linne...par R. R. Castel'...nouvelle edition. Paris. 

Botosaneanu, L. 1983. First record of an anthurid isopod, Cyathura univam sp. n., on the 

South American continent. Bijdragen tot de Dierkunde 53(2):247-254. 

Botosaneanu, L., N. L. Bruce, and J. Notenboom. 1986. Isopoda: Cirolanidae. In L. 

Botosaneanu, ed., Stygofauna Mundi, pp. 412—422. Leiden: E.J. Brill/Dr. W. Backhuys. 

Botosaneanu, L., and J. H. Stock. 1979. Arubolana imula n. gen., n. sp., the first hypogean 

cirolanid isopod crustacean found in the Lesser Antilles. Bijdragen tot de Dierkunde 

49(2):227-233. 

. 1982. Les Cyathura stygobies (Isopoda, Anthuridea) et surtout celles des Grandes et 

des Petites Antilles. Bijdragen tot de Dierkunde 52(1):13—42. 

Bourdon, R. 1972. Sur quelques Bopyridae (Crustacea, Isopoda) parasites de Galatheides. 

Bulletin du Museum national d'Histoire naturelle (3)66, zool. 52:817-838. 

. 1981. Remarques sur le genre Synsynella Hay, avec description de S* integra n. sp. 

(Crustacea, Epicaridea, Bopyridae). Bulletin du Museum national d'Histoire naturelle, Paris 

(4)3(A4):1143-1162. 

Bourdon, R., and J. C. Markham. 1980. A new genus and species of bopyrid isopod infesting 

alpheid shrimps of the genus Synalpheus in the western Atlantic Ocean. Zoologische 

Mededelingen 55( 19):221-230. 

Bovallius, C. 1885. New or imperfectly known Isopoda. Bihang till Kongliga Svenska Vetenskaps- 

Akademiens Handlingar 10(ll):l-32. 

Bowman, T. E. 1956. Una especie nueva de Bopyrella (Crustacea: Isopoda) de Los Roques,

LITERATURE CITED 279 

Venezuela. Novedades Cientijicas, Contribuciones Ocasionales del Museo de Historia Natural La 

Salle, Caracas, Venezuela (Zoologica) 19:1—4. 

. 1965. Cyathura specus, a new cave isopod from Cuba. Studies on the Fauna of Curacao and 

other Caribbean Islands 85:88-97. 

. 1966. Haptolana trichostomay a new genus and species of troglobitic cirolanid isopod 

from Cuba. International Journal of Speleology 2:105—108. 

. 1981. Thermosphaeroma milleri and T. smithi, new sphaeromatid isopod crustaceans 

from hot springs in Chihuahua, Mexico, with a review of the genus. Journal of Crustacean 

Biology 1(1):105-122. 

. 1987. Bahalana mayana, a new troglobitic cirolanid isopod from Cozumel Island and 

the Yucatan Peninsula, Mexico. Proceedings of the Biological Society of Washington 100(3):659- 

663. 

Bowman, T. E., and R. Franz. 1982. Anopsilana crenata, a new troglobitic cirolanid isopod 

from Grand Cayman Island, Caribbean Sea. Proceedings of the Biological Society of Washington 

95(3):522-529. 

Bowman, T. E., and T. M. Iliffe. 1983. Bermudalana aruboides, a new genus and species of 

troglobitic Isopoda (Cirolanidae) from marine caves on Bermuda. Proceedings of the 

Biological Society of Washington 96(2):291-300. 

Bowman, T. E., and B. F. Morris. 1979. Carpias Richardson, 1902, a senior synonym 

of Bagatus Nobili, 1906, and the validity of Carpias minutus (Richardson, 1902) 

(Isopoda: Asellota: Janiridae). Proceedings of the Biological Society of Washington 92(3):650- 

657. 

Boyle, P., and R. Mitchell. 1978. Absence of microorganisms in crustacean digestive tracts. 

Science 200(4346): 1157-1159. 

Brandt, J. F. 1833. Conspectus monographiae Crustaceorum Oniscodorum Latreillii. Bulletin 

de la Societe Imperiale des Naturalistes de Moscou 6:171 — 193. 

Briggs, J. C. 1974. Marine Zoogeography. 475 pp. New York: McGraw-Hill Book Company. 

Bruce, N. L. 1981. Cirolanidae (Crustacea: Isopoda) of Australia: Diagnoses of Cirolana 

Leach, Metacirolana Nierstrasz, Neocirolana Hale, Anopsilana Paulian & Deboutteville, and 

three new genera—Natatolana, Politolana, and Cartetolana. Australian Journal of Marine and 

Freshwater Research 32:945-966. 

. 1984. A new family for the isopod crustacean genus Tridentella Richardson, 1905, 

with description of a new species from Fiji. Zoological Journal of the Linnean Society 80:447- 

455. 

. 1985. Calyptolana hancocki, a new genus and species of marine isopod (Cirolanidae) 

from Aruba, Netherlands Antilles, with a synopsis of Cirolanidae known from the 

Caribbean and Gulf of Mexico. Journal of Crustacean Biology 5(4):707-716. 

. 1986a. Cirolanidae (Crustacea: Isopoda) of Australia. Records of the Australian 

Museum, supplement 6:1-239. 

. 1986b. Revision of the isopod crustacean genus Mothocya Costa, in Hope, 1851 

(Cymothoidae: Flabellifera), parasitic on marine fishes. Journal of Natural History 

20(5):1089-1192. 

Brusca, R. C. 1981. A monograph on the Isopoda Cymothoidae (Crustacea) of the eastern 

Pacific. Zoological Journal of the Linnean Society 73(2): 117—199.


. 1983. A monograph on the isopod family Aegidae in the tropical Eastern Pacific I. 

The genus Aega. Allan Hancock Monographs in Marine Biology 12:1-39. 

. 1984. Phylogeny, evolution and biogeography of the marine isopod Subfamily 

Idoteinae (Crustacea: Isopoda: Idoteidae). Transactions of the San Diego Society of Natural 

History 20(7):99-134. 

Budde-Lund, G. 1885. Crustacea Isopoda Terrestria per Familias et Genera et Species. 319 pp. 

Copenhagen. 

Buss, L. W., and E. VV. Iverson. 1981. A new genus and species of Sphaeromatidae 

(Crustacea: Isopoda) with experiments and observations on its reproductive biology, 

interspecific interactions and color polymorphisms. Postilla 184:1-23. 

Caiman, \V. T. 1904. On the classification of the Crustacea Malacostraca. Annals and 

Magazine of Natural History (7)13:144-158. 

. 1910. On two new species of wood-boring Crustacea from Christmas Island. Annals 

and Magazine of Natural History (8)5:181-186. 

Carpenter, J. H. 1981. Bahalana geracei n. gen., n. sp., a troglobitic marine cirolanid isopod 

from Lighthouse Cave, San Salvador Island, Bahamas. Bijdragen tot de Dierkunde 51 (2):259— 

267. 

Carpenter, J. H., and G.J. Magniez. 1982. Deux asellotes stygobies des Indes Occidentales: 

Neostenetroides stocki n. gen., n. sp., et Stenetrium sp. Bijdragen tot de Dierkunde 52(2):200-206. 

Carvacho, A. 1977. Isopodes de la Mangrove de la Guadeloupe, Antilles Franchises. Studies 

on the Fauna of Curasao and other Caribbean Islands 54( 174): 1—24. 

Chappuis, P.-A., and C. Delamare Deboutteville. 1955. Recherches sur les Crustaces 

souterrains. 7. Les isopodes psammiques de la Mediterranee. Archives de Zoologie 

Experimentale et Generale 91 (1): 103—138. 

. 1956. Etudes sur la faune interstitielle des iles Bahamas recoltee par Madame 

Renaud-Debyser. 1. Copepodes et Isopodes. Vie et Milieu 7(3):373-396. 

Clark, S. T., and P. B. Robertson. 1982. Shallow water marine isopods of Texas. Contributions 

to Marine Science 25(l):45-59. 

Coineau, N., and L. Botosaneanu. 1973. Isopodes interstitiels de Cuba. Resultats des expeditions 

bxospeologiques cubano-roumaines a Cuba 1:191—222. 

Cordiner, C. 1793. Remarkable ruins and romantic prospects, of North Britain. With ancient monuments 

and singular subjects of natural history. 96 plates with letterpress. London: Mazell. 

Costa, A. 1851. Caratteeri di alcuni de'generi e specie nouve segnete nel presente catalogo. In 

F. W. Hope, Catalogo dei crostacei Italiani e di molti altri de Mediterranean pp. 41-48. Naples. 

Coutiere, H. 1908. Sur le Synalpheion Giardi, n. gen., n. sp., entoniscien parasite d'une 

Synalphee. Comptes Rendus Hebdomadaires des Seances de VAcademie des Sciences, Paris 146:1333- 

1335. 

Creaser, E. P. 1936. Crustaceans from Yucatan. Carnegie Institution of Washington Publication 

457:117-132. 

Dana, J. D. 1852. On the classification of the Crustacea Choristopoda or Tetradecapoda. 

American Journal of Science (2)14:297-316. 

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): 1—20.


Delaney, P. M., and R. C. Brusca. 1985. Two new species of Tridentella Richardson, 1905 

(Isopoda: Flabellifera: Tridentellidae) from California, with a rediagnosis and comments 

on the family, and a key to the genera of Tridentellidae and Corallanidae. Journal of 

Crustacean Biology 5(4):728-742. 

Fabricius, J. C. 1793. Volume 2 in Entomologia systematica emendata et aucta...adjectis synonymis, 

locis, observationibus, descriptionibus, 4 vols., 1792-1794. Copenhagen. 

. 1798. Entomologia systematica emendata et aucta...adjectis synonymis, locis, obervationibus, 

descriptionibus. Supplementum. 1798. Copenhagen. 

Frankenberg, D., and R.J. Menzies. 1966. A new species ofasellote marine isopod, Munna 

(Uromunna) reynoldsi (Crustacea: Isopoda). Bulletin of Marine Science 16(2):200-208. 

Fresi, E., and U. Schieke. 1972. Pleurocope dasyura Walker, 1901, and the Pleurocopidae new 

family (Isopoda, Asellota). Crustaceana, Supplement 3:207-213. 

George, R. Y., and J.-O. Stromberg. 1968. Some new species and new records of marine 

isopods from San Juan Archipelago, Washington, U.S.A. Crustaceana 14(3):225-254. 

Gissler, C. F. 1882. Bopyroides latreuticola, a new species of isopod crustacean parasitic on a 

gulfweed shrimp. American Naturalist 16:591-594. 

Glynn, P. W. 1970. A systematic study of the Sphaeromatidae (Crustacea: Isopoda) of Isla 

Margarita, Venezuela, with descriptions of three new species. Memoria de la Sociedad de 

Ciencias Naturales La Salle 30:1-48. 

Glynn P. W., D. M. Dexter, and T. E. Bowman. 1975. Excirolana braziliensis, a Pan-American 

sand beach isopod: taxonomic status, zonation and distribution. Journal of Zoology, London 

175:509-521. 

Glynn, P. W., and C. S. Glynn. 1974. On the Systematics on Ancinus (Isopoda, 

Sphaeromatidae), with the description of a new species from the tropical eastern Pacific. 

Pacific Science 28(4) :401-422. 

Grobben, C. 1892. Zur Kenntniss der Staumbaumer und der Systems der Crustaceen. 

Sitzungsberichte der kaiserlichen Akademie der Wissenschaften. Mathematisch-Naturwissenschaftliche 

Classe, IVien 101 (1 &2) 1:237-274. 

Haller, G. 1880. Ueber einige neue Cymothoinen. Archiv fur Naturgeschichte, Berlin 46(1)375- 

395. 

Hansen, H.J. 1890. Cirolanidae et familiae nonnullae propinquae Musei Hauniensis. Et 

Bidrag til Kundskaben om nogle Familier af isopode Krebsdyr. Kongelige Danske 

Videnskabernes Selskabs Shifter, 6te Raekke, Naturvidenskabelig og mathematisk Afdeling 3:239-426. 

. 1904. On the morphology and classification of the Asellota-group of crustaceans, 

with descriptions of the genus Stenetrium Hasw. and its species. Proceedings of the Zoological 

Society of London, 1904, 2:302-331. 

. 1905a. On the morphology and classification of the Asellota-group of Crustacea, 

with descriptions of the genus Stenetrium Haswell, and its species. Proceedings of the Zoological 

Society of London for 1904 2(2):302-331. 

. 1905b. On the propagation, structure, and classification of the family Sphaeromidae. 

Quarterly Journal of Microscopical Science 49(1):69—135. 

. 1916. Crustacea Malacostraca 3. Danish Ingo If Expedition 3(5): 1—262. 

Harger, O. 1873. [Erichsonia attenuata.] In A. E. Verrill, S. I. Smith, and O. Harger,


Catalogue of the Marine invertebrate animals of the southern coast of New England, and 

adjacent waters, p. 276. In A. E. Verrill and S. I. Smith, Report upon the invertebrate 

animals of Vineyard Sound and adjacent waters, with an account of the physical features 

of the region. In S. F. Baird, Report of Professor S. F. Baird, Commissioner of Fish and Fisheries, 

on the condition of the sea-fisheries of the south coast of New England in 1871 and 1872. 478 pp. 

Washington, D.C.: Government Printing Office. 

. 1879. Notes on New England Isopoda. Proceedings of the United States National Museum 

No.75:157-165. 

. 1883. Reports on the results of dredging, under the supervision of Alexander 

Agassiz, on the east coast of the United States, during the summer of 1880, by the U.S. 

Coast Survey Steamer "Blake," Commander J. R. Bartlett, U.S.N., commanding. Bulletin 

of the Museum of Comparative Zoology at Harvard College 11 (4):91 —104. 

Harper, D. E., Jr. 1974. Chiridotea excavata n. sp. (Crustacea, Isopoda) from marine waters of 

Texas. Contributions to Marine Sciences, Texas A & M University 18:229-239. 

Harrison, K. 1984. The morphology of the sphaeromatid brood pouch (Crustacea: Isopoda: 

Sphaeromatidae). Zoological Journal of the Linnean Society 82:363-407. 

Harrison, K., and D. M. Holdich. 1982. New eubranchiate sphaeromatid isopods from 

Queensland waters. Memoirs of the Queensland Museum 20(3):421-446. 

Hartnoll, R. G. 1966. A new entoniscid from Jamaica (Isopoda, Epicaridea). Crustaceana 

ll(l):45-52. 

Haswell, W. A. 1881. On some new Australian marine Isopoda. Part 1. Proceedings of the 

Linnean Society of New South Wales 5:476-478. 

. 1884. On a new crustacean found inhabiting the tubes of Vermilia (Serpulidae). 

Proceedings of the Linnean Society of New South Wales 9(3):676-680. 

Hay, W. P. 1903. On a small collection of crustaceans from the island of Cuba. Proceedings of 

the United States National Museum 26:429-435. 

. 1917. A new genus and three new species of parasitic isopod crustaceans. Proceedings 

of the United States National Museum 51:569-574. 

Hooker, A. 1985. New species of Isopoda from the Florida Middlegrounds (Crustacea: 

Peracarida). Proceedings of the Biological Society of Washington 98(l):255-280. 

Hurley, D. E., and K. P. Jansen. 1977. The marine fauna of New Zealand: Family 

Sphaeromatidae (Crustacea Isopoda: Flabellifera). New Zealand Oceanographic Institute 

Memoir 63:1095. 

Iverson, E. W. 1982. Revision of the isopod family Sphaeromatidae (Crustacea: Isopoda: 

Flabellifera) I. Subfamily names with diagnoses and key. Journal of Crustacean Biology 

2(2):248-254. 

Ives, J. E. 1891. Crustacea from the northern coast of Yucatan, the harbor of Vera Cruz, the 

west coast of Florida and the Bermuda Islands. Proceedings of the Academy of Natural Sciences 

of Philadelphia, 1891:176-207. 

Jacobs, B.J. M. 1987. A taxonomic revision of the European, Mediterranean and NW. 

African species generally placed in Sphaeroma Bosc, 1802 (Isopoda: Flabellifera: 

Sphaeromatidae). Zoologische Verhandelingen 238:1-71. 

Kaestner, A. 1967. Invertebrate Zoology, vol. 3, 523 pp. New York: Interscience Publisher. 

[Translated and adapted from the second German edition by H. W. Levi and 

L. R. Levi.]


Karaman, S. 1933a. Neue Isopoden aus unterirdischen Gewassern Jugoslawiens. Zoologischer 

Anzeiger 102( 1/2): 16-22. 

. 1933b. Microcerberus stygius, der dritte Isopod aus dem Grundwasser von Skopje, 

Jugoslawien. Zoologischer Anzeiger 102(5/6): 165—169. 

. 1934. Beitrage zur Kenntnis des Isopoden-Familie Microparasellidae. Mitteilungen 

uber Hohlen- und Karstforschung 1934:42-44. 

Kensley, B. 1978. Five new genera of anthurid isopod crustaceans. Proceedings of the Biological 

Society of Washington 91 (3):775-792. 

. 1980. Records of anthurids from Florida, Central America, and South America 

(Crustacea: Isopoda: Anthuridae). Proceedings of the Biological Society of Washington 93(3):725- 

742. 

. 1981. Amsterdam Expeditions to the West Indian Islands, Report 10. Curassanthura 

halma, a new genus and species of interstitial isopod from Curasao, West Indies 

(Crustacea: Isopoda: Paranthuridae). Bijdragen tot de Dierkunde 51(1): 131-134. 

. 1982. Anthuridea (Crustacea: Isopoda) of Carrie Bow Cay, Belize. In K. Rutzler 

and I. G. Macintyre, eds., The Atlantic Barrier Reef Ecosystem at Carrie Bow Cay, 

Belize, 1: Structure and Communities, pp. 321-352. Smithsonian Contributions to Marine 

Sciences 12, 539 pp. 

. 1983. The role of isopod crustaceans in the reef crest community at Carrie Bow Cay, 

Belize. Marine Ecology 5(l):29-44. 

. 1984. The Atlantic Barrier Reef Ecosystem at Carrie Bow Cay, Belize, III: New 

marine Isopoda. Smithsonian Contributions to Marine Sciences 24, iv + 81 pp. 

. 1987a. Further records of marine isopods from the Caribbean. Proceedings of the 

Biological Society of Washington 100(3):559-577. 

. 1987b. A re-evaluation of the Systematics of K. H. Barnard's Review of anthuridean 

isopods. Steenstrupia 13(3):101 —139- 

. 1987c. Harrieta, a new genus for Cymodoce faxoni (Richardson) (Crustacea: Isopoda: 

Sphaeromatidae). Proceedings of the Biological Society of Washington 100(4): 

Kensley, B., and R. Heard. 1985. A new species of the genus Spinianirella Menzies 

(Crustacea: Isopoda: Janiridae) from the western Atlantic. Proceedings of the Biological Society 

of Washington 98(3):682-686. 

Kensley, B., and H. W. Kaufman. 1978. Cleantioides, a new idoteid isopod genus from Baja 

California and Panama. Proceedings of the Biological Society of Washington 91(3):658-665. 

Kensley, B., and M. Schotte. 1987. New records of isopod Crustacea from the Caribbean, 

the Florida Keys, and the Bahamas. Proceedings of the Biological Society of Washington 

100(l):216-247. 

Kensley, B., and P. Snelgrove. 1987. Records of marine isopod crustaceans associated with 

the coral Madracis mirabilis from Barbados. Proceedings of the Biological Society of Washington 

100(1):186-197. 

Koehler, R. 1885. Description d'un Isopode nouveau, le Joeropsis brevicomis. Annates des Sciences 

Naturelles (Paris), Zoologie (6)19:1-7. 

Krayer, H. 1839. Munna, en ny kraebsdyrslaegt. Naturhistorisk Tidsskrifl, Kjebenhavn 2:612— 

616. 

Kussakin, O. G. 1967. Fauna of Isopoda and Tanaidacea in the coastal zones of the


Antarctic and Subantarctic waters. Biological Reports of the Soviet Antarctic Expedition (1955- 

1958) 3:220-389. [English translation by the Israel Program for Scientific Translations, 

Jerusalem, 1968.] 

Lang, K. 1961. Contributions to the knowledge of the genus Microcerberus Karaman 

(Crustacea Isopoda) with a description of a new species from the central Californian coast. 

Arkivfdr Zoologi (2)13(22):493-510. 

Latreille, P. A. 1802. Histoire Naturelle des Crustaces et des Insectes. In Volume 3 of G. L. 

L. de BufTon, 1802-1805, Histoire Naturelle, nouvelle edition, accompagnee des notes. Ouvrage redige 

par C. S. Sonnini, 14 vols. Paris. 

. 1803. Histoire Naturelle des Crustaces et des Insectes. In Volume 5 of G. L. L. de 

Buffon, 1802-1805. Histoire Naturelle, nouvelle edition, accompagnee des notes. Ouvrage redige par 

C. S. Sonnini. 14 vols. Paris 

. 1806. Genera Crustaceorum et Insectorum secundum ordinum naturalem in Familias disposita, 

iconibus exemplisque plurimis explicata, vol. 1, 280 pp. Paris: Amand Koenig. 

. 1817. Les Crustaces, les Arachnides, et les Insectes. In G. L. C. F. D. Cuvier, Le 

Regne Animal, distribue d'apres son organisation, pour servrir de base a I'histoire naturelle des animaux 

et d y introduction a Vanatomie comparee, vol. 3. Paris. 

. 1826. Explication sommaire des planches (Mollusques, Annelides, Crustace's, Arachnides, 

Insectes, Echinoderms, Zoophytes, Ascidies, Polypes, Hydrophytes, Oiseaux) dont les dessins ont ete 

fournis par M. J. C. Savigny. Description de VEgypte, ou recueil des observations et des recherches qui 

ont etefaites en Egypte pendant êxpedition de Varmee Francaise (1798-1801). Paris. 

. 1831. Cours d'Entomologique, ou de Vhistoire naturelle des Crustaces, des Arachnides, des 

Alyriapodes, et des Insectes. 568 pp. Paris. 

Latrobe, B. H. 1802. A drawing and description of the Clupea tyrannus and Oniscus praegustator. 

Transactions of the American Philosophical Society 5:77—81. 

Leach, W. E. 1814. Crustaceology. In Brewster's Edinburgh Encyclopedia, vol. 7, pp. 383-439. 

. 1815. A tabular view of the external characters of four classes of animals, which 

Linne arranged under Insecta; with the description of the genera comprising three of these 

classes into order, etc., and descriptions of several new genera and species. Transactions of 

the Linnean Society of London 2:306—400. 

. 1818. [Rocinela, Livoneca redmanni.J In Volume 5 of F. Cuvier, ed., 1816—1830, 

Dictionnaire des Sciences naturelles. Paris & Strasbourg. 

Leidy, J. 1855. Contributions towards a knowledge of the marine invertebrate fauna of the 

coasts of Rhode Island and New Jersey. Journal of the Academy of Natural Sciences of 

Philadelphia 3:135-152. 

Lemos de Castro, A. 1959. Descrigao de uma nova especie do genero "Ancinus" Milne 

Edwards (Isopoda, Sphaeromidae). Revista Brasileira de Biologia 19(2):215—218. 

Lilljeborg, W. 1851. Norger Crustacear. Ofoersigt af Kongliga Vetenskapsakademiens Forhandlingar, 

Stockholm 8:19-25. 

Linnaeus, C. 1793. In J. C. Fabricius, 1792-1794. Entomologia systematica emendata et 

aucta...adjectis synonymis, locis, observationibus, descriptionibus, 4 vols. Copenhagen. 

Loyola e Silva, J. de. 1960. Sphaeromatidae do Litoral Brasileiro (Isopoda—Crustacea). 

Boletim da Universidade do Parana, Zoologia 4:1 — 182. 

Markham, J. C. 1972a. Two new genera of western Atlantic abdominally parasitizing


Bopyridae (Isopoda, Epicaridea), with a proposed new name for their subfamily. 

Crustaceana, Supplement 3:39-56. 

. 1972b. Four new species of Parathelges Bonnier, 1900 (Isopoda, Bopyridae), the first 

record of the genus from the western Atlantic. Crustaceana, Supplement 3:57-78. 

. 1973. Six new species of bopyrid isopods parasitic on galatheid crabs of the genus 

Munida in the Western Atlantic. Bulletin of Marine Science 23(3) :613-648. 

. 1974. A new species of Pleurocrypta (Isopoda, Bopyridae), the first known from the 

western Atlantic. Crustaceana 26(3):267-272. 

. 1975a. A review of the bopyrid isopod genus Munidion Hansen, 1897, parasitic on 

galatheid crabs in the Atlantic and Pacific oceans. Bulletin of Marine Science 25(3):422-441. 

. 1975b. Bopyrid isopods infesting porcellanid crabs in the northwestern Atlantic. 

Crustaceana 28(3):257-270. 

. 1975c. New records of two species of parasitic isopods of the bopyrid subfamily 

Ioninae in the western Atlantic. Crustaceana 29(l):55-67. 

. 1975d. Two new species of Asymmetrione (Isopoda, Bopyridae) from the Western 

Atlantic. Crustaceana 29(3):255-265. 

. 1977. Description of new western Atlantic species ofArgeia Dana with a proposed 

new subfamily for this and related genera (Crustacea Isopoda, Bopyridae). Zoologische 

Mededelingen 52(9): 107-123. 

. 1978. Bopyrid isopods parasitizing hermit crabs in the northwestern Atlantic Ocean. 

Bulletin of Marine Science 28(1): 102-117. 

. 1985. A review of the bopyrid isopods infesting caridean shrimps in the 

northwestern Atlantic Ocean, with special reference to those collected during the 

Hourglass cruises in the Gulf of Mexico, Memoirs of the Hourglass Cruises 7(3): 1 ~ 156. 

Menzies, R.J. 1951a. A new species of Limnoria (Crustacea: Isopoda) from Southern 

California. Bulletin of the Southern California Academy of Sciences 50(2) :86—88. 

. 1951b. A new subspecies of marine isopod from Texas. Proceedings of the United States 


. 1956a. New abyssal tropical Atlantic isopods, with observations on their biology. 

American Museum Novitates 1798:1 — 16. 

. 1956b. New bathyal Isopoda from the Caribbean with observations on their 

nutrition. Breviora 63:1-10. 

. 1957. The marine borer family Limnoriidae (Crustacea, Isopoda). Bulletin of Marine 

Science of the Gulf and Caribbean 7(2):101-200. 

1:79-206. 

. 1962a. The isopods of abyssal depths in the Atlantic Ocean. Vema Research Series 

. 1962b. The marine isopod fauna of Bahia de San Quintin, Baja California, Mexico. 

Pacific Naturalist 3(11):337-348. 

. 1962c. The zoogeography, ecology, and Systematics of the Chilean marine isopods. 

Lunds Universitets Arsskrift, N.F. Avd. 2, 57(11):1 —162. 

Menzies, R. J., and D. Frankenberg. 1966. Handbook on the Common Marine Isopod Crustacea of 

Georgia. University of Georgia Press, Athens, Georgia. 93 pp. 

Menzies, R. J., and P. W. Glynn. 1968. The common marine isopod Crustacea of Puerto


Rico: A handbook for marine biologists. Studies on the Fauna of Curasao and other Caribbean 

Islands 27(104):1-133. 

Menzies, R. J., and W. L. Kruczynski. 1983. Isopod Crustacea (Exclusive of Epicaridea). 

Memoirs of the Hourglass Cruises 6(1): 1 — 126. 

Miers, E.J. 1880. On a collection of Crustacea from the Malaysian region. Part 4. 

Penaeidea, Stomatopoda, Isopoda, Suctoria, and Xiphosura. Annals and Magazine of Natural 

History (5)5:457. 

Miller, M. A. 1941. The isopod Crustacea of the Hawaiian Islands, II. Asellota. Occasional 

Papers of the Bernice P. Bishop Museum, Honolulu, Hawaii 16( 13):305—320. 

Milne Edwards, A. 1879. Sur un isopode gigantesque, des grandes profondeurs de la mer. 

Comptes Rendus Hebdomadaires des Seances de VAcademie des Sciences 88:21-23. 

Milne Edwards, H. 1840. Histoire Naturelle des Crustaces, comprenant ranatomie, la physiologie et la 

classification de ces animaux, vol. 3. Paris. 

Monod, T. 1926. Les Gnathiidae. Essai monographique (morphologie, biologie, 

systematique). Memoires de la Societe des Sciences Naturelles du Maroc 13:1—667. 

Montagu, G. 1804. Description of several marine animals (Cancer rhomboidalis, C. 

maxillaris, C. phasma, C. palmatus, Oniscus hirsutus, etc) found on the south coast of 

Devonshire. Transactions of the Linnean Society, London 7:61—85. 

Moore, H. F. 1901. Report on Porto Rican Isopoda. U.S. Fish Commission Bulletin for 1900 

2:161-176. 

Moreira, P. S. 1972. Species of Eurydice (Isopoda, Flabellifera) from southern Brazil. Boletim 

do Instituto Paulista de Oceanografico, Sao Paula 21:69-91. 

Miiller, H.-G. 1988. The genus Gnathia Leach (Isopoda) from the Santa-Marta area, 

northern Colombia, with a review of Gnathiidae from the Caribbean Sea and Gulf of 

Mexico. Bijdragen tot de Dierkunde 58(1 ):88-104. 

Negoescu, I. 1979. Cyathura cubana sp. n. (Isopoda, Anthuridea) from the Caribbean Sea 

(Cuban waters). Travaux du Museum d'Histoire naturelle Grigore Antipa 20:157-164. 

Negoescu Vladescu, I. 1983. A study of genus Cyathura from the Cuban freshwaters with the 

description of a new cave species: C. orghidani (Isopoda, Anthuridae). Resultats des expeditions 

biospeologiques cubano-roumaines a Cuba 4:39—45. 

Nierstrasz, H. F. 1931. Die Isopoden der Siboga-Expedition. III. Isopoda Genuina II. 

Flabellifera. Siboga Expedition Monographie 32c: 123-232. 

Nierstrasz, H. F., and G. A. Brender a Brandis, 1925. Bijdrage tot de kennis der fauna van 

Curasao. Epicaridea. Bijdragen tot de Dierkunde 24:1—8. 

. 1929. Papers from Dr. Th. Mortensen's Pacific Expedition 1914—16. 48. Epicaridea 

1. Videnskabelige Meddelelsers fra Dansk Naturhistorisk Forening i Kjebenhavn 87:1-44. 

. 1931. Papers from Dr. Th. Mortensen's Pacific Expedition 1914-16. 57. Epicaridea 

2. Videnskabelige Meddelelsers fra Dansk Naturhistorisk Forening i Kjebenhavn 91:147-225. 

Nordenstam, A. 1933. Marine Isopoda of the families Serolidae, Idotheidae, 

Pseudidotheidae, Arcturidae, Parasellidae and Stenetriidae mainly from the South 

Atlantic. Further Zoological Results of the Swedish Antarctic Expedition 1901-1903 3(1): 1—284. 

Norman, A. M., and T. R. R. Stebbing. 1886. Crustacea Isopoda of the 'Lightning', 

'Porcupine', and 'Valorous' Expeditions, Part 1. Transactions of the Zoological Society of London 

12(4):119-133.


Notenboom, J. 1981. Amsterdam Expeditions to the West Indies Islands, report 12. Some 

new hypogean cirolanid isopod crustaceans from Haiti and Mayaguana (Bahamas). 

Bijdragen tot de Dierkunde 51(2):313—331. 

. 1984. Arubolana parvioculata n. sp. (Isopoda, Cirolanidae) from the interstitial of an 

intermittent river in Jamaica, with notes on A. imula Botosaneanu & Stock and A. aruboides 

(Bowman & IlifTe). Bijdragen tot de Dierkunde 54(1 ):51—65. 

Nunomura, N. 1977. Marine Isopoda from Amakusa, Kyushu (I). Publications from the 

Amakusa Marine Biological Laboratory 4:71-90. 

Ortiz, M., and R. Lalana. 1980. Una nueva especie de isopodo (Crustacea, Isopoda), de los 

manglares de la costa sur de Cuba. Revista Investigaciones Marinas 1(2-3): 160-174. 

Ortiz, M., R. Lalana, and O. Gomez. 1987. Lista de especies y bibliografia de los isopodos 

(Crustacea, Peracarida) de Cuba. Revista de Investigaciones Marinas 8(3):29-37. 

Packard, A. S. 1879. Zoology for Students and General Readers. 719 pp. New York: Henry Holt & Co. 

Pallas, P. S. 1772. In fasc. 9, Spicilegia Zoologica (quibus novae...et obscurae animalium 

species...illustrantur). 2 vols. (1767-) 1774-1780. Berlin. 

Paul, A. Z., and R.J. Menzies. 1971. Sub-tidal isopods of the Fosa de Cariaco, Venezuela, 

with descriptions of two new genera and twelve new species. Boletin de Instituto Universidade 

Oriente 10(l):29-48. 

Paulian, R., and C. Delamare Deboutteville. 1956. Un cirolanide cavernicole a Madagascar 

(Isopode). Memoires de VInstitut Scientifique de Madagascar (A)9:85-88. 

Pearse, A. S. 1932. New bopyrid isopod crustaceans from Dry Tortugas, Florida. Proceedings 

of the United States National Museum 81(1): 1-6. 

. 1950. Bopyrid isopods from the coast of North Carolina. Journal of the Elisha Mitchell 

Scientific Society 66:41-43. 

. 1951. Parasitic Crustacea from Bimini, Bahamas. Proceedings of the United States 

National Museum 101(3280):341-372. 

. 1952. Parasitic Crustacea from the Texas coast. Publications of the Institute of Marine 

Sciences, University of Texas 2(2):5-42. 

Pearse, A. S., and H. A. Walker. 1939. Two new parasitic isopods from the eastern coast of 

North America. Proceedings of the United States National Museum 87:19-23. 

Pennak, R. W. 1958. A new micro-isopod from a Mexican marine beach. Transactions of the 

American Microscopical Society 77:298-303. 

Pennant, T. 1777. British Zoology. 4th Edition, vol. 4. 

Perty, J. A. 1833. In Delectus animalium articulatorum quae in itinere per Brasiliam annis 1817- 

20„.collegerunt..J. B. de Spix...et...C. F. P. de Martius, digessit, descripsit, pingenda curavit M. 

Perty, vol. 3. Monaco. 

Petuch, E.J. 1982. Geographical heterochrony: contemporaneous coexistence of Neogene and 

Recent molluscan faunas in the Americas. Palaeogeography, Palaeoclimatology, Palaeoecology 

37:277-312. 

Pires, A. M. S. 1980. Revalidation and redescription of the genus Carpias Richardson, 1902 

(Isopoda, Asellota). Crustaceana 39(1):95-103. 

. 1981. Carpias harrietae (Isopoda, Asellota), a new species from Florida. Crustaceana 

40(2):206-212.


. 1982. Taxonomic revision of Bagatus (Isopoda, Asellota) with a discussion of 

ontogenetic polymorphism in males. Journal of Natural History 16(2):227—259. 

. 1984. Taxonomic revision and phylogeny of the genus Erichsonella with a discussion 

on Ronalea (Isopoda, Valvifera). Journal of Natural History 18(5):665-683. 

Poore, G. C. B. 1984. Redefinition of Munna and Uromunna (Crustacea: Isopoda: Munnidae), 

with descriptions of five species from coastal Victoria. Proceedings of the Royal Society of 

Victoria 96(2):61-81. 

Poore, G. C. B., and Lew Ton, H. M. 1988. Amakusanthura and Apanthura (Crustacea: 

Isopoda: Anthuridae) with new species from tropical Australia. Memoirs of the Museum of 

Victoria 49:107-147. 

Racovitza, E. G. 1908. Ischyromene Lacazei n.g., n. sp. Isopode mediterraneen de la famille des 

Spheromides (Note preliminaire). Archives de Zoologie Experimental et Generale (4)9, notes et 

revue 3:LX-LXIV. 

Rafinesque-Schmaltz, C. S. 1815. Analyse de la Nature ou Tableau de UUnivers et des Corps 

Organises. 224 pp. Palerma. 

Ray, D. L. 1959 (Editor). Marine boring and fouling organisms. 536 pp. Seattle: University of 

Washington Press. 

Rehm, A., and H.J. Humm. 1973. Sphaeroma terebrans: A threat to the mangroves of 

southwestern Florida. Science 182:173-174. 

Richardson, H. 1897. Description of a new species of Sphaeroma. Proceedings of the Biological 

Society of Washington 11:105-107. 

. 1898. Description of four new species of Rocinela with a synopsis of the genus. 

Proceedings of the American Philosophical Society 37(157):8— 17. 

. 1899. Key to the isopods of the Pacific coast of North America, with descriptions of 

twenty-two new species. Proceedings of the United States National Museum 21:815-869. 

. 1900a. Results of the Branner-Agassiz Expedition to Brazil 2. The isopod Crustacea. 

Proceedings of the Washington Academy of Sciences 2:157-159. 

. 1900b. Synopses of North American invertebrates. 7. The Isopoda. American 

Naturalist 34:207-230. 

. 1901. Key to the isopods of the Atlantic coast of North America with descriptions of 

new and little known species. Proceedings of the United States National Museum 23:493-579. 

. 1902. The marine and terrestrial isopods of the Bermudas, with descriptions of new 

genera and species. Transactions of the Connecticut Academy of Sciences 11:277-310. 

. 1904. Contributions to the natural history of the Isopoda. Proceedings of the United 

States National Museum 27:1-89. 

. 1905. A monograph on the isopods of North America. Bulletin of the United States 

National Museum 54, liii + 727 pp. 

. 1910. Description of a new species of Anilocra from the Atlantic coast of North 

America. Proceedings of the United States National Museum 39:137-138. 

. 1912a. Descriptions of a new genus of isopod crustaceans, and of two new species 

from South America. Proceedings of the United States National Museum 43:201-204. 

. 1912b. Description of a new isopod crustacean belonging to the genus Livoneca from 

the Atlantic coast of Panama. Proceedings of the United States National Museum 42:173-174.


. 1912c. Marine and terrestrial isopods from Jamaica. Proceedings of the United States 


Rioja, E. 1953. Estudios carcinologicos. XXX. Observaciones sobre los cirolanidos 

cavernicolas de Mexico (Crustaceos, Isopodos). Anales del Instituto de Biologia, Mexico 

24(1):147-170. 

Robertson, P. B. 1978. A new species of asellote marine isopod Munna (Uromunna) hayesi 

(Crustacea: Isopoda) from Texas. Contributions in Marine Science 21(1 ):39—46. 

Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker, E. A. Lachner, R. N. Lea, and W. 

B. Scott. 1980. A list of common and scientific names of fishes from the United States and 

Canada (4th edition). American Fisheries Society, Special Publication 12:1-174. 

Roman, M.-L. 1977. Les oniscoides halophiles de Madagascar (Isopoda, Oniscoidea). 

Beaufortia 26:107-152. 

Roux, P. 1828. Crustaces de la Mediterranee et de son littoral, decrits et lithographies par 

iui-meme. Annates des Sciences Naturelles 16: plate 13. 

Sars, G. O. 1882. Oversigt af Norges Crustacea. Forhandlinger i Videnskabsselskabet i Kristiania 

1882 18:1-124. 

. 1897. An account of the Crustacea of Norway, vol. 2, pts. 3-8. Isopoda. 103 pp. Bergen. 

. 1899. An account of the Crustacea of Norway, vol. 2, pts. 13-14. Isopoda. 270 pp. Bergen. 

Say, T. 1818. An account of the Crustacea of the United States. Journal of the Academy of 

Natural Sciences of Philadelphia 1(2):393-401, 423-433. 

Schioedte, J. C, and F. Meinert. 1879. Symbolae ad Monographiam Cymothoarum 

Crustaceorum Isopodum Familiae 1. Aegidae. Naturhistorisk Tidsskrifl (3)12:321-414. 

. 1881. Symbolae ad Monographiam Cymothoarum Crustaceorum Isopodum 

Familiae 2. Anilocridae. Naturhistorisk Tidsskrifl (3)13:1 — 166. 


Familiae 3. Saophridae. 4. Ceratothoinae. Naturhistorisk Tidsskrifl (3)13:281—378. 


Familiae 4. Cymothoidae. Trib. II. Cymothoinae. Trib. III. Livonecinae. Naturhistorisk 

Tidsskrifl (3)14:221-454. 

Schram, F. R. 1986. Crustacea, xiv + 606 pp. New York, Oxford: Oxford University Press. 

Schultz, G. A. 1969. How to know the marine isopod crustaceans, vii + 359 pp. Dubuque, Iowa: 

Wm. C. Brown Co. 

. 1974. Terrestrial isopod crustaceans (Oniscoidea) mainly from the West Indies and 

adjacent regions. I. Tylos and Ligia. Studies on the Fauna of Curaqao and other Caribbean Islands 

45(149):162-173. 

. 1977. Anthurids from the west coast of North America, including a new species and 

three new genera (Crustacea, Isopoda). Proceedings of the Biological Society of Washington 

90(4):839-848. 

. 1979. A new Asellota (Stenetriidae) and two, one new, Anthuridea (Anthuridae) 

from Bermuda (Crustacea, Isopoda). Proceedings of the Biological Society of Washington 

91(4):904-911. 

. 1984. Three new and five other species of Oniscoidea from Belize, Central America 

(Crustacea: Isopoda). Journal of Natural History 18:3-14.


Schultz, G. A., and D. M. Allen. 1982. Prodajus bigelowiensis, new species (Isopoda: 

Epicaridea: Dajidae) parasite of Mysidopsis bigelowi (Mysidacea) from coastal New Jersey, 

with observations on infestation. Journal of Crustacean Biology 2(2):296-302. 

Schultz, G. A., and L. R. McCloskey. 1967. Isopod crustaceans from the coral Oculina 

arbuscula Verrill. Journal of the Elisha Mitchell Scientific Society 83(2): 103—113. 

SimberlolT, D., B.J. Brown, and S. Lowrie. 1978. Isopod and insect root borers may benefit 

Florida mangroves. Science 201:630-632. 

Sivertsen, E., and L. B. Holthuis. 1980. The marine isopod Crustacea of the Tristan da 

Cunha Archipelago. Gunneria 35:1-128. 

Sket, B. 1979. Atlantasellus cavemicolus n. gen., n. sp. (Isopoda Asellota, Atlantasellidae n. 

fam.) from Bermuda. Bioloski Vestnik, Ljubljana 7(2): 175— 183. 

Soika, A. G. 1954. Studi di Ecologia e Biogeografia 12. Ecologia, Sistematica, Biogeografia 

ed Evoluzione del Tylos latreillei Auct. (Isop. Tylidae). Bollettino del Museo Civico di Storia 

Naturale di Venecia 7:63-83. 

Sowinsky, V. K. 1884. Contribution to the crustacean fauna of the Black Sea. Memoires de la 

Societe des Naturalistes de Kieff7:225-288. [In Russian.] 

Stebbing, T. R. R. 1900. On Crustacea brought by Dr. Willey from the South Seas. Willey's 

Zoological Results 5:605-690. 

. 1904. Marine Crustaceans. 12. Isopoda, with description of a new genus. In J. S. 

Gardiner, The Fauna and Geography of the Maldive and Laccadive Archipelagoes, being the account of 

the work carried on and of the collections made by an expedition during the years 1899 and 1900, vol. 2, 

pp. 699-720. Cambridge: Cambridge University Press. 

. 1905. Report to the Government of Ceylon on the pearl oyster fisheries of the Gulf 

of Manaar. Report on the Isopoda collected by Professor Herdman, at Ceylon, in 1902. 

Ceylon Pearl Oyster Fisheries, 1905, supplementary reports 23:1-64. 

Stimpson, W. 1853. Synopsis of the marine Invertebrata of Grand Manan, or the region 

about the Bay of Fundy, New Brunswick. Smithsonian Contributions to Knowledge 6:39-44. 

. 1855. Descriptions of some new marine Invertebrata. Proceedings of the Academy of 

Natural Sciences of Philadelphia 7:385-394. 

Stock, J. H. 1977. Microparasellidae (Isopoda, Asellota) from Bonaire. Studies on the fauna of 

Curacao and other Caribbean islands 51(168) :69—91. 

. 1986. Two new amphipod crustaceans of the genus Bahadzia from 'blue holes' in the 

Bahamas and some remarks on the origin of the insular stygofaunas of the Atlantic. Journal 

of Natural History (4)20:921-933. 

Stone, I., and R. W. Heard. 1989. Excorallana delaneyi n. sp. (Crustacea: Isopoda: 

Excorallanidae) from the northeastern Gulf of Mexico, with observations on adult 

characters and sexual dimorphism in related species of Excorallana Stebbing, 1904. Gulf 

Research Reports) 8(2):199-211. 

Stork, H. A. 1940. A new fresh-water isopod from Curasao. Studies on the Fauna of Curacao, 

Aruba, Bonaire and the Venezuelan Islands 10:147-150. 

Strouhal, H. 1966. Eine neue Halophile Stenophiloscia aus den Rotmeergebiete (Isop. terr.). 

Annalen die Naturhistorischen Museums, Wien 69:323-333. 

Tattersall, W. M. 1905. The marine fauna of the coast of Ireland. Part 5. Isopoda. Scientific 

Investigations for 1904, Fisheries Branch, Ireland 2:1-90.


Thompson, M. T. 1902. A new isopod parasitic on the hermit crab. U. S. Fish Commission 

Bulletin for 1901, pp. 53-56. 

Topp, R. W., and F. H. HofT, Jr. 1972. Flatfishes (Pleuronectiformes). Memoirs of the 

Hourglass Cruises 4(2): 1-135. 

Ul'yanin, B. N. 1875. [Crustacea of Turkestan]. Imperatorskoe Obshchestvo Lyubitelei 

Estestvoznaniya Antropologhii i Etnoghrafii, vol. 2, pt. 6. Moscow. [In Russian.] 

Upton, N. P. D. 1987a. Asynchronous male and female life cycles in the sexually dimorphic, 

harem-forming isopod Paragnatkia formica (Crustacea: Isopoda). Journal of Zoology, London 

212:677-690. 

. 1987b. Gregarious larval settlement within a restricted intertidal zone and sex 

differences in subsequent mortality in the polygynous saltmarsh isopod Paragnatkia formica 

(Crustacea: Isopoda). Journal of the Marine Biological Association of the United Kingdom 

67(3):663-678. 

Vandel, A. 1952. Etude des isopodes terrestres recoltes au Venezuela par le Dr. G. 

Marcuzzi, suivie de considerations sur le peuplement du Continent de Gondwana. Memorie 

del Museo Civico di Storia Naturale di Verona 3:59-203. 

Vanhoffen, E. 1914. Die Isopoden der Deutschen Siidpolar-Expedition 1901-1903. Deutsche 

Sudpolar-Expedition 1901-1903, 25 (Zoologie) 7:447-598. 

Van Name, W. G. 1936. The American land and fresh-water isopod Crustacea. Bulletin of the 

American Museum of Natural History 71:1—535. 

Veillet, A. 1945. Recherches sur le parasitisme des crabes et des Galathees par les 

Rhizocephales et les Epicarides. Annales de Vlnstitut oceanographique, Paris, new series 

22(4):193-341. 

Wagele, J.-W. 1979. Morphologische Studien an Eisothistos mit Beschreibung von drei neuen 

Arten (Crustacea, Isopoda, Anthuridea). Mitteilungen aus dem Zoologiscken Museum der 

Universitat Kiel 1(2): 1-19. 

. 1981. Zur Phylogenie der Anthuridea (Crustacea, Isopoda). Mit Beitragen zur 

Lebensweise, Morphologie, Anatomie und Taxonomie. Zoologica 132:1-127. 

. 1982. On Apanthuretta lathridia n. sp. (Crustacea, Isopoda, Anthuridea) from Cuba 

Bijdragen tot de Dierkunde 52(l):43-48. 

. 1983. On the origin of the Microcerberidae (Crustacea: Isopoda). Zeitschrift fur 

zoologische Systematik und Evolutionsforschung 21 (4):249—262. 

. 1985. New west Atlantic localities for the stygobiont paranthurid Curassanthura 

(Crustacea, Isopoda, Anthuridea) with description of C. bermudensis n. sp. Bijdragen tot de 

Dierkunde 55(2):324-330. 

Walker, A. O. 1901. Contributions to the Malacostracan fauna of the Mediterranean. Journal 

of the Linnean Society of London, Zoology 28:290-307. 

Waterman, T. H., ed. 1960. The Physiology of Crustacea, vols. 1, 2, 670, 681 pp. New York & 

London: Academic Press. 

Williams, E. H., and L. B. Williams. 1980. Four new species of Renocila (Isopoda: 

Cymothoidae), the first reported from the New World. Proceedings of the Biological Society of 

Washington 93(3):573-592. 

. 1982. Mothocya bohlkeorum, new species (Isopoda: Cymothoidae) from West Indian 

cardinalfishes (Apogonidae). Journal of Crustacean Biology 2(4):570-577.


. 1985a. A new cymothoid isopod, Glossobius hemiramphi, from the mouth of the 

ballyhoo, Hemiramphus brasiliensis (Linnaeus) (Exocoetidae), in the Caribbean Sea. 

Crustaceana 48(2): 147-152. 

. 1985b. Cuna insularis n. gen. and n. sp. (Isopoda: Cymothoidae) from the gill 

chamber of the sergeant major Abudefduf saxatilis (Linnaeus) (Osteichthyes) in the West 

Indies. Journal of Parasitology 71 (2):209-214. 

. 1986. Kuna Nomen Novum for Cuna Williams and Williams, 1985, preoccupied by Cuna 

Hedley, 1902. Journal of Parasitology 72(6):879. 

Williams, L. B., and E. H. Williams. 1981. Nine new species of A nilocra (Crustacea: Isopoda: 

Cymothoidae) external parasites of West Indian coral reef fishes. Proceedings of the Biological 

Society of Washington 94(4): 1005-1047. 

Wilson, G. D. F. 1987. The road to the Janiroidea: Comparative morphology and evolution 

of the asellote isopod crustaceans. Zeitschrifi fir Zfiologische Systematik und Evolutionsforschung 

25(4):257-280.

Index 

abbreviata, Bopyrina, 110, 267 

Abudefduf saxatilis, 171, 175, 186 

abudefdufi, Anilocra, 171, 175 

Abyssianira dentifrons, 263 

Acanthocope spinosissima, 263 

acanthopkora, Domecia, 111 

Acanthopleura granulata, 215 

acanthura, Anopsilana, 124, 125 

acanthuri, Anilocra, 171, 175, 176 

Acanthurus 

bahianus, 171, 176 

chirurgus, 171, 176 

acanthurus, Macrobrachium, 112 

crenulata, 64, 65, 267 

setosa, 64, 65 

Achelion occidentalism 110 

acuminata, Nerocila, 171, 172, 173, 190, 266 

268 

forma acuminata, 190 

forma &yter, 190 

acuta, Anilocra, 268 

acutirostrata, Stenobermuda, 106, 270 

acutirostris, Processa, 113 

acutitelson, Dynamenella, 213, 214 

ilagfl, 116, 117 

antillensiSy 117 

incisa, 268 

-4i?£fl (Aega), 116, 117 

deshaysiana, 117, 266, 268 

ecarinata, 117, 268 

-dtfgfl (Rhamphion), 116, 117 

^wtata, 117, 119 

tenuipes, 117, 119 

Aegidae, 115, 116, 262 

Aetobatus narinari, 166 

Pseudione, 113, 270 

Upogebia, 112 

Agaricia, 88, 166 

agaricicola, Metacirolana, 153, 154 

A£

294 INDEX 

angulata 

Dynamene, 214 

Dynamenella, 214 

angustifrons, Hexapanopeus, 111 

Anilocra, 170, 174, 175 

abudefdufi, 171, 175 

acanthuri, 171, 175, 176 

acuta, 268 

chaetodontis, 171, 175, 177 

fAnwiw, 172, 175, 177 

haemuli, 172, 173, 175, 177 

kolacantki, 172, 175, 179 

holocentri, 175, 179 

laticauda, 268 

myripristis, 172, 175, 180 

/wr/i/i, 173, 175, 180 

annandalei, Sphaeroma, 234 

annaoides, Antarcturus, 264 

annulicornis, Pandalus, 113 

annulipes, Pagurus, 113 

anomala, Aporobopyrina, 110, 267 

anops, Creaseriella, 137 

Anopsilana, 124, 273 

acanthura, 124, 125 

ArMWH, 124, 125, 266 

owiata, 124, 125 

cubensis, 124, 126 

;m, 124, 127 

radicicola, 124, 127 

annaoides, 264 

jloridanus, 269 

Antennuloniscus dimeroceras, 263 

Antheluridae, 275 

Anthomuda, 17, 23, 291 

stenotehon, 23, 270 

Anthuridae, 16, 270 

Anthuridea, 2, 14, 15, 16, 244, 263, 269 

antiguai, Plesionika, 113 

antillense, Exosphaeroma, 229, 230 

antillensis 

Aega, 117 

Dromidia, 111 

Excorallana, 161, 162, 268 

Paranthura, 69 

Antrolana, 273 

Apanthura? 17, 25 

cracenta, 25 

cram, 25, 26 

harringtoniensis3 25, 26, 270 

Apanthuroides? 17, 26 

millae, 27 

Aphysina fistulariSy 223 

lachneri, 171, 188 

maculatus, 171, 193 

townsendi, 171, 193 

Apogonidae, 187 

Aporobopyrina, 110 

anomala, 110, 267 

Aporobopyrus, 110 

collardi, 275 

curtatus, 110 

arbuscula, Oculina, 88 

Archosargus probatocephalus, 122 

Arcturella, 252 

bispinata, 252, 269 

#wiflte, 252, 269 

Arcturidae, 251, 252, 264 

caribbaeus, 264 

purpureus, 264 

arenatus, Priacanthus, 173, 183 

Argeia, 110 

atlantica, 110 

Argeiinae, 107 

Ariusfelis, 171, 190 

Armadilloniscus; 247 

m'/zai, 247 

Iscfinomesus, 263 

Petrolisthes, 110 

armillatuSy Alpheus, 112 

arndti, Dies, 207 

aruboidesy Arubolana, 144, 145, 270 

Arubolana, 144, 273 

aruboides, 144, 145, 270 

/ma/a, 144, 145 

parvioculata, 144, 145 

ascensionis, Holocentrus, 172, 179 

Aselloidea, 75 

Asellota, 15, 15, 73, 263, 267, 269 

Astacilla, 252 

cymodocea, 252, 253 

/ajtf//

Atlantasellidae, 75 

Atlantasellus, 75 

cavernicolus, 75, 269, 270 

atlantica 

Argeia, 110 

Megalops, 172, 183 

attenuata, Erichsonella, 256, 258, 269 

aurolineatum, Haemulon, 172, 178 

Azygopleon, 110 

schmitti, 110, 267 

bacescui, Phycolimnoria, 276 

Bagatus, 83 

Bahalana, 124, 127, 128, 273 

cardiopus, 128 

geracei, 128 

mayana, 128 

bahianus, Acanthurus, 171, 176 

bajonado, Calamus, 122 

Balanopleon, 110 

tortuganus, 110 

Balistes vetula, 122 

balthica, Idotea, 259 

barbadensis, Micropanope, 111 

barbarae, Geogerceis, 215 

6anzar

296 INDEX 

Cancrion carolinus, 111 

capistratus, Chaetodon, 171, 177 

caraibica, Cymothoa, 182, 268 

Caranx, 122, 171 

hippos, 171, 183 

latus, 171, 183 

ruber, 171, 183 

carbonarium, Haemulon, 172, 178 

carcinus, Macrobrachium, 112 

cardiopus, Bahalana, 128 

caribaeus, Dactylokepon, 111 

caribbaeus, Arcturus, 264 

caribbea, Storthyngura pulchra, 263 

Ianirella, 263 

Malacanthura, 45 

Ischnomesus, 263 

Macrostylis, 263 

caribea, Uromunna, 94 

carinata, 'Cerceis,' 211, 268 

carolii, Metaphrixus, 111 

carolinense, Tozeuma, 112 

carolinus, Cancrion, 111 

Gzr/u

Colopisthus, 144, 146 

parvus, 147, 270 

confixa, Cortezura, 31 

Conilera cylindracea, 268 

Conilerinae, 123, 139 

conklini, Phaeoptyx, 173, 188 

constricta, Munida, 112 

convexa, Eurydice, 147, 148, 149, 268 

coralicolay Joeropsis, 88, 267 

Corallanidae, 115, 157 

corallicola, Minyanthura, 53 

corallinus, Pylopagurus, 112 

Cortezura, 17, 29 

confixa, 31 

penascoensis, 31 

cracenta, Apanthura, 25 

crassa, Virganthura, 73 

Creaseriella, 124, 137, 273 

anops, 137 

crenata, Anopsilana, 124, 125 

crenulata, Accalathura, 64, 65, 267 

crenulitelson, Cirolana, 132, 133 

cromis, Pogonias, 173, 190 

cruciaria, Astalione, 110 

cruris, Apanthura, 25, 26 

cruentatus, Epinephelus, 172, 179 

crumenophthalmus, Selar, 173, 183 

Cryptoniscoidea, 107 

Crytoniscidae, 107, 109 

cubana, Cyathura (Cyathura), 33 

cubense, Munidion, 111 


tfocme/a, 119, 120 

cuborientalis, Cyathura (Stygocyathura), 34, 35 

culebrae, Vandeloscia, 247, 251 

cumanensis, Malacanthura, 45 

cumulus, Agarna, 173 

cuprea, Diopatra, 256 

curacaoensis, Hippolyte, 110 

Curassanthura, 64, 67, 270 

bermudensis, 67, 270 

canariensis, 270 

Afl/mfl, 67, 68, 270 

curassavica, Cyathura (Stygocyathura), 35 

o/rn, Haliophasma, 41 

curtatus, Aporobopyrus, 110 

cuvieri, Galeocerdo, 122 

cyaneus, Chromis, 172, 177 

Cyathura, 17, 31, 270, 273 

/>o/z7a? 267 

(Cyathura), 31 

cubana, 33 

(Stygocyathura), 31, 33, 35, 270 

cuborientalis, 34, 35 

curassavica, 35 

hummelincki, 35 

motasi, 35, 36 

orghidani, 35, 36 

parapotamica, 35, 36 

salpiscinalis, 35, 38 

sbordonii, 35, 38 

specus, 35, 38 

univam, 35, 38 

Cyclograpsus interger, 111 

cylindracea, Conilera, 268 

Cymodoce, 226, 227 

barrerae, 227 

ruetzleri, 227 

Cymodocea, 223, 253 

cymododea, Astacilla, 252, 253 

Cymothoa, 170, 172, 182 

caraibica, 182, 268 

«OM, 172, 173, 182, 268 

INDEX 297 

wrfrem, 171, 172, 173, 182, 183, 268 

Cymothoidae, 115, 169, 170 

Cynoscion, 172, 183 

nebulosus, 172, 183 

Dactylokepon 

caribaeus, 111 

sulcipes, 267 

Dajidae, 107 

Dardanus jucosus, 1122 

Dasyatis americana, 122, 166 

decorata, Mesanthura, 53 

deformans, Synsynella, 110, 113, 268, 270 

delaneyi, Excorallana, 160, 161 

Dendrotiidae, 263 

Dendrotion hanseni, 263 

dentata, Aega, 117, 119 

dentifrons, Abyssianira, 263 

deplanata, Ceratothoa, 180 

depressa, Panoplax, 111 

depressus, Ancinus, 268 

deshaysiana, Aega, 117, 266, 268 

Desmosoma magnispina, 263 

Desmosomatidae, 263 

destructor, Sphaeroma, 235 

desultor, Asymmetrione, 110 

rfifl/M*, Paradella, 224, 266, 268 

Dicropleon 

periclimenis, 111 

Dictyota, 219 

Z)r>j, 207 

flnw/rf, 207 

barnardi, 207

298 INDEX 

dimeroceras, Antennuloniscus, 263 

diminuta, Exosphaeroma, 229, 231 

diogenes, Petrochirus, 110 

Diopatra cuprea, 256 

Diplophryxus, 111 

Discerceis, 210, 211 

linguicauda, 213 

dispar, Paraliomera, 111 

distorta, Leidya, 111 

Domecia 

acanthophora, 111 

hispida, 111 

Dromidia antillensis, 111 

dubitans, Angliera, 91 

Dynamene angulata, 214 

Dynamenella, 210, 213, 214, 224 

acutitelson, 213, 214 

var. glabrothorax, 214 

var. typica, 214 

angulata, 214 

perforata, 213, 215, 270 

quadrilirata, 213, 215 

Dynameninae, 204, 210, 211 

ecarinata, Aega, 117, 268 

Echinothambema ophiuroides, 263 

Echinothambematidae, 263 

edithae, Paracerceis, 218, 221 

Edo tea 

lyonsi, 269 

montosa, 269 

samariensis, 276 

edulis, Processa, 113 

edwardsi, Plesionika, 113 

Edwinjoycea horologium, 252, 269 

eglanteria, Raja, 122 

Eisothistos, 16, 38, 39 

petrensis, 39 

ten, 39 

mm, Plesionika, 113 

Entoniscidae, 107, 109 

Entophilinae, 107 

Eophrixus subcaudalis, 111, 267 

Epicaridea, 4, 14, 107, 267 

Epinephelus, 122, 172, 183 

cruentatus, 172, 179 

>/z^, 172, 179 

guttatus, 172, 179 

*7, 122, 172, 190 

morio, 122 

Erichsonella, 255, 256, 257 

attenuata, 256, 258, 269 

Jiliformis, 256, 258, 269 

tropicalis, 257 

floridana, 256, 258, 269 

isabelensis, 269 

Eriphia gonagra, 111 

Eubranchiatae, 203 

Eurycopidae, 263 

Eurydice, 143, 147 

con^xfl, 147, 148, 149, 268 

littoralis, 148, 149, 268 

personata, 147, 149, 270 

piperata, 147, 149, 268 

Eurydicinae, 123, 139, 143 

excavata, Chiridotea, 269 

Excirolana, 144, 149, 150 

braziliensis, 150, 266, 268 

mayana, 150, 153, 268 

Atrial, Cymothoa, 172, 173, 182, 268 

Excorallana, 157, 159, 161 

antillensis, 161, 162, 268 

berbicensis, 161, 162 

delaneyi, 160, 161 

fissicauda, 161, 162 

mexicana, 160, 161, 268 

oculata, 161, 163 

quadricornis, 161, 165, 270 

sexticornis, 161, 165 

subtilis, 160 

tricornis, 266, 268 

occidentalis, 167 


warmingii, 161, 167 

exilipes, Palaemonetes, 113 

Exocoettus, 172, 184 

Exosphaeroma, 226, 229, 231 

fl/Afl, 229 

antillense, 229, 230 

diminuta, 229, 231 

productatelson, 229, 231 

yucatanum, 229, 231 

exotica, Ligia, 247, 249 

faber, Chaetodipterus, 171, 190 

fasciata, Mesanthura, 47, 49 

faustinum, Macrobrachium, 112 

/flxwii, Harrieta, 232, 268 

felis,Arius, 171, 190 

filiforme, Syringodium, 253, 260 

Jiliformis, Erichsonella, 256, 258, 269 

fimbriata 

Pleurocryptella, 112 

Processa, 113 

fissicauda, Excorallana, 161, 162 

fistularis, Aphysina, 223 

Flabellifera, 2, 14, 114, 115, 268, 269 

Jlavolineatum, Haemulon, 122, 172, 178

Jlinti, Munida, 111 

jloridana 

Erichsonella, 256, 258, 269 

Pleurocrypta, 112 

florid anus 

Allodiplophryxus, 267 

Antarcturus, 269 

Thor, 110, 111 

jloridensis 

Carpias, 267 

Gnathic, 268 

Mesanthura, 53, 267 

Paranthura, 69, 71, 267 

Pleurocope, 98, 267 

joetens, Synodus, 173, 183 

foliatus, Parathelges, 112 

formosa, Kupellonura, 267 

formosus, Alpheus, 111, 112 

fritzmuelleri, Synalpheus, 110, 111 

jucorum, Latreutes, 112 

Jucosus, Dardanus, 112 

Julvus, Epinephelus, 172, 179 

Jurcifer, Paranthias, 173, 179 

Galathea rostrata, 112 

galathinus, Petrolisthes, 110 

Galeocerdo cuvieri, 122 

geminsula, Amakusanthura, 18 

Geocerceis, 210, 215 

barbarae, 215 

geracei, Bahalana, 128 

Genes rhombeus, 172, 187 

giardi, Synalpheion, 113 

giganteus, Bathynomus, 131, 268 

Gigantione 

mortenseni, 111, 267 

uberlackerae, 267 

Ginglymostoma cirratum, 122, 158 

glabrothorax, var., Dynamenella acutitelson, 214 

Glossobius, 170, 172, 183, 184 

hemiramphi, 172, 184 

impressus, 172, 184 

Paracerceis, 218, 221 

Paraleptosphaeroma, 210, 266 

Cwato, 238 

beethoveni, 238, 239 

Jloridensis, 268 

gonzalezi, 238, 239 

Johanna, 238, 239, 276 

magdalenensis, 238, 239 

puertoricensis, 238, 239 

ra^z, 238, 241 

samariensis, 238, 241 

triospathiona, 238, 241 

ye/

300 INDEX 

hemiramphi3 Glossobius, 172, 184 

Hemiramphidae, 187 

Hemiramphus 

bermudensis, 188 

brasiliensis, 172, 184 

hemphilli, Synalpheus, 110, 111 

hendleri, Pendanthura, 56 

herbstii, Panopeus, 111 

herrerai, Microcharon, 93 

heterocarpus, Plesionika, 113 

heterochaelis, A Ipheus, 112 

Heteromesus bijurcatus, 263 

Hexapanopeus angustifrons, 111 

Hippolyte 

curacao ensis 3 110 

pleutacanthus, 110, 111 

Zostericola, 110 

hippos, Caranx, 171, 183 

Hirundichthys speculifer, 172, 184 

hispida, Domecia, 111 

holacanthi, Anilocra, 172, 175, 179 

Holacanthus tricolor, 172, 179 

holocentri, Anilocra, 175, 179 

Holocentrus ascensionis, 172, 179 

hopkinsi, Mesanthura, 47, 51, 267 

Horoloanthura irpex, 267 

horologium, Edwinjoycea, 252, 269 

hummelincki, Cyathura (Stygocyathura), 35 

Hydroniscus quadrifrons, 263 

Hyperphrixus castrensis, 267 

Hypoconcha 

sabulosa, 111 

spinosissima, 111 

Hyporhamphus unifasciatus, 172, 188 

hyptius, Stegophryxus, 113, 268 

Hyssuridae, 16, 58, 60 

fanirella 

caribbica3 263 

vemae, 263 

Afotai, 255, 259 

balthica, 259 

metallicay 259, 269 

Idoteidae, 251, 252, 255 

Idoteinae, 255, 256 

Iliacantha 

liodactyla, 111 

subglobosa3 111 

imbricata, Parapagurion, 112 

impressa, Politolana, 140 

impressus, Glossobius, 172, 184 

imswe, Kupellonura, 60 

imw/a, Arubolana, 144, 145 

/nriftZj 4*£A, 268 

indica, Limnoria, 194, 195 

infundibulata, Paranthura, 69, 71, 270 

insulae, Limnoria, 195 

insular is 

Alcirona, 158 

A'ttrtfl, 170 

Rocinela, 119, 120, 268 

integra, Synsynella, 268 

interger, Cyclograpsus, 111 

intermedius, Palaemonetes, 113 

loninae, 107 

Iridopagurus, 112, 113 

zm, 113 

z'm, Iridopagurus, 113 

irmae, Haliophasma, 43 

fr/fcx, Horoloanthura, 267 

irrasa, Munida, 111 

irritans, Munidion, 111 

isabelensis, Erichsonella, 269 

Ischnomesidae, 263 

Ischnomesus 

armatus, 263 

caribbicus, 263 

multispinis, 263 

Ischyromene, 210, 214, 217 

bamardi, 218 

iVfljflrfl, Epinephelus, 122, 172, 190 

jacobus, Myripristis, 172, 180 

jacqueti, Sclerocrangon, 110 

Janiridae, 80, 81, 263 

Janiroidea, 75, 79, 80, 81 

Joeropsidae, 80, 87 

Joeropsis, 87, 88 

bifasciatus, 88 

coralicola, 88, 267 

personatus, 88, 90 

rathbunae, 88, 90, 267, 270 

JOAHHM, G/ifltfifl, 238, 239, 292 

jonesi, Anopsilana, 124, 127 

kadiakensis, Palaemonetes, 113 

kensleyi, Mexicope, 81, 267 

kohleri, Mesosignum, 264 

AWAJI, Alcirona, 158, 268, 270 

tfiwifl, 170, 184 

insularis, 171, 186 

Kupellonura, 60 

formosa, 267 

imswe, 60 

lachneri, Apogon, 171, 188 

Lachnolaimus maximus, 122 

Laminaria, 201 

lamprotaenia, Anchoa, 171, 187 

laodicense, Gnathostenetroides, 78

lasallae, Astacilla, 252, 253 

lata, Parabopyrella, 112 

lathridia, Amakusanthura, 18, 20 

laticauda, Anilocra, 268 

laticeps, Skuphonura, 58 

latreillei, Tylos, 247, 250 

Latreutes Jucorum, 112 

latreuticola, Probopyrinella, 112, 268, 270 

latus, Caranx, 171, 183 

/flw#?, Astacilla, 269 

Laurencia, 219 

Aimini, 111, 270 

distorta, 111 

Leiostomus xanthurus, 172, 183, 187, 

190 

Lepisosteus spatula, 172, 190 

leptorhynchus, Pandalus, 113 

lewisi, Chalixanthura, 27, 29 

Licranthura, 16, 43 

amyle, 43 

Lfcifl, 247, 249 

baudiniana, 247, 249 

wrrtiai, 247, 249 

302 INDEX 

mayana 

Bahalana, 128 

Excirolana, 150, 153, 268 

mcclendoni, Synalpheus, 110, 111 

Megalops atlantica, 172, 183 

menziesi, Metacirolana, 153, 154 

Mesantkura, 17, 45 

bivittata, 47 

decorata, 53 

fasciata, 47, 49 

JloridensiSy 53, 267 

hopkinsit479 51, 267 

looensis, 47, 51 

paucidens, 47, 51 

pulchra, 47, 52, 267 

punctillata, 47, 53 

reticulata, 47, 53 

Mesosignidae, 264 

Mesosignum kohleri, 264 

Metacirolana, 144, 153 

agaricicola, 153, 154 

Afl/ifl, 153, 154 

menziesi, 153, 154 

sphaeromiformis, 153, 154 

metallica, Idotea, 259, 269 

Metaphrixus carolii, 111 

mexicana, Excorallana, 160, 161, 268 

mexicanus, Microcerberus t 269 

Mexicope, 80, 81 

kensleyiy 81, 267 

Mexilana, 273 

mgrayiy Serolis, 202, 268 

Microcerberidae, 243, 244 

Microcerbendea, 14, 243, 269, 273 

Microcerberus, 244, 273 

littoralis, 244 

mexicanuSy 269 

minutuSy 244 

mirabiliSy 244 

nunezi, 244 

renaudiy 244 

simplex, 244 

syrtiaiSy 244 

Microcharon, 90, 91 

herrerai, 93 

phreaticus, 93 

sabulurrty 91 

Micropanope barbadensis, 111 

Microparasellidae, 80, 90, 273 

Microphrys bicornutus, 110 

//nV^r, Munida} 112 

millae, Apanthuroides, 27 

milleri, Santiay 99, 267 

minocule, Stenetrium, 100, 102 

minusy Synalpheus, 110, 113 

minuta, Cirolana, 132, 135 

Cfl#wj, 82, 84, 270 

MacrostyliSy 263 

Microcerberusy 244 

Minyanthura, 16, 53 

corallicola, 53 

mirabilis 

MadraciSy 29, 41 

Microcerberusy 244 

Miratidoteay 255, 259 

bruscai> 260 

Monacanthus ciliatuSy 172, 190 

montaguiy Pandalus, 113 

montosa, Edoteat 269 

morioy Epinephelus, 122 

Gigantione, 111, 267 

Parabopyrellay 112, 267 

mosaica, Cassidinidea, 208 

motasiy Cyathura (Stygocyathura), 35, 36 

Mothocya, 170, 187 

bohlkeorum, 171, 173, 187, 188 

wflwfl, 172, 187, 188 

Mugil cephalus, 172, 190 

multilineatus, Chromis, 172, 177 

multipunctata, Limnoria, 195, 196 

multispinis, Ischnomesus, 263 

constricta, 112 

TKnrt, 111 

irrasas 111 

longipeSy 112 

772 £/w, 112 

schroederiy 112 

simplex, 110 

stimpsoni, 111 

valida, 110 

Munidion 

cubensCy 111 

irritans, 111 

longipedisy 112, 267 

Munna, 93 

petronastes, 94 

Munnidae, 80, 93 

Munnogoniumy 96 

wilsoni, 96 

Mycteroperca 

bonaci, 122 

venenosa, 122 

Myripristis jacobus, 172, 180

myripristis, Anilocra, 172, 175, 180 

Nalicora, 157, 168 

rapax, 169, 268 

nana, Mothocya, 172, 187, 188 

Nannoniscidae, 264 

Nannoniscus camayae, 264 

narinari, Aetobatus, 166 

Natatolana, 139 

gracilis, 140 

nebulosus, Cynoscion, 172, 183 

Nemanthura, 43 

Neoanthura coeca, 263 

Neopanope 

packardii, 111 

ttxana sayi, 111 

Neostenetroides, 77, 78 

stocki, 78 

Nerocila, 170, 172, 188 

acuminata, 171, 172, 173, 190, 266, 268 

forma acuminata, 190 

forma aj/er, 190 

ninae, Armidilloniscus, 246, 247 

nfzwtf, T^/ar, 247, 250 

normanni, Alpheus, 112 

northropi, Palaemon, 113 

nunezi, Microcerberus, 244 

nuttingi, Paracerceis, 218, 223 

obtruncata, Cirolana, 132, 135, 268 

occiden talis 

Achelion, 110 

Cleantioides, 256 

Excorallana tricornis, 167 

Parathelges, 112 

ocellatus, Chaetodon, 171, 177 I 

Excorallana, 161, 163 

Rocinela, 119, 120, 266, 268 

Oculina arbuscula, 88 

Ocyurus chrysurus, 172, 183 

owfram, Cymothoa, 171, 172, 173, 182, 183, 

268 

ohione, Macrobrachium, 112 

Olencira praegustator, 268 

Zigia, 247, 249 

Macrobrachium, 112 

Ontilorpheus, 124, 139 

stebbingi, 139 

Oniscidea, 1, 4, 14, 15, 246, 247, 262 

ophiuroides, Echinothambema, 263 

Orbioninae, 107 

orghidani, Cyathura (Stygocyathura), 35, 

36 

Orthopristis 

chrysoptera, 172, 183 

raA^r, 122, 173, 179 

ovalis 

Cassidinidea, 207, 208, 268 

Lironeca, 268 

Ovobopyrus alphezemiotes, 267 

oxyophthalmus, Paguristes, 112 

Pachygrapsus transversus, 111 

packardii, Neopanope, 111 

Paguristes 

oxyophthalmus, 112 

tortugae, 112 

Pagurus 

annulipes, 113 

bonairensis, 110, 113 

brevidactylus, 112, 113 

longicarpus, 110, 113 

provenzanoi, 110, 112, 113 

Palaemon 

northropi, 113 

pandaliformis, 113 

Palaemonetes 

exilipes, 113 

intermedius, 113 

kadiakensis, 113 

paludosus, 113 

pugio, 113 

vulgaris, 113 

paludosus, Palaemonetes, 113 

pandalicola, Probopyrus, 112, 266 

pandaliformis, Palaemon, 113 

Pandalus 

annul ico rni s, 113 

Ari, 113 

leptorhynchus, 113 

montagui, 113 

pandionis, Synalpheus, 111, 113 

Panopeus herbstii, 111 

Panoplax depressa, 111 

Parabopyrella 

lata, 112 

mortenseni, 112, 267 

richardsonae, 112, 267 

thomasi, 112 

Parabopyriscus stellatus, 267 

Paracerceis, 210, 218, 223 

cflurfate, 219, 268, 270 

var. brevipes, 219 

roA«ifl*, 219, 220 

wfifAae, 218, 221 

£/>wii, 218, 221 

Hatting, 218, 223

304 INDEX 

Paradella, 210, 214, 223 

dianae, 224, 266, 268 

plicatura, 223, 224 

quadripunctata, 223, 224 

tumidicauda, 223, 226 

Paradynamene benjamensis, 268 

Paragnathia, 237 

Paraleptosphaeroma, 207, 208 

îww, 210, 266 

Paralimnoria, 193, 199 

andrewsi, 199 

forma A, 200 

forma B, 201 

forma typica, 200 

Paraliomera dispar, 111 

Paramunnidae, 80, 96 

Paranthias farcifer, 173, 179 

Paranthura, 64, 69 

antillensis, 69 

barnardiy 69, 71 

floridensis, 69, 71, 267 

infundibulatay 69, 71, 270 

Paranthuridae, 16, 64, 263, 270 

Parapagurion imbricata, 112 

Parapagurus, 112 

parapotamica, Cyathura (Stygocyathura), 35, 

Para theIges 

foliatus, 112 

occidentalism 112 

piriformis, 112, 270 

tumidipes, 112, 270 

partiti, Anilocra, 173, 175, 180 

partitus, Pomacentrus3 173, 180 

parva, Cirolana, 132, 135, 268 

parvioculata, Arubolana, 144, 145 

parviramus, Pseudione, 276 

parvus, Colopisthus, 147, 270 

patulipalma, Stenetrium, 100, 102 

paucidens, Mesantkura, 47, 51 

pectiniger, Synalpheus, 110, 111, 113 

penascoensis; Cortezura, 31 

Pendanthura, 17, 56 

hendleri, 56 

tanaiformis, 56, 270 

penna, Calamus, 122 

perforata, Dynamenella, 213, 215, 270 

Periclimenes 

americanus, 111, 113 

longicaudatus, 113 

periclimenis, Dicropleon, 111 

personata, Eurydice, 147, 149, 270 

personatus, Joeropsis, 88, 90 

petrensis, Eisotkistos, 39 

Petrochirus diogenes, 110 

Petrolisthes 

armatus, 110 

galathinus, 110 

marginatus, 110 

petronastesy Munna, 94 

pfefferi, Limnoria, 195, 198 

Phaeoptyx 

conklini, 173, 188 

pigmentaria, 173, 188 

phreaticuSy Microcharoriy 93 

Phreatocoidea, 14 

Phycolimnoria, 193, 201 

bacescuiy 276 

clarkae, 201 

Phyllodurinae, 107 

pigmentariay Phaeoptyx, 173, 188 

piperata, Eurydice, 147, 149, 268 

piriformis, Parathelges, 112, 270 

planicauda, Cleantioides, 256, 266, 268 

Platybranchiatae, 203 

platycauda, Limnoria, 195, 198 

Plesionika 

antiguai, 113 

edwardsiy 113 

mw, 113 

heterocarpus, 113 

martia, 113 

p leuracanthus, Hip poly te, 110, 111 

Pleurocope, 97 

floridensisy 98, 267 

Pleurocopidae, 80, 81, 96 

Pleurocryptella fimbriata, 112 

plicatura, Paradella, 223, 224 

plicifera, Callispongia, 221 

plumieri, Haemulon, 172, 179 

Pogonias cromis, 173, 190 

polita 

Cyathura, 267 

Politolana, 140, 143, 268 

Politolana, 139, 140 

impressa, 140 

ô/ito, 140, 143, 268 

Pomacentrus partitus, 173, 180 

Pontonia margarita, 113 

Porce liana say ana, 110 

A>n7^, 88, 90 

poteriophora, Akidognathia, 264 

praegustator, Olencira, 268 

Priacanthus arenatus, 173, 183 

probatocephalus, Archosargus, 122 

Probopyria alphei, 112, 267

Probopyrinella 

heardiy 267 

latreuticola, 112, 268, 270 

Probopyrus pandalicola, 112, 266 

Processa 

acutirostris y 113 

canaliculata y 113 

eduliSy 113 

Jimbriatay 113 

tenuipes, 113 

processaey Urobopyrusy 113, 268 

Prodajus cf. bigelowiensis, 268 

productatelson, Exosphaeroma3 229, 231 

provenzanoiy Pagurus> 110, 112, 113 

psamathus, Anglieray 91 

Pseudasymmetrione, 113 

Pseudione 

qffinis, 113, 270 

cognata, 268 

parviramuSy 276 

upogebiae, 268 

Pseudioninae, 107 

Ptilanthura tricarina, 267 

puertoricensis, Gnathia} 238, 241 

pugilatoty Uca, 111 

GnathostenetroideSy 77', 267 

PalaemoneteSy 113 

pulchra 

caribbedy Storthyngura, 263 

Mesanthura, 47, 52, 267 

punctatuSy Carpias, 82, 85 

punctillatdy Mesanthura, 47, 53 

purpureus, Arcturus3 264 

PylopaguruSy 110 

corallinus, 112 

quadricornisy Excorallana, 161, 165, 270 

quadridentata, Sphaeroma, 234, 268 

quadrifrons, Hydroniscus, 263 

quadriliratdy Dynamenella, 213, 215 

quadripunctata, Paradella, 223, 223 

racovitzai, Angliera, 91 

radicicola 


Xylolana, 157 

i?a/a eglanteridy 122 

rapaXy Nalicora, 169, 268 

rathbunae 

JoeropsiSy 88, 90, 267, 270 

Lysmata, 112 

rafAi, Gnathic, 238, 241 

redmanni, Lironecay 172, 173, 186, 268 

regalis, Scomberomorus, 173, 187 

regina, Astacilla, 252, 253 

renaudiy Microcerberus, 244 

Renocila, 170, 191 

bowmani, 173, 191 

«/!/!!, 171, 191 

waldneriy 173, 191, 193 

reticulatas Mesanthura, 47, 53 

INDEX 305 

reynoldsiy Uromunnay 94, 95, 266, 267 

RhamphioUy see i4«ga (Ramphion) 

Rhizophora mangle, 235 

rhombeus, GerreSy 172, 187 

Rhyscotus, 247, 249 

fexwuij, 247, 249 

richardsonae, Parabopyrella, 112, 267 

ricordiy Sesarma, 111 

nVrf/z, Vandeloscia, 247, 251 

Ritkropanopeus karrisiiy 111 

Rocinela, 116, 119 

cubensisy 119, 120 

insularis, 119, 120, 268 

oca/ate, 119, 120, 266, 268 

jignafa, 119, 120, 266, 268 

rostratay Galathea, 112 

Caranx, 171, 183 

Orthopristisy 122, 173, 179 

ruetzleri, Cymodoce, 227 

sabulosa, Hypoconcha, 111 

sabulum, Microcharony 91 

salpiscinalis, Cyathura (Stygocyathura), 35, 38 

samariensis 

Edotea, 276 

GnafAia, 238, 251 

Santiay 98 

ffiiV/m, 99, 267 

Santiidae, 80, 98 

Sargassum, 84, 201 

saseboensiSy Limnoria, 195, 198 

saxatilis, Abudefdufy 171, 175, 186 

sayana, Porcellana, 110 

ja>^", Neopanope texana, 111 

sbordoniiy Cyathura (Stygocyathura), 35, 

38 

Schizobopyrina urocaridis, 113, 268 

schmittiy Azygopleon, 110, 267 

schoepfi 

Alutera, 171, 190 

Chilomycterusy 172, 190 

schroederi, Munida, 112 

sciuruSy Haemulon, 172, 179 

Sclerocrangon jacqueti, 110 

Scomberomorus 

cavalla, 173, 187

306 INDEX 

Scomberomorus (cont.) 

maculatus, 173, 187 

regain, 173, 187 

scopulosa, Chalixanthura, 27, 29 

sedentarius, Chaetodon, 171, 177 

Selar crumenophthalmuSy 173, 183 

Serolidae, 114, 115, 201 

Serolis, 202 

mgrayi, 202, 268 

Serranus tigrinus, 173, 191, 193 

serrata, Spinianirella, 263 

serratum, Stenetrium, 100, 102 

serricaudus, Carpias, 82, 87 

Sesarma ricordi, 111 

seticornis, Stenorhynchus, 110 

setifer, Macrostylis, 264 

setosa, Accalathura, 64, 65 

sexticorniSy Excorallana, 161, 165 

Amakusanthura, 18, 21 

Rocinela, 119, 120, 266, 268 

signijica, Amakusanthura, 18, 23 

Microcerberus, 244 

Munida, 110 

simulata, Limnoria, 195, 198 

Skuphonura, 17, 58 

laticeps, 58 

lindae, 267 

snanoi, Storthyngura, 263 

somala, Haptolana, 138 

Sparisoma viride, 122 

spathulicarpusy Stenetrium, 100, 104 

spatula, Lepisosteus, 172, 190 

speculifer, Hirundichthys, 172, 184 

specus, Cyathura (Stygocyathura), 35, 38 

Speocirolana, 273 

Sphaerolana> 273 

Sphaeroma, 226, 232, 234 

annandalei, 234 

destructor, 235 

quadridentata, 234, 268 

terebrans, 234, 235, 268 

ttWfen, 234, 235 

Sphaeromatidae, 114, 115, 202, 204 

Sphaeromatinae, 204, 226 

sphaeromiformis, Metacirolana, 153, 

154 

Sphoeroides maculatus, 173, 190 

Sphyraena barracuda, 122 

j/>i»ata, Arcturella, 252, 269 

Spinianirella serrata, 263 

spinosissima 

Acanthocope, 263 

Hypoconcha, 111 

Oncilorpheus, 139 

Stenetrium, 100, 104, 267, 270 

Stegias clibanarii, 113, 270 

Stegophryxus hyptius, 113, 268 

steindachneri, Haemulon, 122 

Astrapogon, 171, 188 

Parabopyriscus, 267 

Stenetriidae, 99 

Stenetrioidea, 99 

Stenetrium, 99, 100 

bowmani, 100 

minocule, 100, 102 

patulipalma, 100, 102 

serratum, 100, 102 

spathulicarpus, 100, 104 

stebbingi, 100, 104, 267, 270 

Stenobermuda, 99, 106 

acutirostrata, 106, 270 

Stenorhynchus seticornis, 110 

stenotelson, Anthomuda, 23, 270 

stimpsoni, Munida, 111 

stocki, Neostenetroides, 78 

pulchra caribbea, 263 

snanoi, 263 

striata, Yvesia, 246 

striatus, Chaetodon, 171, 177 

Stygocyathura, see Cyathura (Stygocyathura) 

subcaudalis, Eophrixus, 111, 267 

subglobosa, Iliacantha, 111 

subtilis, Excorallana, 160 

sulcipes, Dactylokepon, 267 

surinamensis, Batrachoides, 171, 190 

surinamicum, Macrobrachium, 112 

symmetricus, Ambidexter, 113 

synagris, Lutjanus, 172, 183 

synalphei 

Bopyrione, 110, 267 

Hemiarthrus, 111, 267 

Synalpheion giardi, 113 

Synalpheus 

bousfieldi, 110 

brevicarpus, I ] 0 

AiwwW, 110, 111, 113 

fritzmuelleri, 110, 111 

goodei, 110, 111 

hemphilli, 110, 111

longicarpus, 110, 111, 113 

mcclendoni, 110, 111 

minus, 110, 113 

pandionis, 111, 113 

pectiniger, 110, 111, 113 

Synodus foe tens, 173, 183 

Synsynella, 110, 113 

choprae, 113, 268, 270 

deformans, 110, 113, 268, 270 

Integra, 268 

Syringodium, 166, 232 

filiforme, 253, 260 

syrticus, Microcerberus, 244 

tanaiformis, Pendanthura, 56, 270 

Tecticeps, 204 

Tecticipitinae, 204 

/l^, 117, 119 

Processa, 113 

tenuis, Colanthura, 65, 270 

tenuistylis, Lironeca, 171, 186, 187 

terebrans, Sphaeroma, 234, 235, 268 

ten, Eisothistos, 39 

testudinea, Thalassia, 251 

texana, Lironeca, 268 

texensis, Rhyscotus, 247, 249 

Thalassia, 166, 232 

testudinea, 251 

Thermarcturus, 252, 255 

venezuelensis, 255 

thomasi, Parabopyrella, 112 

floridanus, 110, 111 

manningi, 111 

f/Wn, Bopyrinella, 110 

tigrinus, Serranus, 173, 191, 193 

tortugae, Paguristes, 112 

tortuganus, Balanopleon, 110 

Tozeuma carolinense, 112 

transversa, Ceratothoa, 268 

transversus, Pachygrapsus, 111 

tricarina, Ptilanthura, 267 

tricarinata, Booralana, 275 

trichostoma, Haptolana, 138 

Clibanarius, 110, 112, 113 

Holacanthus, 172, 179 

tricornis, Excorallana, 161, 165, 266, 268 

occidentalis, 167 


Tridentella, 236 

virginiana, 236 

Tridentellidae, 115, 235, 236 

triospathiona, Gnathia, 238, 241 

tripunctata, Limnoria, 199 

/n/ow, Carpias, 82, 87 

Troglocirolana, 273 

tropicalis 

Erichsonella filiformis, 2bl 

Haplomesus, 263 

Lironeca, 268 

INDEX 307 

tuberculata, Limnoria, 194, 195, 199, 268 

tuberculatus, Chiton, 229 

tumidicauda, Paradella, 223, 226 

tumidipes, Parathelges, 112, 270 

Turbinaria, 166, 219 

7>/Wj 247, 250 

/

308 INDEX 

Virganthura, 64, 73 

crassa, 73 

virginalis, Gnathia, 238, 243 

virginiana, Tridentella, 236 

viridari, Alpheus, 112 

viride, Sparisoma, 122 

vittatus, Clibanarius, 110, 112 

vulgaris, Palaemonetes, 113 

waldneri, Renocila, 173, 191, 193 

walkeri, Sphaeroma, 234, 235 

warmingii, Excorallana, 161, 167 

wegeneri, Tylos, 247, 250 

wilsoni, Munnogonium, 96 

wolffi, Bopyrissa, 110 

wurdemanni, Lysmata, 112 

xanthurus, Leiostomus, 172, 183, 187, 190 

Xenanthura, 60 

brevitelson, 62, 267 

A>/

6 - Sphaeromatidae::“Cute As Buttons”

Create successful ePaper yourself

Delete template?

Save as template?