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Guide to the
MARINE ISOPOD CRUSTACEANS
of the Caribbean
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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
CIP
British Library Cataloging-in-Publication Data available
Manufactured in the United States of America 10298 SOo 7s 6) Dae 4 mee 98 97 96 95 94 93 92 91 90 89
oo The paper used in this publication meets the minimum requirements of the American National Standard for Performance of Paper for Printed Library Materials Z39.48-1984
Contents
l Introduction
] HISTORIC BACKGROUND 3 GEOGRAPHIC AREA COVERED IN THIS GUIDE 4. ARRANGEMENT OF THE GUIDE AND HOW TO USE IT
5 | ACKNOWLEDGMENTS 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
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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
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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 Muséum 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
AESTHETASG. 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.
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GLOSSARY
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.
9
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.
MANCA. 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.
MULTIARTICULATE. 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.
GLOSSARY 11
PRODUCED. Extended or lengthened.
PROPODuUS. Article 6 of pereopod.
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.
SCLEROTIZED. 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.
SUBCHELATE. Having a subchela, forming a pincerlike structure, especially by the dactylus folding back on the propodus.
SUTURE. AA 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.
TRACHEATE. 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.
TRIUNGUICULATE. 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.
UNIARTICULATE. 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 Calman, 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 | 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 ISOPODA
Key to suborders of Isopoda
1. Parasitic on crustaceans; body of 2 nearly always asymmetrical PE hed Pere ek, IN eae SIME, Sales hale aiclincd brea Epicaridea Free-living or parasitic on fishes; body of ? bilaterally symmetrical, or if parasitic: 2 semewhat Gistorted) A505 - pct 40 - = ee epee ne ee 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’ >... 4 e5 Fo <b oe er tocar 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 imterstitial farmisys). eset s orden ake 3s AS bicenneted oe beve Anthuridea @!¢ Uropods terminal, exopod lacking; minute interstitial forms cealidinttiansd sie Baas viasteetide ahyetien see syeleieg Microcerberidea
6. Antennules minute; terrestrial forms, with pleopods tracheate
SN pe Ce ar reer ROR Al SST ard ry Sowa Ue ros pie Bait ioe3 Oniscidea Antennules rarely minute; aquatic forms, pleopods never tracheate .. 7 : : : 75"\ 7. Uropods ventral, operculiform, covering pleopods .......... Valvifera ?'~ Uropods never operculitorm;over pleopods) 0 .)5.00. 21st ce eee 8
8. Uropods lateral or ventrolateral, forming tailfan with pleotelson;
pleopods 1 and 2 rarely operculiform <......... 0. 6c. on Flabellifera ¢ !\\ Uropods terminal or subterminal; pleopods | and 2 variously aperculilorian specu sna gttie asses eso ce ei eee CeCe eae Asellota 9+ ’
* 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 Asellota) 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 * ANTHURIDAE
Key to families of Anthuridea
1. Mouthparts adapted for piercing and sucking, together forming conclike-structire! ..29 744 sv i beeetere as. Paranthuridae p 64 Mouthparts adapted for cutting, lamina dentata and molar usually
presemty:. 0. casktenes Hace ere: cnet te cn salsa ote 2 2. Pereopod | subchelate, with propodus expanded; pleonites generally fused; if free; much: shorter than wide ©. .... ..\0.2 284 20a Anthuridae J Pereopods 1-3 subchelate, subsimilar; pleonites free, often as long as WIE IA EL TAIL ae rte coe ens SRE Grate ata eae a Oaee ror Hyssuridae pco
Family Anthuridae Leach, 1814
DIAGNOSIS Mouthparts adapted for cutting. Pereopod | usually markedly different from remaining pereopods, subchelate with propodus more or less inflated. Exopod of pleopod | operculiform, covering remaining pleopods. Pleonites 1—5 fused, with fusion marked ventrolaterally 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 Pleoped I, only exopod operculiform, 2... 2/5. 605+ vata -s---%-=5 6
2. Antennal peduncle bearing serrate process ............-- Licranthura +3 Antennal peduncle lacking any serrate process ..............-.+-- 3
3.. Pleopod Jim: Y rami to Some depree fused 2... 36. Eisothistos 32 Pleopod: iin; 2; ram. trees") - 5.04 seb om se cee ee eee 4
4. Pereopods | and 2 subchelate, of similar size, propodi not noticeably inflated (12 24! < ooeigads Saba 5 aglaw Sete ake eee) «Site 5 Pereopod | much larger and propodus more expanded, than pereopod 2 s2...ssigekite wadeanst anh eee ernie ee Minyanthura
Amakusanthura 17
5. Integument noticeably pitted; mandible on one side lacking molar, on
Other side wath spleate Molar TH. Re en ek Apanthuroides p- ab Integument not pitted; mandibles similar on both sides Chalixanthura_ p- 2F Be Melson awathissiale/SEALGEVSE (oe ep a am ay bine oe wt ¥en spe ens Anthomuda Telson with two statocysts, or lacking statocysts ................. i 7+ Pereopods 4—7; carpi roughly rectangular 2... .:. ..... ..)R¥ekas Jed 8 Pereopods 4—7, carpi triangular, with anterior margin considerably Shorter than: pOstenior Margihy 205 acd ase ariewid des 1 Wasorw were 9 3: Maxallipedal palpof tworarticless: “220 ys Pali. css Haliophasma Masallipedal palp*of three articles 2... 222 2.0.05%5 sic. s. Malacanthura 9. Maxillipedal palp of one article; mandibular palp of one article BR pe RNC cle S aeRO PANEL dears GIES efeal kay Gee cts Gores aia SP Pendanthurag. sc Maxillipedal palp of more than one article; mandibular palp of more HINARI CONG ATERELE? (os oh osc sb NEI te ela capn Mes Mp2 ear a tered a Ry ee aerate 10 LO: ¢Miasallipedal palpiof two articles \...4.: 2 sls ds25 oh see oe dete ee oe 1] Maxllipedalspalprot three articles... 2.2442 <slc ss cane dees oe « 12 iesMandibularpalp of two,articles,. .. 2... 220 Mataania hones Cortezura P- 24 Mancibular palprot threearucles: . (v6). 2b. ana son ois sh «as Cyathura f. 3] 12. Cephalon with midventral process at base of mouthparts .. Skuphonura Cephalon lacking midventral process')!.26°.4.0:0).g0ditcue half 6 i 13. Pleon lacking dorsal grooves or lines indicating boundaries of fused pleonites; species-specific persistent dorsal pigment pattern usually PUES IU eect hs erent Heh hers iectrocte ect the arts evap any cdl osene -adbvatone Mesanthura Pleon with dorsal grooves or lines indicating boundaries of fused pleonites; persistent pigment pattern lacking ................. 14 14. Pleon with complete dorsal lines separating pleonites Da oie cane eects eta Sita cc Nar eT ae slee oe edi ee, SaaS Apanthura po Pleon with incomplete dorsal line between pleonites 4 and 5 PP Se eee EN tN Gt aE bette ocak epee ae ate Amakusanthura ,\*°'*
Amakusanthura Nunomura, 1977
DIAGNOSIS Integument sometimes with pigment. Eyes present. Antennular flagellum of three articles. Antennal flagellum of two to four articles. Man- dibular palp of three articles, terminal articles bearing distal spines. Max-
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 | 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 REE Ae Mere metre eT Sk FAO HUM VE End ie CEN magnifica Telson dorsally flat, not thickened; uropodal exopod distinctly notched on barely excavate distallys pjis sss ots oe ao one my ere te 2
2. Integument pigmented; uropodal endopod length 1.5 times basal width
sinat idl dfn Reka Betas ee CSU Gs ENE VA SEUSS a SE ace tL MSIE RMN Rear Re eaae a signata
Imtesument not pigmented)... inside sedlk eip atl eieh etd ree 3
3: Uropodal) exopod! distimetly notched... sic oc \in. - eae et Oe. eee 4 Uropodal exopod, outer margin weakly excavate; mandibular palp
anticle:J*bearing three spines mess aaselt cielo icin) eect gee lathridia
4. Mandibular palp article 3 bearing three spines .............. significa
Mandibular palp article 3 bearing two spines ............-. geminsula
Amakusanthura geminsula (Kensley, 1982) Figure 3A—E
DIAGNOSIS. 2: 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. d: 4.8 mm. Eyes larger than ?. Antennular flagellum of 8—9 articles. Pereopod 1, carpus with distal lobe
Amakusanthura geminsula 19
Figure 3. Amakusanthura geminsula: A, 2; B, pereopod 1; C, telson; D, uropodal sympod and endopod; £, uropodal exopod. Amakusanthura lathridia: F, pereopod 1; G, telson; H, uropodal exopod; J, 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 * ANTHURIDAE
Amakusanthura lathridia (Wagele, 1982) Figure 3F—I
DIAGNOSIS @ 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 2: 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
Figure 4. Amakusanthura magnifica: A, pereopod 1; B, telson; C, uropodal sympod and endopod (setae omitted); D, uropodal
exopod (setae omitted)
Amakusanthura signata 21
Figure 5. Amakusanthura magnifica: A, pleonites in lateral view; B, pleon in dorsal
view.
away posteriorly. d: 10.3 mm. Antennular flagellum of 24 articles. Pereopod 1, carpus with sclerotized distal rounded lobe; propodal palm with strong sclerotized tubercle longer than in 2; numerous setae on mesial surface. Pleopod 2, copulatory stylet of endopod not reaching beyond ramus.
RECORDS Off Georgia, 17-137 m; off Florida, 7-11 m; Cuba; Gulf of Mexico.
Amakusanthura signata (Menzies and Glynn, 1968) Figure 6A—E
DIAGNOSIS @: 4.9 mm. Integument with strong patches of pigment on cephalon; pigment sparse on pereonites; two bars on pleonite 6. Antennular
22 ANTHURIDEA * ANTHURIDAE
Figure 6. Amakusanthura signata: A, 2; B, telson; C, pereopod 1; D, uropodal sympod and endopod; £, uropodal exopod. Amakusanthura significa: F, telson; G, uropodal sympod and endopod; H, uropodal exopod; /, 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. ¢: 4.5 mm. Eyes larger than in ¢. 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
DIAGNOSIS @ 5.0 mm. Antennular flagellum of three articles. Antennal 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 | 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 @ 8.8 mm. Antennular flagellum of three or four articles. An- tennal flagellum of six articles. Pereopods | 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, 2°. Apanthura cracenta: B, 2; C, telson; D, uropodal endopod and sympod; £, 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 cracenta 25
Apanthura Stebbing, 1900
DIAGNOSIS Integument sometimes pigmented. Eyes present. 2: 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. ¢ 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
i, Uropodal'exopod distally notched or‘excavate ...... 2. feos ee. ees 2 Uropodal exopod distally entire; uropodal endopod length subequal to WIE D oa Cie ee Fe Ae gE Se ee Ae ieda Mintel act crucis
2. Uropodal exopod distally notched; uropodal endopod length 2.4 times ate tuyaniel igs os Ns Sess Spal Secgced SS atecun agai Sepe raee, iS sBe a ntte cracenta
Uropodal exopod faintly excavate or sinuate; uropodal endopod length abouimiavmice Hasaliwad the sa.1.21 aca 2. Sc cacucray Pods sigra harringtoniensis
Apanthura cracenta Kensley, 1984 Figure 7B—E
DIAGNOSIS @: 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 2!/3 basal length. Pleotelson lanceolate. d: 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 ANTHURIDEA * ANTHURIDAE
RECORDS ‘Turks and Caicos Islands, 1 m; Carrie Bow Cay, Belize, interti- dal on reef crest, 2 m.
Apanthura crucis (Barnard, 1925) Figure 7F,G
DIAGNOSIS ?: 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. ¢: 6.2 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 @ 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 Qi,
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 @: 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. d: 5.0 mm. Integumental pigment stronger than in @, in irregular brown patches. Eyes larger than in 2. 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 6, resulting in mouthpart reduction. Mandibular palp of three articles; molar small, lamina dentata, and incisor present. Mouthparts reduced in 6. Per- eopods 1-3 similar, propodi barely inflated. Pereopods 4—7, carpi triangular. Pleopod 1, exopod broader than endopod, operculiform, or both rami form- ing operculum. Pleopods 2—5, exopods biarticulate. Pleonites short, free, longer in d than in 2. Uropodal rami, margins moderately to strongly in- cised or serrate. Telsonic margin serrate posteriorly; statocysts lacking.
Key to species of Chalixanthura
1. Body pigmented; uropodal exopod deeply incised ........... scopulosa Body not pigmented; uropodal exopod weakly serrate .......... lewisi
Figure 8. Apanthuroides millae: A, 2; B, right mandible; C, left mandible; D, maxilliped; £, uropodal exopod; F, pleopod 1; G, pereopod 1.
Cortezura 29
Chalixanthura lewisi Kensley and Snelgrove, 1987 Figure 9A—D
DIAGNOSIS @: 3.1 mm. Integument lacking pigment. Antennular flagellum of three articles. Antennal flagellum of seven articles. Maxillipedal endite large, apically acute. Pereopod | propodus slightly expanded, palm with few (3) simple setae. Pereopod 7 propodus with two elongate anterodistal fringed spines. Uropodal and telsonic margins serrate. d: 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 @.
RECORDS Barbados, in Madracis mirabilis coral, 9-15 m.
Chalixanthura scopulosa Kensley, 1984 Figure 9E—J
DIAGNOSIS ¢: 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. d: 2.6 mm. Eyes considerably larger and with more ommatidia than in 2. 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, 36; B, 2; C, uropodal exopod; D, pleopod 1. Chalixanthura scopulosa: E, 3; F, 2; G, maxilliped; H, pereopod 1; J, pleopod 1; J, uropodal exopod.
Cyathura (Cyathura) ai
Cortezura confixa (Kensley, 1978) Figure 10
DIAGNOSIS 2 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 Cubagua 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 2. 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 SES Sts Dade A Le BR oe Seth. wen Peis eae ieee Stygocyathura Pleonite 6 dorsally demarked from telson; articulation of uropodal exopod relatively elongate; exopod adpressed dorsally to telson ies ebb te mntin see NN aie tes ANE alata oe Sate eral, HARMS Cyathura
Cyathura (Cyathura) Norman and Stebbing, 1886
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 | removed); C, antennule; D, pleon, lateral view; £, 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 2: 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. d: 5.5 mm. Antennular flagellum of four 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).
34 ANTHURIDEA * ANTHURIDAE
J
Figure 11. Cyathura (Cyathura) cubana: A, 2; B, pereopod 1, 2. Cyathura (Stygocyathura) cuborientalis: C, pereopod 1, 2; D, telson. Cyathura (Stygocyathura) curassavica: EF, pereopod 1, 2; F, telson. Cyathura (Stygocyathura) hummelincki: G,
pereopod 1, @; H, telson. Cyathura (Stygocyathura) motasi: I, telson; J, pereopod 1, ae
Cyathura (Stygocyathura) cuborientalis Botosaneanu and Stock, 1982 Figure 11C,D
DIAGNOSIS 2 6.8 mm. Pereopod 1, propodal palm straight, bearing about
Cyathura (Stygocyathura) hummelincki 215)
TABLE |. CARIBBEAN SPECIES OF Cyathura (Stygocyathura), THEIR TOTAL LENGTHS (MM) AND LOCALITIES
C. (S.) cuborientalis 2 6.8 Cuba
C. (S.) curassavica & 7.0; 292 Curacao
C. (S.) hummelincki 6 4.75.09 8:5 Aruba
C. (S.) motasi 6 6:85 9100 Haiti
C. (S.) orghidani 28:0 Cuba
C. (S.) parapotamica ovis. 931692401 Jamaica
C. (S.) salpiscinalis 6 5:6; Lares Haiti
C. (S.) sbordonii 6, 2 90 Vera Cruz, Mexico C. (S.) specus 6) 18.0; 2 19:8 Cuba
C. (S.) univam 2 10.0 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, 2? 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 Curacao, from pits and wells.
Cyathura (Stygocyathura) hummelincki Botosaneanu and Stock, 1982 Figure 11G,H
DIAGNOSIS ¢ 4.75 mm, ? 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.
36 ANTHURIDEA * ANTHURIDAE
Figure 12. Cyathura (Stygocyathura) curassavica:
pleon.
Cyathura (Stygocyathura) motasi Botosaneanu and Stock, 1982 Figure 111,J
DIAGNOSIS <6 6.8mm, 2 10.0 mm. Pereopod 1, propodal palm gently sinu- ate, bearing 11—18 marginal pectinate setae. Pleotelsonic margins tapering evenly to slightly notched apex; angle of apex less than 90°.
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.
Cyathura (Stygocyathura) parapotamica Botosaneanu and Stock, 1982 Figure 13D,E
DIAGNOSIS 2 4.1 mm (ovig. ? 3.6 mm). Pereopod 1, propodal palm sinu- ate, bearing eight marginal pectinate setae. Pleotelson gently tapering to notched apex; angle of apex about 90°.
RECORDS Jamaica, from river alluvia.
Figure 13. Cyathura (Stygocyathura) orghidani (from Negoescu, 1983): A, telson; B, pereopod 1, 2. Cyathura (Stygocyathura) sbordonii: C, pereopod 1, ° (from Argano, 1971). Cyathura (Stygocyathura) parapotamica: D, telson; E, pereopod 1, ?. Cyathura (Stygocyathura) salpicinalis: F, pereopod 1, 2; G, telson. Cyathura (Stygocyathura) specus: H, 2; I, telson and uropods; J, pereopod 1, 2. Cyathura (Stygocyathura) univam: K, telson.
38 ANTHURIDEA * ANTHURIDAE
Cyathura (Stygocyathura) salpiscinalis Botosaneanu and Stock, 1982 Figure 13F,G
DIAGNOSIS 6 5.6 mm, 2 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 2 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, 2 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 @ 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 ¢ 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 2, rami fused, together forming operculum; d rami separate. Pleonites free, short, longer in d than in 2. Telson lacking statocysts.
REMARKS Wagele (1979) first recorded species of Evzsothistos 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. @ telson with middorsal spines; d pereopod | propodal palm with 19 Gi AURSOUES tee nie eile Suagece ane 2 ke ooo 1s eeu Gm ata aN teri
2 telson lacking middorsal spines; d pereopod | propodal palm with MRS SIOIINE SR ls Chia os spoc sens Meat: «she ieccusen a, eee ie ras oo ale anegey ae petrensis
Eisothistos petrensis Kensley, 1984 Figure 14A—E
DIAGNOSIS @: 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 | rami fused for '/12 of length. d: 2.0 mm. Antennular flagellum of eight articles. Antennal flagellum of six articles. Per- eopod | propodal palm with 11 fringed spines. Telson narrower than in &, 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 @: 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 | rami fused for */+ of length. Uropodal exopod with one or two strong slightly recurved spines on dorsal surface.
40
ANTHURIDEA * ANTHURIDAE
i
: SYK Bee:
Figure 14. Eisothistos petrensis: A, 2; B, uropodal sympod and endopod; C, uropodal exopod; D, pleopod 1; E, pereopod 1, ¢. Eisothistos teri: F, telson and
uropod; G, pleopod 1; H, pereopod 1, d.
Telson with eight or nine slightly recurved middorsal teeth becoming longer posteriorly. d: 2.0 mm. Antennular flagellum of eight articles. Antennal
Haliophasma curri 4]
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 | exopod operculiform. Pleonites 1— 5 short, fused; pleonite 6 usually demarked from telson. Latter with two basal statocysts. d often with more elongate form than ?. Antennular flagellum multiarticulate. Eyes larger.
Key to species of Haliophasma
1. Telson posteriorly narrowly rounded; dactylus of pereopod | dentate sige MN NE AE ey Sees ence Dene Cara anes ey Par eee we pee valeriae
Haliophasma curri Paul and Menzies, 1971 Figure 15A—C
DIAGNOSIS @ 7.0 mm. Antennular flagellum of three articles. Antennal flagellum of five articles. Mandibular palp article 3 with five spines. Max- 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.
42 ANTHURIDEA ¢ ANTHURIDAE
Figure 15. Haliophasma curri: A, 2; B, pereopod 1; C, uropod. Haliophasma valeriae: D, 2; E, pereopod 1; F, uropod.
Haliophasma valeriae Paul and Menzies, 1971 Figure 15D-F
DIAGNOSIS 2 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.
RECORDS Off Venezuela, 95 m.
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 @ 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
DIAGNOSIS. Eyes present. Mandibular palp of three articles. Maxillipedal palp of three articles, terminal article usually broadly ovate. Pereopod 1, propodus expanded. Pereopods 4—7 with carpi roughly rectangular. Pleopod
44 ANTHURIDEA * ANTHURIDAE
Figure 16. Licranthura amyle: A, 2; 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.
Mesanthura 45
Malacanthura caribbica Paul and Menzies, 1971 Figure 17
DIAGNOSIS @ 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 @: 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. d: Eyes larger than in 2. 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 2, extending onto ventral surface.
REMARKS Méesanthura 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, 2; B, °, lateral view; C, mandible; D, maxilliped; £, uropod; F, pleopod 1.
Mesanthura bivittata 47
Key to species of Mesanthura (° only)
1. Pigment in tiny evenly scattered chromatophores over body; cephalon with solid patch of pigment; mandibular palp article 3 with seven SOUT COMI. co UMMeR a ceo His Hees og eee tn cst ot PRE cS punctillata
Pismentinot evenly scattered over body = ..q.cn2... 40 See oes See 2s Z
2. Pereonites 4 and 5 with patch of pigment, sometimes with unpigmented areammommiccdle Of pateliterc.. 0 <=/. sgt ee etce «shape apie s 2 sm ose Skt 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
STAG SPLINES gee sehoee ae ete, Ee. o's =, ae does Wats Sak tome eer Se paucidens Pereonites 4 and 5 with unpigmented area in pigment patch; pleon lackingjtransverse pigmentdines ...)9. «sens. os Species went hea 4
4. Pigment in obvious double longitudinal bands on pereon and pleon; mandibular palp article 3 with nine spines ................ bivittata Picmentnotin obvious double bands. . 2 njcui-l2 soc nste ees Bee ey oo D
5. Unpigmented area in middle of pigment patch of pereonites 1—3;
mandibular palp article 3 with 10 spmes: 4......... 55. sa... .- pulchra No unpigmented area in middle of pigment of pereonites 1—3; mandibular palp article S"with eight spines”... 2.22242...» ©- looensis
6. Pigment of pereon in fine reticulate lines; mandibular palp article 3 WAGE SIXUS[ONMIES® </o).<cpate NMC a 10 ve =.+ « ahtnee name» 6 = ees Aeon Sete reticulata Pigment of pereon not in fine reticulate lines; mandibular palp article 3 SAL PlaN TOU SUING Ses Se cotedar hates = <> + het nears) cis, MaaPe ae arae oye 95, wv eg @ 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 2? 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 hopkinsi; D,
BI
Mesanthura bivittata; B, Mesanthura fasciata; C
3
Figure 18. A
Mesanthura looensis.
Mesanthura fasciata 49
e wi) qo rd
ed ene
Se wes vee
:
Figure 19. A, Mesanthura paucidens; B, Mesanthura pulchra; C, Mesanthura punctillata; D, Mesanthura reticulata.
Mesanthura fasciata Kensley, 1982 Figures 18B, 20E—H
DIAGNOSIS @ 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; B, 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 ¢ 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, 59°m:
Mesanthura looensis Kensley and Schotte, 1987 Figures 18D, 21A—D
DIAGNOSIS 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, | m.
Mesanthura paucidens Menzies and Glynn, 1968 Figures 19A, 21E-I
DIAGNOSIS @: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 | propodal palm with step. d: 6.4 mm. Antennular flagellum of seven articles. Pigment more diffuse than in 2 but retaining five pleonal bars.
RECORDS Looe Key, Florida, 5—6 m; Carrie Bow Cay, Belize, 15.2 m; Pu- erto Rico, intertidal; Jamaica.
52 ANTHURIDEA * ANTHURIDAE
o
Figure 21. Mesanthura looensis: A, mandible; B, maxilliped; C, pereopod 1; D, uropodal exopod. Mesanthura paucidens: E, mandible; F, maxilliped; G, pereopod 1, 2; H, pereopod 1, 6; J, uropodal exopod. Mesanthura pulchra: J, mandible; K, maxilliped; L, pereopod 1, d; M, pereopod 1, 2; N, uropodal exopod.
Mesanthura pulchra Barnard, 1925 Figures 19B, 21J—N
DIAGNOSIS 2: 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. d: 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, | m; Puerto Rico, intertidal 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. floridensis.
Mesanthura punctillata Kensley, 1982 Figures 19C, 22A—F
DIAGNOSIS 2: 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 | propodal palm with step. d: 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 { 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 @: 1.7 mm. Antennular flagellum of one article. Antennal
Figure 22. Mesanthura punctillata: A, mandible, 2; B, mandible 6; C, maxilliped; D, pereopod 1, 2; E, pereopod 1, 6; F, uropodal exopod. Mesanthura reticulata: G, mandible; H, maxilliped; J, pereopod 1; J, uropodal exopod.
Minyanthura corallicola 55
Figure 23. Minyanthura corallicola: A, 2; 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. d: 1.3 mm. Eyes larger than
in 2. Antennular flagellum of three articles. Antennal flagellum of two articles.
RECORDS Carrie Bow Cay, Belize, 6-24 m; Barbados, 9-15 m; Jamaica.
56 ANTHURIDEA * ANTHURIDAE
Pendanthura Menzies and Glynn, 1968
DIAGNOSIS 2: 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. Maxillipedal 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. d: 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 @.
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 bey 2 nak seeps Sees mot ee eee tanaiformis Dorsal pigmentation on cephalon, pereonite 2, and pleon; pereopod 1,
propedal palmulacking rounded lobe fonts = 2 se hendleri
Pendanthura hendleri Kensley, 1984 Figure 24A—E
DIAGNOSIS 2: 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. d: 2.8 mm. Pigmentation as in ?. Pereopod 1, propodal palm gently convex, lacking rounded lobe.
RECORDS Carrie Bow Cay, Belize, 9-23 m; Twin Cays, Belize, 0Q—2 m; Pan- ama, 30 m.
Pendanthura tanaiformis Menzies and Glynn, 1968 Figure 24F-H
DIAGNOSIS Ovigerous 2: 2.9 mm. Dorsal pigmentation a dense red-brown
Figure 24. Pendanthura hendleri: A, 2; B, mandible; C, maxilliped; D, pereopod 1, 3; E, pereopod 1, 2. Pendanthura tanaiformis: F, 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. d: 2.8 mm. Pigmentation as in 2. 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 | similar to
Figure 25. Skuphonura laticeps: A, 3; B, cephalon and pereonite 1, lateral view; C, antennule, d; D, pereopod 1, d; £, antenna; F, maxilliped.
60 ANTHURIDEA * HYSSURIDAE
following pleopods, not operculiform. Pleonites 1—5 freely articulating, rela- tively elongate. Pleotelson lacking statocysts.
Key to genera of Hyssuridae
1. Pleotelson not covered by uropodal exopods; maxillipedal rami basally EEE) a. Son SE sists aS he ed ES ey, Send ol SPs Ae Sree ae Kupellonura
Pleotelson covered completely by uropodal exopods; maxillipedal rami basally fused) ic) c-f5.1.) errs cose oar te ea ae cess ea tor earns 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 @: 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. d: 2.7 mm. Eyes larger than in 2. 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, | 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, 2°; B, cephalon and pereonite 1, 3; C, maxilliped;
D, pereopod 1, ; E, pereopod 1, 3.
62 ANTHURIDEA * HYSSURIDAE
Figure 27. Kupellonura imswe: A, 2 cephalon, dorsal view; B, posterior pleon, dorsal view; C, pleon, ventral view; D, cephalon and pereonites | and 2, lateral view.
Xenanthura brevitelson Barnard, 1925 Figure 28
DIAGNOSIS 2: 4.0 mm. Ommatidia of eyes in longitudinal row of three or four groups. Antennular flagellum of three articles. Antennal flagellum of three articles. Mandibular palp of single short article bearing single seta. Maxillipedal rami basally fused, palp of three or four obscurely separated articles. Pereopods 1—3 similar, carpus of pereopod 2 triangular, with strong triangular projecting lobe posterodistally; propodi expanded, palm incised into several rounded lobes. Uropodal exopods circular, overlapping dorsally and covering telson; endopod projecting beyond exopods ventrally, mesial margin with step, distally rounded. Telson shorter than uropodal exopod, posterior margin truncate to faintly concave. 6: 3.5 mm. Antennular flagellum of seven articles.
RECORDS Off Georgia, 20-145 m; off Florida, 8-10 m; Turks and Caicos Islands, 1 m; St.Thomas, U.S. Virgin Islands, 50-60 m; Gulf of Mexico.
Figure 28. Xenanthura brevitelson: A, @; B, pereopod 1; C, uropodal endopod; D,
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 fused. Pleotelson with single basal statocyst, or lacking statocyst.
Key to genera of Paranthuridae
1. Pereopod»7 lacking in‘adult. s/o. 92) sashes sao dee-.> so ee eee 2 Pereopod. 7 present inadult: 2. a. %ielas om gees ole sd tok cates oe eee 3 2: EVES (PLESEME (6.05. jn Sees. Sao, seeks cobs tonlere sale Sie eins etd Ss < fete Colanthura Byes absent. is... ci eee Seem mL ans oo Ee Curassanthura
3. Antennular and antennal flagellum of more than 10 anticles: 37.5 ae: dae ee eae os Shee Meee Misia eee eee Accalathura
4. Antennal flagellum a single (rarely two or 3) flattened article;
maxilhpedal palp of onevor twetartcles’ 219. ..210025 os Paranthura Antennal flagellum of seven articles; maxillipedal palp of three articles eR Te ere ate Lerner 2. Sa GOs RRS OS cee ks Virganthura
Accalathura Barnard, 1925
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 widthy (iho este fac siecnedenn eee eh Nera Chane ene tate ketene setosa
Colanthura tenuis 65
palp of two articles, endite almost reaching end of palp. Pereopod | 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 2: 16.0 mm. Antennular flagellum of about 26 articles. Anten- nal flagellum of about 18 articles. Uropodal exopod narrow, parallel sided. Telson apically subacute. d: 15.0 mm. Pleopod 2, copulatory stylet of endo- pod apically acute, with subapical “heel.”
RECORDS Off North 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 2: 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. d: 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 : 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, 2; B, uropodal sympod and endopod; C, uropodal exopod; D, telson. Accalathura setosa: E, 2; 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. ¢: 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 | 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) 49 umes ereatest width. 25). of ac2e scr o3 gas bermudensis Telson tapering, but posterior fourth not abruptly narrowed; uropodal exopod! lengih 2-amimies ereatest Widths. ~.. <--ee - +25. --4-- halma
Curassanthura bermudensis Wagele, 1985 Figure 30G,H
DIAGNOSIS @ 3.0 mm. Pereopod | 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, 2; B, uropod; C, pereopod 1. Curassanthura halma: D, @; E, telson; F, uropodal exopod. Curassanthura bermudensis: G, telson; H, uropodal exopod (from Wagele, 1985).
Curassanthura halma Kensley, 1981 Figure 30D-F
DIAGNOSIS @ 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 SeMEALe ) Pee Se OMe ae ER on Sh oe Re Side ice infundibulata Telson posteriorly rounded; uropodal exopox ovate, margins entire .. 2
Za wropodaltendoped longer than wide ...5.¢.,. 445-00: 6 s-e206-% +s fos 3 Wropodal-endopad as lone as wide... ..4. digs. s wscace cise antillensis
3. Uropodal exopod elongate-elliptical; telson parallel sided for half length
Uropodal exopod ovate, outer margin sinuate; telson evenly elliptical SRS rite il eres Sete akon ek eS ahh Op eae bidnt ga rece A a barnardi
Paranthura antillensis Barnard, 1925 Figure 31A—F
DIAGNOSIS 2 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: A, 2; B, telson; C, uropodal exopod; D, uropodal sympod and endopod; E, pereopod 1, 2; F, pereopod 2, 2. Paranthura barnardi: G, 2; H, telson; J, pereopod 1, 2; J, 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 @ 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.
RECORDS Off Venezuela, 95 m.
Paranthura floridensis Menzies and Kruczynski, 1983 Figure 32A—E
DIAGNOSIS @ 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 Off Sanibel Island, Florida, Gulf of Mexico, 73 m.
Paranthura infundibulata Richardson, 1902 Figure 32F-J
DIAGNOSIS @ 8.2 mm, 6 6.0 mm. Integument with red-brown pigmenta- tion; broad irregular patch between eyes running onto bases of antennules; pereonites | 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 floridensis: A, 2; B, pereopod 1, 2; C, uropodal sympod and endopod; D, uropodal exopod; E, telson. Paranthura infundibulata: F, 2; G, telson; H, pereopod 1, 2; J, uropodal exopod; J, uropodal sympod and endopod.
ASELLOTA ns
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
DIAGNOSIS’ Eyes present. Mandibular palp of three articles. Maxillipedal palp of three articles; endite present. Pereopods 1—3 subchelate, pereopod | 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
DIAGNOSIS ? 6.8 mm. Antennular flagellum of three articles. Antennal 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 USS. 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 2. One pair of pleopods in @, and one or two pairs of pleopods in 6 forming operculum over remaining respiratory pleopods. Pleopod 2 in 3 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,
Figure 33. Virganthura crassa: A, 2; B, maxilliped; C, uropodal endopod and sympod; D, telson; E, uropodal exopod.
Atlantasellus cavernicolus 15
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 <d: Pleopod 1, peduncles separate, rami uniarticulate; pleopod 2 having biarticulate exopod and copulatory endopod; pleopod 3 biramous, opercular. 2: 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. d: Pleopod 1 of two articles; pleopod 2 with sympod, small exopod, and uniarticulate copulatory endopod. °: Pleopods | 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.
“eIOT[PSY Jopsoqgns ay} Jo sarpruueysadns inoy ay} ut ¢—] spodogyd jo uostreduiory “FE aunB1q
\: U O ¢ GOd03 1d SY poo
| GOd03 1d
Gnathostenetroides pugio CE
Superfamily Gnathostenetroidoidea Kussakin, 1967
DIAGNOSIS 4d: Pleopod | peduncles fused, uniarticulate rami separate, op- ercular; pleopod 2 small, with biarticulate exopod and copulatory endopod; pleopod 3, broad endopod biarticulate, exopod slender, uniarticulate. 2: Pleopod 1 absent; pleopod 2 rami fused to form operculum; pleopod 3 as in
OF
Family Gnathostenetroididae Kussakin, 1967
DIAGNOSIS Uropods with short sympod, rami relatively well developed. 3: 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. 2: 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 d, larger than following pereopods. Pleon having one very short free pleonite. Pleotelson with lateral margins subparallel, ending in strong tooth posteriorly; posterolateral margins sinuous, rounded. Mand- ible in d with strong projection arising proximolateral to incisor, visible dor- sally; in 2, projection short, not visible dorsally.
Gnathostenetroides pugio Hooker, 1985 Figure 36A—C
DIAGNOSIS 6 3.2 mm, @ 1.6 mm. Eyes of five ommatidia each. d: Per- eopod 1, merus with anterior margin strongly produced into narrow triangu- lar process; propodus broad, palm demarked by strong denticulate spine and
78 ASELLOTA ¢ GNATHOSTENETROIDIDAE
Figure 35. Atlantasellus cavernicolus: A, 2; B, pereopod |. (From Sket, 1979).
bearing row of nine more slender spines, posterior margin setose. 2: 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 2 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 1.46mm, 2 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, 36; B, mandible; C, pereopod 1. Neostenetroides stocki (from Carpenter and Magniez, 1982): D, 2; 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 | and 2 together forming operculum; pleopod 3 endopod uniarticu-
80 ASELLOTA * JANIROIDEA
late, exopod biarticulate. 2: Pleopod 1 absent; pleopod 2, rami fused to form operculum; pleopod 3 as in 6.
Key to families of Janiroidea
1: Eyes (Gf present):on lateral processes of céphalon .. ..17....-. 42.025 2 Eyes dorsolateral on cephalon, not on lateral processes ............. 6 2. Uropods with large, easily visible sympod and rami ............... 3 Uropods with sympod minute or absent, rami short ............... 5
3. Pleon posteriorly markedly produced; some pereonites produced
laterally into, fingerlike:processes: {27.2.2 Jew t ees Pleurocopidae Pleon posteriorly rounded, barely produced; pereonites not laterally
produced into prominent fingerlike processes: .....-5.5.---2- ease 4
4: \Peréopod' lSubchelate “sth 5 gas Oc as sete en ee Santiidae
Pereopod. ] ambulatory, biunguiculate 52% os -s.6 2. .tiee ee Mexicope
5. Pleopod 1 6 distally sagittate; anus covered by pleopod 1 d, or
2 ber) 008 ee ee aera kta, ety Se Rr Paramunnidae
Pleopod | 6 distally truncate; anus exposed .............. Munnidae
6. Uropodal rami minute, smaller than squat sympod ...... Joeropsidae
Uropodal rami elongate, sympod variable, generally elongate ....... 7
7. Eyes lacking; pereopods all similar, ambulatory .... Microparasellidae Eyes usually present; pereopod | prehensile, subchelate, pereopods
2= 7am bUlAaLOry , sch hae ven Puateeee sae ee see Re Janiridae
Family Incertae Sedis
Mexicope Hooker, 1985
DIAGNOSIS_ Eyes on short lateral lobes of cephalon. Scale present on anten- nal peduncle. Coxae short, visible in dorsal view on pereonites 2—7. Per- eonites laterally acute, not markedly produced. Pereopod | ambulatory, dac- tylus biunguiculate. Pleon consisting of one short pleonite plus broad pleotelson. Uropods large, with elongate peduncle, endopod, and exopod.
ASELLOTA * JANIRIDAE 81
Figure 37. Mexicope kensleyi: A, 2; B, pereopod 1.
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, 2 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. Mandibles with palp and well-developed molar. Maxilliped with articles 1—3 at least as wide as endite, markedly broader than articles 4 and 5. Pereopod | prehensile, subchelate, sexually dimorphic, larger in ¢ than in 2. Pereopods 2—7 ambulatory, dac- 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 ¢)
L..,Propodisdistallly, Acute stoic has Rahn et Says si SS see as, seo 2 Propodusydistally broads ooo ike oe scccs ois Sut a ona ans on 4 2. Propodus apically acute, with proximal tooth ................ minutus Propodus’ lacking ‘teeth: 38 2'..18. See ane sh. IO RSE eee 3
3. Carpus with two strong distal teeth; propodus (when chela closed) FEACHING. TOUMEHUS.. voles cineca sews a pe ieee eee cae oe algicola Carpus with two strong and one small teeth; propodus (when chela
closed’) reaching proximallhalfof carpus 22-2: +245. 20 ec: serricaudus 4. Propodus with distinct teeth, ots. iach yhoo es > Propodus lacking distinct teeth. otic cre hake ee 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 GME tho ons See t nA sd GAR socio agers Soe soe geen e: punctatus 6. Carpus with two distal teeth; dactylus minute ................. triton Carpus with three distal teeth; dactylus small, but not obsolete ..... |
7. Propodus distally truncate; carpus with middle tooth of palm distally faintly “bilobed 5 eS eer hee eee eee ee brachydactylus Propodus distally faintly bilobed; carpus with middle tooth of palm distally narrowly“rounded OY i2ee2 ene ee ee ae 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 algicola (Miller, 1941) Figure 38A,B
DIAGNOSIS 6 2.9 mm, ovigerous 2 2.0 mm. Frontal margin straight. Per- eopod | in 6, carpus distally not much broader than proximally, with two teeth. Propodus reaching back to merus in mature male. Pleopod | in 4, rami with outer lobe distally acute but not produced. Uropod longer than pleotelson. Pigment in scattered red-brown chromatophores.
RECORDS Looe Key, Florida, 1—1.5 m; Yucatan, Mexico; Carrie Bow Cay, Belize, 0—2 m; Puerto Rico; Jamaica; Venezuela. Indo-west Pacific.
Carpias bermudensis Richardson, 1902 Figure 38C,D
DIAGNOSIS. 6 2.7 mm. Frontal margin straight. Pereopod | in adult ¢ al- most 1.6 times body length; carpus distally broadened, with rounded pos- terior area, palm with tooth at outer distal angle, larger tooth at midlength followed by deep notch; propodus with two teeth on flexor margin, widening to truncate distal margin. Pleopod 1 6, outer distal lobe narrowly acute, somewhat produced. Operculum of 2 with distal margin emarginate.
84 ASELLOTA * JANIRIDAE
RECORDS’ Bermuda; eastern and southern coasts of Florida, 1.5—15 m.
Carpias brachydactylus Pires, 1982 Figure 38E
DIAGNOSIS <6 1.6 mm. Pereopod | 4, 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 | 2, outer distal lobe slightly produced, rounded. Operculum of 2 wider than long, distal margin broadly emarginate.
RECORDS Puerto Rico, 1.5 m.
Carpias harrietae Pires, 1981 Figure 38F,G
DIAGNOSIS. 6 2.3 mm. Closely resembling C. brachydactylus. Pereopod | 6, carpus with posterodistal area somewhat expanded, with strong tooth defin- ing outer margin of palm, middle tooth rounded, inner tooth distally faintly bifid; propodus widening to shallowly bilobed distal margin. Pleopod | 6, inner distal lobe rounded, outer lobe narrowly acute. Operculum of @ 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 6 1.9 mm, ovigerous 2 1.8 mm. Pereopod | ¢, carpus distally widening, palm defined by strong triangular tooth, two inner teeth of palm separated by rounded notch; propodus with strong proximal tooth on flexor margin, distally produced into small triangular lobe below dactylus. Pleopod 1 6, outer distal lobe narrowly triangular and produced well beyond inner lobe. Uropod subequal to pleotelson in length.
RECORDS Bermuda, on Sargassum.
Carpias punctatus 85
Figure 38. Carpias algicola: A, 2; B, pereopod 1, d. Carpias bermudensis: C, D, pereopod 1, 3. Carpias brachydactylus: E, pereopod 1, 3. Carpias harrietae: F, G, pereopod 1, 6.
Carpias punctatus (Kensley, 1984) Figure 39 B,C
DIAGNOSIS 6 2.2mm, @ 2.8 mm. Dorsal integument with patchy reticulate pattern of large dark brown chromatophores. Frontal margin faintly convex.
86 ASELLOTA * JANIRIDAE
Figure 39. Carpias minutus: A, pereopod 1, 6. Carpias punctatus: B, 3; C, pereopod 1, 3. Carpias serricaudus: D, pereopod 1, 3. Carpias triton: E, F, pereopod 1, ¢.
Pereopod | 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 | 6, outer distal lobe narrowly triangular, reaching well beyond inner lobe. Oper- culum of 2 with distal margin shallowly concave. Uropod half length of pleotelson; latter with posterior margin a broadly rounded lobe between uropodal bases.
RECORDS Carrie Bow Cay, Belize, intertidal to 15.2 m.
Joeropsis 87
Carpias serricaudus Menzies and Glynn, 1968 Figure 39D
DIAGNOSIS 6 1.6mm, 2 1.5 mm. Pereopod | 6, 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 4, 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 d 2.3 mm. Very similar to C. algicola. Pereopod | d, carpus with two strong basally broad distal teeth; propodus extending back to merus in adult 6, widening to broadly rounded distal margin; dactylus minute. Pleopod | 6, 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 5 og eR SO een eRe nen octane a Mr ak aL eS enatae te bifasciatus
3. Rostrum evenly convex; outer uropodal ramus longer than inner EE eco Sch eR on eo Pee EE Ee re Noe rathbunae Rostrum anteriorly shallowly notched; outer uropodal ramus shorter tian Grime. setae. sco canes etc n Bn oie Sieve sins Weer enESeeTS coralicola
Joeropsis bifasciatus Kensley, 1984 Figure 40A—F
DIAGNOSIS 6 2.5 mm, @ 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 2 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 Cay, 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 2 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 2 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., 29-33 m.
Figure 40. Joeropsis bifasciatus: A, 6; B, pleopod 1, 3; C, uropod; D, operculum, ; E, antennule; F, antenna. Joeropsis coralicola: G, 3. Joeropsis personatus: H, 3. Joeropsis rathbunae: I, 3.
90 ASELLOTA * MICROPARASELLIDAE
Joeropsis personatus Kensley, 1984 Figure 40H
DIAGNOSIS 6 2.2 mm, @ 2.0 mm. Body glabrous. Lateral margin of cephalon serrate. Rostrum triangular. Lateral margins of pleotelson serrate. Antennal flagellum of five articles. Apex of 2 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 Porites 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 2 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 2 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. 3, pleopod 1 narrow, not overlapping external part of pleopod 2; maxillipedall palp:of five articles’: 2); 34 1)--)-1- -re Microcharon 3, pleopod 1 broad, almost completely covering pleopod 2; maxillipedal palp of four articles, terminal article ending in pointed FUEOGESS. 5c eons hats leh ole atone ee wa. electra cet eee on yet yet nee er te Angliera
Microcharon sabulum 9]
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- nai acute process. Pleopod | in d forming broad lamella.
Angliera psamathus Kensley, 1984 Figure 41A—D
DIAGNOSIS. d6 1.0mm, 2 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 _ ‘T'wo 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. d, pleopod | 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 1.4mm, 2? 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, 3; B, right, and part of left, mandible; C, maxilliped; D, pleopod 1, 3. Microcharon sabulum: E, 3; F, mandible; G, maxilliped; H, pleopod 1, 6.
Munna 93
dopod of pleopod 3 with three distal plumose setae. Uropodal sympod stout, longer than rami.
RECORDS Carrie Bow Cay, 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 | 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 | enormous in 6, as compared LEE OE NTR OS Raat eee One ane ay ON ae He Munna
Munna Kreyer, 1839
DIAGNOSIS Body dorsally with numerous setae and/or articulating spines. Antennular flagellum with two distal articles each with single aesthetasc, terminal article minute. Mandibular molar strong, subcylindrical, distally truncate, with accessory setae; palp reaching beyond incisor, article 2 with few serrate spines. Pereopod | sexually dimorphic; pereopods 2—7 not (or barely) sexually dimorphic, dactyli with accessory claw. Pleopod 3, article 2 of exopod reaching well beyond endopod.
94 ASELLOTA * MUNNIDAE
Munna petronastes Kensley, 1984 Figure 42A—D
DIAGNOSIS d 1.1 mm, 2 1.0 mm. Pereopod | in d enormously enlarged, carpochelate. Pleopod | in d with distolateral angles projecting, acute. Body with anterodorsal U-shaped pigment band, and two converging bands on posterior pereon.
RECORDS Carrie Bow Cay, Belize, intertidal to 2 m, usually on corals.
Uromunna Menzies, 1962c
DIAGNOSIS Body with few if any dorsal setae, without articulating spines. Terminal antennular flagellar article not minute, bearing single aesthetasc. Mandibular molar strong, cylindrical, distally truncate, lacking accessory setae; palp present or absent. Pereopod | not sexually dimorphic, small. Per- eopod 2 (rarely 2—7) sexually dimorphic, carpi and propodi broader in 6 than in 2. Pleopod 3, article 2 of exopod not reaching beyond endopod.
Key to species of Uromunna
1. Larger uropodal ramus parallel sided, about 3.5 times longer than basal width; inner uropodal ramus tiny, obscured by pleotelsonic TELA DORN aS. 5 oaths BP wwe eee a cee ie ie ver ke shoals har ars eet ae reynoldsi Larger uropodal ramus tapering, about 1.5—2.0 times longer than basal width; inner uropodal ramus smaller than outer, but visible beyond
pleotelsonie mare <5 soccer ose ee ee caribea
Uromunna caribea (Carvacho, 1977) Figure 42E,F
DIAGNOSIS 6 1.5 mm, 2 1.5 mm. Propodus of pereopod 1 1.5—2 times longer than wide. Operculum of @ distally truncate. Shorter uropodal ramus visible beyond pleotelsonic margin, with single seta. Pigmentation in re- ticulation on cephalon and pereon; with six marginal patches on pleon.
RECORDS ‘Turks and Caicos Islands, 1 m; Canal de la Belle Plaine, Guadeloupe, in water of 25%o.
Uromunna reynoldsi 95
Figure 42. Munna petronastes: A, 3; B, uropod; C, pereopod 1, 3; D, pereopod | ¢ (C and D same scale). Uromunna caribea: E, 3; F, larger uropodal ramus. Uromunna reynoldsi: G, pereopod 1, 6; H, larger uropodal ramus; J, 6.
Uromunna reynoldsi Frankenberg and Menzies, 1966 Figure 42G—I
DIAGNOSIS <6 1.5 mm, @ 1.6 mm. Propodus of pereopod 1 two or three times longer than wide. Operculum of ¢ 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 | prehensile; pereopods 2—7 ambulatory. Pleopod 1 3 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, 2 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 | in 6 not sagittate. Uropod pedunculate, inserted laterally or slightly dorso- or ven- trolaterally on pleotelson; biramous, or with one ramus fused with sympod.
Pleurocope 97
Figure 43. Munnogonium wilsoni: A, 2; B, pereopod 1, 3. Pleurocope floridensis: C, 3; D, antennule 6; FE, pereopod | 6. Santia milleri: F, 3; G, pereopod 1, 3; H, maxilliped.
Pleurocope Walker, 1901
DIAGNOSIS Eyes present on lateral peduncle. Antennular peduncle of two articles; flagellum of four articles. Antennal peduncle of five (six) articles, scale lacking; flagellum of six or seven articles. Mandibular palp lacking. Pereopod | subchelate. Pereopods 2—7 uniunguiculate.
98 ASELLOTA * SANTIIDAE
Figure 44. Pleurocope floridensis: adult in dorsal
view.
Pleurocope floridensis Hooker, 1985 Figure 43C—E, 44
DIAGNOSIS. 6 1.15 mm, ¢ 0.96 mm. Body ovate, tapering posteriorly. In- tegument very finely tuberculate. Mesiodistal lobe on antennal peduncle ar- ticle 3 bearing five distal setae. Pereon lacking long dorsal setae. Pereopod 1 subchelate, but almost carpochelate. Pleon consisting of single segment, pos- teriorly narrowly tapered and produced. Uropodal rami as long as sympod.
RECORDS Turks and Caicos Islands, 1 m; Carrie Bow Cay, Belize, 3—10 m; Florida Middlegrounds, Gulf of Mexico, 55 m.
Family Santiidae Wilson, 1987
DIAGNOSIS Antennular flagellum with at most, three articles, antennular scale sometimes present. Pereopod 1 prehensile. Pereopods 2—7, dactyli biunguiculate. Coxae visible at least on pereonites 5—7. Pleon consisting of single short pleonite plus pleotelson. Uropods pedunculate, biramous, inser- ted dorsally or laterally. (One species of Santia possesses a uniramous
uropod. )
Santia Sivertsen and Holthuis, 1980
DIAGNOSIS Cephalon about twice wider than long. Eyes present. Antennu- lar peduncle of three articles. Pereonites laterally rounded, sometimes bear- ing short lateral spines.
ASELLOTA ¢ STENETRIIDAE 99
Santia milleri (Menzies and Glynn, 1968) Figure 43F—H
DIAGNOSIS 6 and 2 1.0 mm. Eye on short lateral process of cephalon. Mandibular palp of three articles. Maxillipedal palp articles all of similar width. Pereopod | 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 | small, peduncles fused, rami separate, uniarticu- late; pleopod 2 small, copulatory; pleopod 3 biramous, opercular. 2: Pleopod 1 absent; pleopod 2 rami and peduncle fused to form operculum; pleopod 3
as in oC.
Family Stenetriidae Hansen, 1905a
DIAGNOSIS’. <d: Pereopod | frequently much bigger than in 2, with distinc- tive lobes and teeth. Pleopods 1 and 2 reduced; pleopod | protopodite short, fused, rami separate. Pleopod 2, endopod elongate, flexed, exopod short. Pleopod 3 exopod basally broad, distally narrowed; endopod broad, biarticu- late. 2: 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 24s eat ee ee een Stenobermuda Rostrum short, basally broad, anteriorly truncate or broadly rounded; two or three very short free pleonites anterior to pleotelson ee Se teh Se NCE RG, Naa eA. PUNTERS oa Stenetrium
100 ASELLOTA * STENETRIIDAE
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! ommmatidia, renitorm 2,,.0.103 1, boc). h . ee 3
2. Eyes of four ommatidia; d pereopod 1 small, propodus not unusually
21.01 (6 Mima Lea Oem eR cS gone IN MOH AE Ser RCA a minocule Eyes of more than four ommatidia; 3 pereopod 1, propodus broad,
WHEL Wale soa et sian at oe Tn ee ee Nk patulipalma
3) ‘Pleotelsonie. margins. serrate’. sess. 6 eee ee os eee 4
Fleotelsonic*margins entire We" eh. . 4504 Be ee 5
4. Rostrum convex, with fine marginal teeth; pereopod 1 6, propodal
palne with*three(straight teethe Avs. 2. en iol).. atemne Sauee bowmani Rostrum truncate; pereopod | 3, propodal palm with two teeth, outer
toothelongate; cued) i S20. 20a, deen). Une %, nao serratum
5. Pereopod 1 6, carpus produced, apically acute .............. stebbingi
Pereopod | 6, carpus produced, apically rounded ....... spathulicarpus
Stenetrium bowmani Kensley, 1984 Figure 45
DIAGNOSIS 6 5.0 mm, 2 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. d: Pereopod | propodus broad, palm with three teeth, outermost
Stenetrium bowmani 101
Figure 45. Stenetrium bowmani: A, 6; B, pereopod 1, 3; C, pereopod 1, 2D; pleopod 1, d; E, pleopod 2, 4; F, pleopod 3; G, pleopod 4; H, operculum, &.
102 ASELLOTA ¢ STENETRIIDAE
longest, slender. 2: Pereopod | 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 6 2.8 mm, 2 3.7 mm. Eye of four ommatidia. Anterolateral lobes of cephalon blunt, barely produced. Rostrum poorly defined, truncate. 3: Pereopod 1, carpus produced posterodistally into broadly rounded lobe; propodus broad, palm demarked by strong spine, with six low rounded teeth. ?: Pereopod | 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 6 2.0 mm, @ 2.7 mm. Eyes of about 10 ommatidia in cluster. Rostrum poorly defined, truncate. Two basal articles of maxillipedal palp not enlarged. d: Pereopod | unknown. 2: Pereopod | 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. 6 4.0 mm, ovigerous 4.9 mm. Rostrum truncate. Pereonites 1—5 with acute anterolateral angles. Pleotelsonic lateral margins with five teeth. d: Pereopod 1, propodus broad, palm with three teeth, outermost elongate, curved; dactylus reaching well beyond outermost palmar tooth. 2: Pereopod 1, propodus much smaller than in d, palm bearing series of about nine fringed spines. Color: tiny red-brown chromatophores arranged in re-
A, 3; B, pereopod 1| 4; C, pereopod | 2. Stenetrium Q. Stenetrium serratum: F, 3; G, pereopod | Oe.
Figure 46. Stenetrium minocule: patulipalma: D, 2; E, pereopod 1 pereopod | Gd.
104 ASELLOTA * STENETRIIDAE
ticulate 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, ¢ 4.1 mm. Rostrum truncate. d: 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. 2: 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 ¢ 4.1 mm. Rostrum truncate. Ommatidia of eye more bunched than in S. spathulicarpus. 6: Pereopod | 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. 2: 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.
thulicarpus: A, 3; B, pereopod 1, 3, many setae removed; rium stebbingi: D, 3; E, pereopod 1, 9; F, G, H, variation
Figure 47. Stenetrium spa C, pereopod 1, 2. Stenet in pereopod 1, d.
106 ASELLOTA © STENETRIIDAE
ae.
Figure 48. Stenobermuda acutirostrata: A, 2; B, pereopod 1, °.
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, | m.
EPICARIDEA 107
Suborder Epicaridea Latreille, 1831
DIAGNOSIS’ Predominantly ectoparasites of marine crustaceans, feeding on blood. Eyes sessile, usually present in d, often reduced or lost in 2. Anten- nae and antennules reduced; mouthparts reduced, forming a suctorial cone containing pair of piercing stylets formed from modified mandibles. Maxillae 1 and 2 reduced or lost. All mouthparts may be lost, and replaced by pro- boscis. d small and isopodlike. 2 undergo considerable distortion or reduc- tion, often to unsegmented sacs of eggs in some forms. Ostegites usually re- tained. Two larval mancalike stages, epicaridium and cryptoniscium (Figure 49), characteristic of entire suborder.
REMARKS The epicarideans are ectoparasites of other crustaceans, with the juveniles often using copepods as intermediate hosts. Sexual dimorphism is marked, the males being symmetrical with unambiguous segmentation, and considerably smaller than the often highly distorted females. In these, body segmentation is often obscured, with body segments often expanded on one side and reduced and compressed on the other. The marsupium of the ovigerous female, except in the Cryftoniscidae and Entoniscidae, is made up of broadly lamellar oostegites, is relatively enormous and often obscures the rest of the body structure.
Crustacean hosts of the epicarideans are found in four classes: Ostracoda, Copepoda, Cirripedia, and Malacostraca, and in nine orders of the Mal- acostraca: Leptostraca, Stomatopoda, Mysidacea, Cumacea, Tanaidacea, Isopoda, Amphipoda, Euphausiacea, and Decapoda. The Epicaridea have been divided into two superfamilies, the Bopyroidea, containing families Bopyridae, Dajidae, and Entoniscidae, and the Cryptoniscoidea, containing
the Crytoniscidae. In the Bopyridae (Figure 50), the often asymmetrical adult female shows
some segmentation. Seven pereopods may be present only on one side, their number being variable on the other. This is largest of the epicaridean fam- ilies, containing over 400 species (Markham, 1974). Ten subfamilies have been recognized: the monotypic Entophilinae parasitizes galatheid crabs; six subfamilies, the Argeiinae, Bopyrinae, Bopyrophryxinae, Ioninae, Pseu- dioninae, and Orbioninae are all branchial parasites of decapod crustaceans; two subfamilies are abdominal parasites, the Phyllodurinae on callianassid mud-shrimps, and the Athelginae on hermit crabs; the Hemiarthrinae are known from the dorsal and ventral abdominal surfaces, and from the branchial chamber of caridean shrimps.
The Dajidae are ectoparasites pelagic mysidaceans, euphausiaceans, and decapod caridean shrimps. Adult females are often found unattached in plankton and pelagic samples. When attached, dajids are found on the
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SW hs =a) is Soe > ae OP BASE ON, ey ZS 2 WIAA ALES
IVE
OSS
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Figure 49. A, epicaridium larva, lateral view; B, epicaridium larva, ventral view;
C, cryptoniscium larva, ventral view (from Bonnier, 1900).
EPICARIDEA 109
Figure 50. A, caridean shrimp with bopyrid parasite in branchial chamber. Probopyrus pandalicola: B, 2 and ¢ in dorsal view, same scale; C, d enlarged; D, °, 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)
Synalpheus pectiniger Coutiére North Carolina to Florida; Bahamas; Stenorhynchus seticornis (Herbst) Hispaniola; Jamaica; Bonaire; Curacao; Belize; Gulf of Mexico Balanopleon tortuganus Markham, 1973 Munida simplex Benedict
Tortuga Island
Jamaica
Aporobopyrina anomala 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
Bopyrella harmopleon Bowman, 1956 Synalpheus brevicarpus (Herrick) Synalpheus fritzmuelleri Coutiére Synalpheus hemphilli Coutiére Synalpheus minus Say Venezuela; Brazil
Bopyrina abbreviata Richardson, 1904 Hippolyte curacaoensis Schmitt Hippolyte pleuracanthus (Stimpson) Hippolyte zostericola (Smith)
Argeia atlantica Markham, 1977 Sclerocrangon jacqueti (A. Milne Edwards) Bahamas; Newfoundland Astalione cruciaria Markham, 1975b Clastotoechus vanderhorsti (Schmitt) U.S. Virgin Islands Bopyrione synalphei Bourdon and Asymmetrione clibanarii Markham, Markham, 1980 1975d Synalpheus goodei Coutiére Clibanarius tricolor (Gibbes) Synalpheus bousfieldi Chace Florida; Bahamas; Ascension Synalpheus brevicarpus (Herrick) Island Synalpheus pectiniger Coutiére Florida; Haiti; Curacao; Gulf of Mexico Bopyrissa wolffi Markham, 1978 Clibanarius tricolor (Gibbes) Clibanarius vittatus (Bosc)
North Carolina to Florida; Belize; West Indies; Gulf of Mexico Bopyrinella thorii (Richardson, 1904) Thor floridanus Kingsley Florida; Curacao; Yucatan Peninsula, Mexico
Asymmetrione desultor Markham, 1975d Pagurus bonairensis Schmitt Pagurus longicarpus Say Pagurus provenzanoi Forest and de
Saint Laurent
Bermuda; North Carolina to Florida; Bahamas; Puerto Rico; Gulf of Mexico
Pylopagurus sp. North Carolina; Florida Keys; Curacao; Bonaire Azygopleon schmitti (Pearse, 1932) Synalpheus brooksi Coutiére Synalpheus hemphilli Coutiére Synalpheus longicarpus Coutiére
Cabirops sp. Synsynella deformans Hay Bermuda
Cancricepon choprae (Nierstrasz and
Synalpheus mcclendoni Coutiére 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; Curacao; Gulf of Mexico Cancrion carolinus 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 brooksi Coutiére Synalpheus goodei Coutiére Synalpheus hemphilli Coutiére Synalpheus longicarpus (Herrick) Synalpheus mcclendoni Coutiére Synalpheus pandionis Coutiére Synalpheus pectiniger Coutiére North Carolina to Florida; Yucatan Pensinsula, Mexico; Belize; Hispaniola; Curacao
1
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 Coutiére Synalpheus hemphilli Coutiére Synalpheus longicarpus (Herrick) 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 floridanus 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 Munida flinti Benedict Munida stimpsoni A. Milne Edwards Cuba; Venezuela
Munidion irritans Boone, 1927 Munida irrasa A. Milne Edwards Florida Keys; Belize
(continued )
112
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 and Brender a Brandis, 1929) Alpheus formosus Gibbes Alpheus heterochaelis (Say) U.S. Virgin Islands; Gulf of Mexico Parabopyrella thomasi (Nierstrasz and Brender a Brandis, 1929) Tozeuma carolinense Kingsley U.S. Virgin Islands
Parapagurion imbricata Markham, 1978 Paguristes tortugae Schmitt Parapagurus sp.
Cuba; Colombia
Parathelges foliatus Markham, 1972b Clibanarius vittatus (Bosc)
Pagurus brevidactylus (Stimpson)
Barbados; Curagao; Trinidad
Parathelges occidentalis Markham,
1972b
Clibanarius tricolor (Gibbes)
Tridopagurus sp.
Pylopagurus corallinus (Benedict)
North Carolina; Florida Keys; Bahamas; Venezuela
Parathelges piriformis Markham, 1972b Paguristes oxyophthalmus Holthuis Pagurus brevidactylus (Stimpson) Pagurus provenzanoi Forest and de
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
Probopyrinella latreuticola (Gissler, 1882) 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 intermedius 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) Tridopagurus 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; Gulf of Mexico Stegias clibanarii Richardson, 1904 Clibanarius tricolor (Gibbes) Bermuda
113
Stegophryxus hyptius Thompson, 1902 Iridopagurus sp. Pagurus annulipes (Stimpson) Pagurus bonairensis Schmitt Pagurus brevidactylus (Stimpson) Pagurus longicarpus Say Pagurus provenzanoi Forest and de Saint Laurent Massachusetts to Florida; Curacao Synalpheion giardi Coutiére, 1908 Synalpheus longicarpus Herrick Yucatan, Mexico Synsynella choprae (Pearse, 1932) Synalpheus brooksi Coutiére Synalpheus longicarpus (Herrick) Synalpheus minus (Say) Synalpheus pandionis Coutiére Bermuda; North Carolina to Florida; Bahamas; Haiti; U.S. Virgin Islands; Gulf of Mexico Synsynella deformans Hay, 1917 Synalpheus brooksi Coutiére Synalpheus longicarpus (Herrick) Synalpheus pectiniger Coutiére Bermuda; Carolinas to West Indies; Gulf of Mexico Urobopyrus processae Richardson, 1904 Ambidexter symmetricus Manning and Chace Processa acutirostris Nouvel and Holthuis Processa canaliculata Leach Processa edulis (Risso) Processa fimbriata Manning and Chace Processa tenuipes Manning and Chace Caribbean; Gulf of Mexico; Brazil; Mediterranean; Eastern Atlantic
114 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 PlEOREISOM yee elas wie a clei maccnite. oe ork Rom eret Sines cee ete ae ee 2
2. Pleon consisting of one or two free pleonites plus pleotelson; body usually dorsally strongly convex; pleopods subequal fae oh Ge: Ghent nara idie Met areln ome Wie aia cana te oF eet menee 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: jicn0 win ore | cis erie wage ac eee eee Serolidae
3. Uropodal ramy flattened, generally not reduced” 7.2.00) - 52 4 Uropodal rami reduced, exopod often hooklike .......... Limnoriidae
FLABELLIFERA * AEGIDAE IPSs
4. Pereopods 4—7 prehensile, with dactyli longer than propodi; antennae reduced, with no clear distinction between peduncle and flagellum
SM as ed eS PEC PRT eS thie kala wide dan vig male subse 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
Biedeusteges Ye Pmeis aes te shies s «ts Hote oeiigadiiee ave Aegidae Maxilliped lacking distal recurved hooks; pereopods 1—3 ambulatory or AGMOSEMCAKIY, PRevensile’ ipo cf eci1 SS ayongy. ebm aise the 99 weenie « 6
6. Maxilliped lacking, or with very reduced endite; maxilla | a strongly RetNeABe MENA D. Sei Segoe Ai an eter ae wists aos Sys Corallanidae Maxilliped with strong endite; maxilla 1 not strongly falcate ........ |
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 | 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 | 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 pleotelson. Pleon of four or five free pleonites 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 SABO Aes Use bs season, CR ae ee eae cs ase Rocinela
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.
Aega (Aega) ecarinata Ie Wg
Key to species of Aega (Aega)
RP es eONP EI UNOS Sete ics aa. 0 aio ls isbn ua YY Gig ai Ba tepore en ok 2 deshaysiana BERS SSCA ALE Be eH Oe ah be Sa oo oe ops ayo ey herve divine ole bon eR ecarinata
Key to species of Aega (Rhamphion)
1. Posterior margin of pleotelson distinctly dentate .............. dentata Posterior margin of pleotelson at most faintly crenulate ....... tenuipes
Aega (Aega) deshaysiana (H. Milne Edwards, 1840) Figure 51A
DIAGNOSIS ? 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
DIAGNOSIS ¢ 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; C, Aega (Rhamphion) dentata; D, 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.
Rocinela 119
Aega (Rhamphion) dentata Schioedte and Meinert, 1879 Figure 51C
DIAGNOSIS @ 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 2 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
EV CSMEOILIC MOUS 57.07724. sc Gli DARSs SOE UL ie ae ieaies ten ee ere oculata By CS et COMM GUOUS. % 5. 3d. ocie sueltns sfliogy a brecee Sle Gisiaacs & ccehans ss OR ere: 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 * AEGIDAE
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 2 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 ¢ 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 2 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
Figure 52. A, Rocinela cubensis; B, Rocinela insularis; C, Rocinela oculata; D, Rocinela signata.
122 FLABELLIFERA * CIROLANIDAE
single spine on posterior margin. Pleotelson 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 of jew- 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 Bal- istes vetula, on saucereye porgy Calamus calamus; Jamaica, on French grunt Haemulon flavolineatum, 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. Pleon 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 PROP ITIP) ML ie eee eae Ws ARE, Spm 4 le UO wre «TS Eurydicinae Clypeus flattened, not projecting; pleonite 5 lacking free lateral margin, avenapped*by-pleonite’ 4) 2020 oe Pek. ek el dav Se ES 2
2. Pereopods 1—3 with ischium and merus not anterodistally produced; antennal peduncular articles 4 and 5 subequal; secondary unguis present on percopodal dactyli- y+. -Pe Ts eee ee ae. Cirolaninae
Pereopods 1—3 with ischium and merus anterodistally produced; antennal peducular articles 3 and 4 subequal; no secondary unguis GHePEneOPOUAlGAClYIN ' 8 s1s.8 sete Picea sod Mas oe eo eA a a ares 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 d with copulatory stylet articulating basally.
Key to genera of Cirolaninae
1. Pleopods having accessory branchial filaments ............ Bathynomus Pleopods lacking accessory branchial filaments .................... 2 eet COO ene LCUMNONMIGE 0.8 4 hon edicles averse Mele d gkn ek Aew sine Os es 7 Picopad -ltnereperculiforni shoe! 220m. Gis he eh. Bi gee oreei . 3 3. Pleopods 3-5, endopods lacking, or with very few, marginal setae ... 4 Only endopod of pleopod 5 lacking marginal setae ........... Cirolana
(continued )
124 FLABELLIFERA * CIROLANIDAE
Key to genera of Cirolaninae (Continued)
4. Merus of pereopod | posterodistally produced; merus of pereopods 2
and 3 anterodistally produced s'.. c 2 sn: eae meee Bahalana Meri ‘of pereopods. 1—3- not markedly produced ......4..5022 +5598 5
5. ‘Animal able .to.conglobates 2 cepts pene Yee ee tae Creaseriella Animal unabletejconclobatelusescis tastes alowed: Asean 6
6G. Mandibular palp directed’ antenorly” ... 2. 5..5.-.-----.9:- Anopsilana Mandibular palp-directed posteriorly .....-5..-24-.200-5458 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 antegumental: pigment, |< f<.0 6 f.6, we2ho cee etm ory 3 2. Frontal lamina distally rounded; projectune {25.°..-. 25. 5-22. browni Frontal lamina distally acute, not projecting .................. jonest 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 cubensis
Posterior margin of pleotelson with 10—12 spines; found in cave on Haiti
©) ehce: eB, 6s! epee (a, 8/8) 6. 6) (aL 6) Sm, .0) «0! ee) ae |e) iWiqiej) © fefle| (elim) se) 1) 8 le a) e's) a) .e) (ell eie/ce! e-em imam
acanthura
5. Posterior margin of pleotelson with eight spines; found in cave on Grand Caymai: :.6 aka 5 tase pyedonsthe aioeie aie ee eee crenata Posterior margin of pleotelson with four spines; found in cave on Haiti radicicola
Bie 68) 0) Ge Om ee Oe) Sh Si ep ee, wel in esfee) ee) a6) a) fo) @ (6) eo (ame. le louie. ote 6) (a) ese mies (ee cee ane
Anopsilana crenata 125
somewhat expanded. Antennular peduncle of two articles. Maxillipedal en- dite with two coupling hooks. Pereopod | prehensile, pereopods 2—3 weakly prehensile, pereopods 4—7 ambulatory. Pleopod 2 d, copulatory stylet artic- ulating at base of endopod. Pleopods 3-5, exopods biarticulate, endopods lacking marginal setae. Uropodal sympod produced along mesial margin of endopod.
Anopsilana acanthura (Notenboom, 1981) (was fla L Lome. Figure 53A,B
DIAGNOSIS 6 7.0 mm. Lacking eyes and integumental pigment. Frontal lamina anteriorly rounded. Posterior margin of pleotelson with 10—12 spines.
RECORDS Well at Marigot, Haiti.
Anopsilana browni (Van Name, 1936) Figure 53C,D
DIAGNOSIS. 6 11.1 mm, 2 10.0 mm. Eyes well developed, pigmented. Dor- sal integument strongly pigmented with red-brown chromatophores. Frontal lamina as wide as long, anteriorly rounded. Cephalon with two fused mid- dorsal tubercles near posterior margin. Pereonites and pleonites each with row of tubercles near posterior margin. Pleotelson triangular, with scattered dorsal tubercles, apex rounded, with eight spines.
RECORDS River in Santa Clara Province, Cuba (freshwater); Sittee River and Salt Creek, Stann Creek District, Belize (brackish water). Golfo de Nicoya, Pacific Costa Rica, in red mangroves.
Anopsilana crenata Bowman and Franz, 1982 Figure 53E,F
DIAGNOSIS 6 6.2 mm. Lacking eyes and integumental pigmentation. Frontal lamina longer than wide, anteriorly rounded. Posterior margin of pleotelson with eight spines.
RECORDS West Bay Cave, Grand Cayman Island.
126 FLABELLIFERA * CIROLANIDAE
Figure 53. Anopsilana acanthura: A, 2; B, anterior cephalon. Anopsilana browni: C, &; D, anterior cephalon. Anopsilana crenata: E, 2; F, anterior cephalon. Anopsilana cubensis: G, 2; H, anterior cephalon.
Anopsilana cubensis (Hay, 1903) Figure 53G,H
DIAGNOSIS ¢ 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 ¢ 5.9 mm. Eyes well developed and pig- mented. Dorsal integument strongly pigmented with almost solid central area on pereonites 1—7. ¢ cephalon with three low tubercles near posterior margin; pereonite 1 with four to six low tubercles. 2 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 6.3mm, 2 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 Débarasse, a spring near Jérémie, 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 d 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.
128 FLABELLIFERA ¢ CIROLANIDAE
Key to species of Bahalana
1. Pleopods 3—4, endopods lacking setae; maxillipedal endite with one coupling hook sat tin Ge SOMERS als haat on tes eae eae sete cardiopus Pleopods 3—4, endopods with few setae; maxillipedal endite with one or twovcoupling NOOKS 4.....025 eae. s: aeele «Burg te eo eee 2
2. Antennular peduncle, article 3 longest; maxillipedal endite with two CouplingyhoOoksSgee 2 ~ seica sucht a age = «bac. ae ee geracei
Antennular peduncle, article 2 longest; maxillipedal endite with one COUPER HOON teste san css ask, cana tale, Be 6 bt we os en mayana
Bahalana cardiopus Notenboom, 1981 Figure 55A,B
DIAGNOSIS ¢ 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 15.0 mm, d 8 mm. Maxillipedal endite with two coupling hooks. Pereopod 1, 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 2 8.4 mm, d 10.0 mm. Clypeus acutely pointed. Antennular peduncle, article 2 longest. Maxillipedal endite with one coupling hook. Per-
Bathynomus 129
B
Figure 54. Anopsilana jonesi: A, 36; B, anterior cephalon; C, pleopod 2, 3; D, pleopod 3. Anopsilana radicicola: E, 3; F, anterior cephalon.
eopod 1, meral projection rudimentary. Pleopods 3—4, endopods with few marginal setae, pleopod 5 lacking marginal setae. Uropodal exopod narrow, 73 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, 2; B, pleopod 3. Bahalana geracei: C, 2; D, pleopod 3; E, pereopod 1; F, pereopod 2; G, anterior cephalon. Bahalana mayana: Hy 2.
Calyptolana 131
Figure 56. Bathynomus giganteus.
Bathynomus giganteus A. Milne Edwards, 1879 Figure 56
DIAGNOSIS Up to 280 mm. Large pigmented eyes present, not visible dor- sally. Antennule with small exopod distally on peduncular article 3. Pleopods with marginal setae on all rami; pleonites 3 and 4 with epimera produced posteriorly. Posterior margin of pleotelson with median tooth and five or six teeth on each side.
RECORDS Gulf of Mexico; Caribbean; Bahamas; Florida Keys; 360—2300
m.
REMARKS Gut-content analysis of these deep-water giants has shown them to be scavengers, commonly feeding on dead fish, cephalopods, crabs, and polychaete worms.
Calyptolana Bruce, 1985
DIAGNOSIS. All pereopods ambulatory; each dactylus with secondary un- guis. Pleopod | operculate, longer than following pleopods. All pleopodal rami except endopod of pleopod 5 with marginal setae.
132 FLABELLIFERA ¢ CIROLANIDAE
Calyptolana hancocki Bruce, 1985 Figure 57
DIAGNOSIS 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 Pleotelsom postenionly narmowed Hills, 29g Ie 2 ase eee 3 See ee 4
2. Uropodal endopod broad, distally rounded, lacking marginal spines aoa dp hs SRhe nas Spe anc spovena ve wy oe a NRL ann a eee a eee ca obtruncata
earn elie 20h OE Ta aaa One GARE Chg) UR As Nich tr eee 3
3. Posterior margin of pleotelson bearing spines; uropodal endopod not lenibate eh Fig a. Tea BT TAI ee Le ee ke minuta Posterior margin of pleotelson faintly crenulate, lacking spines; uropodal ‘endoped strongly dentate 52. 5.-2..5. =1 nen crenulitelson
4. Uropodal endopod evenly tapering to acute apex; uropodal exopod length about four times greatest width. 2%. 425... 6.26427 e05 albidoida Uropodal endopod, outer margin convex; uropodal exopod length less thanfour times 'greatest width yw ieHeliin Se eee parva
Cirolana crenulitelson 133
Figure 57. Calyptolana hancocki: A, 2; B, ventral pleon.
bearing marginal setae except endopod of pleopod 5. Copulatory stylet on endopod of ¢ 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 | 3d, endopod markedly narrowed in distal half. Pleopod 2 d, 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 Off Lucaya, Grand Bahama, 180-220 m.
Cirolana crenulitelson Kensley and Schotte, 1987 Figure 58D-F
DIAGNOSIS <6 7.0mm, 2 7.0 mm. Antenna reaching posteriorly to anterior of pereonite 3. Pleopod 2 6, copulatory stylet reaching by '/s its length be-
134 FLABELLIFERA * CIROLANIDAE
Figure 58. Cirolana albidoida: A, 3; B, uropod; C, pleotelsonic apex. Cirolana crenulitelson: D, 2; E, pleotelsonic apex; F, uropod. Cirolana minuta: G, 3; H, anterior cephalon; /, uropod.
yond rami. Uropodal endopod with mesial margin broadly convex, serrate, apically acute; exopod about 2.5 times longer than wide, mesial margin weakly convex, apically acute. Posterior margin of pleotelson truncate, faintly crenulate, lacking spines.
RECORDS Carrie Bow Cay, Belize, 36 m.
Cirolana parva 135
Cirolana minuta Hansen, 1890 Figure 58G-—I
DIAGNOSIS 6 8.9 mm, ¢ 5.0 mm. Antenna reaching posteriorly to per- eonite 3. Pleopod 2 d, copulatory stylet reaching by about '/ of its length beyond rami. Uropodal endopod, mesial margin broadly convex, with apical angle of about 90°; exopod about three times longer than wide, parallel sided for proximal two-thirds, with marginal spines, apically narrowly rounded. Pleotelson with posterior margin subtruncate to broadly rounded, with about eight marginal spines.
RECORDS St. Thomas, U.S. Virgin Islands; Tobago; 180-220 m.
Cirolana obtruncata Richardson, 1901 Figure 59A,B
DIAGNOSIS. 6 11.3 mm, 2 11.0 mm. Antenna reaching posteriorly to mid- dle of pereonite 3. Pleopod 2 6, 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 6.9mm, 2 7.9 mm. Cephalon with furrow between eyes hav- ing middorsal posterior deflection. Antenna reaching posteriorly to pereonite 4. Pleopod | 6, endopod narrowed in distal third. Pleopod 2 6, copulatory stylet reaching by '!/s 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, 2; B, uropod. Cirolana parva: C, 2; D, uropod; E, pleotelson and uropods.
Haptolana 137
Figure 60. Cirolana parva,
anteroventral cephalon.
Creaseriella Rioja, 1953
DIAGNOSIS Antennular peduncle of two articles (articles 1 and 2 fused). Antennal peduncle of five articles. Pereopods all ambulatory. Penial rami fused to form short stout process. Pleopod 2 in d with copulatory stylet inser- ted at base of endopod. Pleopods 3—5, endopods lacking marginal setae. Pleonite 5 lacking free lateral margins.
Creaseriella anops (Creaser, 1936) Figure 61A,B
DIAGNOSIS 6 15.5 mm. Animal able to conglobate. Eyes lacking. Frontal lamina pentagonal, longer than wide, with transverse ridge at widest point. Maxillipedal endite with four or five coupling hooks. Pleotelson wider than long, posterior margin very broadly rounded and finely crenulate. Both uropodal rami distally rounded, margins bearing spines and setae, sympod produced along mesial margin of endopod.
RECORDS Several caves and cenotes on the Yucatan Peninsula, Mexico.
Haptolana Bowman, 1966
DIAGNOSIS Mandibular palp directed posteriorly. Antennular peduncle of two articles, basal article expanded. Pereopods all prehensile, with dactyli closing in propodal groove. Pleopods 3—5, exopods with partial suture and marginal setae; endopods undivided, lacking marginal setae. Pleonite 4 over- lapping pleonite 5 laterally.
138 FLABELLIFERA * CIROLANIDAE
Figure 61. Creaseriella anops: A, adult; B, frontal lamina. Haptolana trichostoma: C, adult; D, frontal lamina. Oncilorpheus stebbingi: E, 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, 2/3 width of endopod, both rami with marginal setae and spines.
RECORDS Cave in Camaguey Province, Cuba.
REMARKS A second species of Haptolana, H. somala Messana and Chelazzi, has been described from northern Somalia in Africa.
Natatolana 139
Oncilorpheus Paul and Menzies, 1971
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 @ 11.0 mm. Body narrow, about five times longer than wide. Pleotelson triangular, bearing faint middorsal longitudinal ridge, apex nar- rowly rounded.
RECORDS Off Venezuela, 73 m.
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,
MEKUS: ANGUCATOUS MAUCENEM) 2.5 cons. cates eek note oe oes 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 6, copulatory stylet articulating basally on endopod.
140 FLABELLIFERA ¢ CIROLANIDAE
Natatolana gracilis (Hansen, 1890) Figure 62
DIAGNOSIS. 6 8.0 mm. Antennule short, not reaching distal end of antennal 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. 3d 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 2 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, 2; B, frontal lamina; C, pleotelson; D, uropod; E, pereopod 7.
142 FLABELLIFERA * CIROLANIDAE
\
ok a f
\ F
Figure 63. Politolana impressa: A, 3, lateral view; B, uropod, (setae and spines omitted). Politolana polita: C, 3; D, pereopod 1; E, 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.
Politolana polita 143
Figure 64. Politolana polita, anteroventral cephalon.
Politolana polita (Stimpson, 1853) Figures 63C—F, 64
DIAGNOSIS 6 27.0 mm, 2 29.0 mm. Antennule barely reaching distal end of antennal peduncle. Frontal lamina basally slender, anteriorly expanded, just visible in dorsal view. Coxae lacking impressed oblique line. Uropodal endopod distal to emargination narrowed, margin convex. Pleotelson pos- teriorly broadly rounded.
RECORDS Bay of Fundy, Canada, to Florida Keys, 2—600 m; Gulf of Mexico.
Subfamily Eurydicinae Stebbing, 1905
DIAGNOSIS Clypeus projecting. Pleonite 5 with free lateral margins (except in Xylolana). Penes prominent. Pleopod 2 of d with copulatory stylet artic- ulating subbasally, medially, or subapically.
Key to genera of Eurydicinae
1. Uropodal sympod not produced along mesial margin of endopod Eurydice Uropodal sympod produced along mesial margin of endopod ....... 2
2. Rostrum prominent, fused with frontal lamina, separating antennal SES aie Bole hy SES COS eo ae RO tors ree RE EOS ee ree Try ar 5
(continued )
144 FLABELLIFERA * CIROLANIDAE
Key to genera of Eurydicinae (Continued)
3: -Pleon: of five ireepleanites. plus pleotelson .).. 2... ...<=- soeae oe wees 4 Pleon of three free pleonites plus pleotelson ......:...20%:- Colopisthus
4. Endopods of pleopods 3—5 lacking marginal setae, or with no more than three marginal setae; copulatory stylet of endopod of pleopod 2
in; ©, articulating SUB emmmMANy (oc... sc see see occ we ess Arubolana Marginal setae lacking only on endopod of pleopod 5; copulatory stylet
of pleopod 2 endopod in ¢ articulating basally ........ Metacirolana
5. Clypeus conical; uropodal endopod lacking notch in outer distal margin
tee Sal BRE APSR wt aR Ny st ae PREG 7a emt alee cage ce Xylolana Clypeus flattened; uropodal endopod with notch in outer distal margin
gin aE AMPS ees Saran cs kus Wy Syeras a MONG aes eae SURES ara ie Oi il a Excirolana
Arubolana Botosaneanu and Stock, 1979
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 | and 2 and sometimes pereopod 3 prehensile; pereopods 4—7 ambulatory. Pleopods | and 2, rami undivided. Pleopod 2 3 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. Byes: present, Salk yiycta 33.2 GUN AR ee parvioculata VCS. ASETIE a0) car. tayay'e' arcs at a ae a crea a ee caren 2
2. Pleotelson posteriorly rounded; rostrum not distinct in dorsal view
BS TAS Chaka te See ay ee ee aruboides
Arubolana parvioculata 145
Arubolana aruboides (Bowman and Iliffe, 1983) Figure 65A—C
DIAGNOSIS 6 3.9mm, 2 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. ¢ and 2 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 crenulations 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, ¢ 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.
146 FLABELLIFERA * CIROLANIDAE
Figure 65. Arubolana aruboides: A, 3; B, pereopod 1; C, pereopod 2. Arubolana imula: D, 2. Arubolana parvioculata: E, 3; F, pleopod 2, ¢.
Colopisthus 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.
Eurydice convexa 147
Colopisthus parvus Richardson, 1902 Figure 66A
DIAGNOSIS @ 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. ¢ 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
imenrontal lamina distally truncate to faimtly bilobed .........5...-.-. z Frontal lamina lanceolate, distally acute: 2.52 e 4-2 2 chan: personata
2. Posterior margin of pleotelson between notches rounded, with four INOdEerAbe SPIES! a. Shree ee HEE EE ake was Deis P98 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, 2 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. ¢d: Plicate process on antennal flagellar articles about '/s length of article. Pleopod 2 with copulatory stylet distally blunt, reaching well beyond rami.
148 FLABELLIFERA * CIROLANIDAE
Figure 66. Colopisthus parvus: A, 2. Eurydice convexa: B, 3; C, uropod; D, antennule; E, pleotelsonic apex. Eurydice personata: F, 3, lateral view; G, pleotelsonic apex. Eurydice piperata: H, 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.
Eurydice personata Kensley, 1987a Figures 66F,G; 67C
DIAGNOSIS <6 6.0 mm, ovigerous 2 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. ¢: 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.
Eurydice piperata Menzies and Frankenberg, 1966 Figure 66H
DIAGNOSIS 6 4.0 mm, ? 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. d: Plicate organ on antennal flagellar articles about '/eé 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
150 FLABELLIFERA * CIROLANIDAE
Figure 67. Eurydice convexa: A, anteroventral cephalon; B, frontal lamina. Eurydice
personata: C, frontal lamina.
lacking marginal setae. Pleonite 5 with free lateral margins, not overlapped by pleonite 4. Uropodal sympod produced along mesial margin of endopod.
Key to species of Excirolana
1. Pleotelson with two anterior hollows clearly joined by impressed line;
uropodal endopod about half length of exopod .......... braziliensis Pleotelson with two anterior hollows not connected by impressed line; uropodal endopod about two-thirds length of exopod ....... mayana
Excirolana braziliensis Richardson, 1912a Figures 68A—C, 69A—C
DIAGNOSIS 6 6.0 mm, 2 7.5 mm. Frontal lamina very slender between antennal bases, widening anteriorly into rounded structure between anten-
VY
.?)
Figure 68. Excirolana braziliensis: A, 6; B, pleopod 2 6; C, uropod, (setae and spines omitted). Excirolana mayana: D, 2; E, pleopod 2 ¢; F, uropod, setae and spines omitted.
152 FLABELLIFERA * CIROLANIDAE
Figure 69. Excirolana braziliensis: A, anteroventral cephalon; B, pleotelson; C,
uropodal endopod. Excirolana mayana: D, anterodorsal cephalon; FE, anteroventral cephalon.
nular bases, joined to rostrum by very slim isthmus. Clypeus distally sub- acute. Uropodal endopod about half length of exopod. Pleotelson with two lateral hollows defined and connected by clear impressed line; posterior mar- gin evenly convex, bearing numerous plumose setae.
Metacirolana {533
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 mayana (Ives, 1891) Figures 68D-—F, 69D,E
DIAGNOSIS 6 8.2mm, 2 10.0 mm. Frontal lamina between antennal bases about half anterior width. Clypeus anteriorly rounded. Uropodal endopod about 7/3 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 d than in
a
Key to species of Metacirolana
cts OME POSECHOE Ly LEUMEALC oho ances 4 >, s ckepoyeye ePeiahs josie dis: Afo Shale xs halia Melson posteriorly rounded Or angulate . ... . o..<cjs.sjoi ois ieee eis. > wiecarar 2 2. Posterfr margin of telson an obtuse angle ................ agaricicola SCEScrta OS LELIOU yah OUNCE sn ote a0e ais: bos vine sie eto bi © (ob lap eh es nse ae rete 3
3. Posterior margin of telson narrowly rounded; uropodal rami, margins SEM OITGUI A CUCIMEAEG eri Pela, Stan ee Oe aia nl ace lb yes cvelays waist menziest Posterior margin of telson broadly rounded; uropodal rami, margins BBSOULELY Cente x. seis On eee Lie. Sr hace sphaeromiformis
154 FLABELLIFERA * CIROLANIDAE
Metacirolana agaricicola Kensley, 1984 Figure 70A—C
DIAGNOSIS d 2,6 mm, ovigerous 2 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 2 2.7 mm. Antennular flagellum of 10 articles in 2, 14 in d. Antennal flagellum of 10 articles in 2, 11 in ¢. 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 6 2.3 mm, ovigerous 2 2.4 mm. Antennular flagellum of six articles in 9, eight in 6. Antennal flagellum of nine articles in 2, 10 in 6. Posterior margin of telson broadly rounded, finely dentate. Uropodal exopod half width of endopod, margins dentate, apically acute; endopod, margins
dentate, apically acute.
RECORDS Carrie Bow Cay, Belize, intertidal to 30 m, usually in coral rubble.
Metacirolana sphaeromiformis (Hansen, 1890) Figure 71C,D
DIAGNOSIS. 6 2.5 mm, 2 3.2 mm. Antennular flagellum of three articles. Antennal flagellum of eight articles. Posterior margin of telson narrowly
Metacirolana sphaeromiformis 193
Figure 70. Metacirolana agaricicola: A, 2; B, uropod; C, maxilliped. Metacirolana halia: D, uropod; E, 3; F, &.
rounded, obscurely dentate. Telson with low rounded middorsal ridge and pair of lateral ridges. Uropodal exopod more than half width of endopod, margin dentate; exopod distally broadened, margin dentate, with few sen- sory spines.
156 FLABELLIFERA * CIROLANIDAE
Figure 71. Metacirolana menziesi: A, 2; B, uropod. Metacirolana sphaeromiformis: C, 2; D, uropod.
RECORDS Looe Key, Florida, intertidal reef crest; Turks and Caicos Is- lands, | m; St. Thomas, U.S. Virgin Islands.
Xylolana Kensley, 1987a
DIAGNOSIS Frontal lamina and rostrum fused, broad, separating antennu- lar bases. Clypeus conical, projecting. Maxillipedal endite reduced, lacking
Alcirona iay7/
coupling hooks; palp of five articles. Copulatory stylet in d 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 6 2.6mm, 2 3.3 mm. Body about four times longer than great- est width. Uropodal exopod about */3 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 Cays, Belize, in dead red mangrove roots, | 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
ee Maxilla | with single strong falcate spime. 2.20. 2/60) .13% es ee ee sa soe 2 Maralla 1, with: two large talcate:spines: «. 24. :2) Pejsn. w ser ivi 2 Alcirona
2. Maxilla 2, distal article slender-elongate; article 2 of maxillipedal palp
COVES S| ae rae thelial suse Seer Nite tp ia aera ete 1 pew Nalicora Maxilla 2 distally bluntly bilobed; article 3 of maxillipedal palp longest soc, GRUPPE gees ae ae ay, Pate ais arena rote ater Ota Excorallana
Alcirona Hansen, 1890
DIAGNOSIS Antennular peduncle of two articles. Mandible lacking molar. Maxilla | 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 jonly @. 2... 10.0.4 46. c eee krebsi
Alcirona insularis Hansen, 1890 Figure 73A
DIAGNOSIS. 6 and @, 5.0 mm. Posterior half of body, especially in d, 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 <6 10 mm, ovigerous 2 15.5 mm. Posterior half of body, espe- cially ind 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 ¢ specimen of Alcirona from Panama Bay (Pa- cific) has the characteristic pereopod | of A. krebsii, but has the rows of stuff 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, B, 3; C, maxilliped; D, pleopod 2 d; 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-
160 FLABELLIFERA « CORALLANIDAE
Figure 73. Alcirona insularis: A, pereopod 1. Alcirona krebsi: B, 3; C, maxilla 1; D, pereopod 1.
hensile, pereopods 4—7 ambulatory. All rami of pleopods bearing marginal setae. Pleotelson often with characteristic spination and tuberculation; 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
MPP ESAGOULIPUOUSE. |. Haste eck clas qslstettiihk « Jul+ siertintis ® Gate ae 5 Deas 2 Eves Well SGpabdateted.. shisha, os at Aeids, 4 Unie. eden Me eM eee 4 Beepex of plcotclson with deep slit .. ..\...Jcnusisglees co oe eee fissicauda Pese MaMa PUCOLEISONSCMENS. y.2 02 cos Stacie tanta s ae «i ceeimioe S Oe EO oe es 3 pepelecotcison-with lateral ancision:.......05.. i. 52 2. eas she Gees » oculata Picoteisonmacking lateral incision: ~ . .< = a)..s is. see sas 6 eee warmingit Peercotcison with lateral incision” =. o 6.0. 3. ko tees neces es 6 Picoteison lacking fateral incision’. <2. soso eee ea wee yess es 5 5. Frontal lamina linguiform, anteriorly rounded ............. berbicensis Frontal lamina posteriorly with faintly concave margins, anteriorly SELEY KS he Metre Ce Oe PRE PUR ere MEP mee APENAYIBN Meng GEA delaneyi 6. Frontal lamina strongly grooved for entire length ................. 7 Frontal lamina with,ventral surface flat. )s< i.cca.... fs.0 22 250s ncn 8
7. Pleotelson with medial row of small tubercles flanked by row of larger Pubercles may postenon Wale 7.362) 2) no o1 od cepa slaeiaw es mexicana Pleotelson lacking rows of tubercles in posterior half ........ antillensis
8. Frontal lamina distinctly bell shaped; d 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; d cephalon with two pairs of tubercles and basal antennulararticle each with tubercle: <.c.. .1. <:cr.6 os: 9 sisiam are sexticornis
Frontal lamina anteriorly narrowly rounded, length about twice basal width; ¢d cephalon with three tubercles, tubercles lacking on AACIEINULALMOASES! “(Sec URE sa: Sais etatoe nls oleae sere s tricornis tricornis
162 FLABELLIFERA * CORALLANIDAE
Excorallana antillensis (Hansen, 1890) = E aculicauda (Miers 18 I) Figure 74A—D
DIAGNOSIS. 6 11.0 mm, ovigerous 2 15.0 mm. Cephalon unornamented. Eyes well separated. Frontal lamina parallel sided, length twice basal width, anteriorly broadly rounded. Pleonites 2—4, posterolateral margins tubercu- late, middorsal posterior margin excavate with strong middorsal tubercle; pleonite 5 with strongest tubercles submedian, posterior margin not exca- vate. Pleotelson with low middorsal ridge, strong basal tubercles; lateral inci- sions present; two submedian patches of spines; apex narrowly rounded.
RECORDS Florida Keys; St. Thomas, U.S. Virgin Islands; Puerto Rico; Quintana Roo, Mexico; Carrie Bow Cay, Belize, 5—18 m.
Excorallana berbicensis Boone, 1918 Figure 74E,F
DIAGNOSIS 6 9.9 mm, 2 12.0 mm. Eyes well separated. Frontal lamina about twice longer than wide, posteriorly parallel sided, widening anteriorly to broadly rounded apex. Subadult 6 cephalon unornamented except for pair of very low tubercles mesial to eyes. Posterior margins of pleonites 3—5 very faintly tuberculate. Pleotelson with submedian pair of low tubercles basally; lacking lateral incisions; apex rounded.
RECORDS Guyana; French Guiana; Guadeloupe.
Excorallana fissicauda (Hansen, 1890)
DIAGNOSIS 11 mm. Cephalon unornamented. Eyes contiguous. Frontal lamina unknown. Pleonite 5 with three strong mesial, and several smaller lateral tubercles on posterior margin. Pleotelson with two strong submedian basal tubercles; lateral incision lacking; posterior margin with deep open incision.
RECORDS St. Thomas, U.S. Virgin Islands.
REMARKS This species was described from a single specimen, and has not been recorded since. After examining the holotype, Paul Delaney (in litt.) suspects that the terminal incision of the pleotelson may be the result of an injury.
Excorallana oculata 163
F
Figure 74. Excorallana antillensis: A, 6; B, frontal lamina; C, maxilla 2; D, maxilliped. Excorallana berbicensis: E, 3; F, frontal lamina.
Excorallana oculata (Hansen, 1890) Figure 75A,B
DIAGNOSIS 6 6.9 mm, 2 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.
Figure 75. Excorallana oculata: A, 3; B, frontal lamina. Excorallana quadricornis: C, 3; D, frontal lamina. Excorallana sexticornis: E, 36; F, frontal lamina. Excorallana tricornis tricornis: G, 3; H, frontal lamina. Excorallana warmingi: I, 3; J, frontal
lamina.
Excorallana tricornis tricornis 165
Excorallana quadricornis (Hansen, 1890) Figures 75C,D; 76A—C
DIAGNOSIS 6 13.2 mm, 2 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 7.9mm, ovigerous 2 6.9—8.3 mm. Eyes well separated. Basal antennular peduncular article in d 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, 2 9.9 mm. Basal antennular article narrow, not dilated. Cephalon in d 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.
166 FLABELLIFERA * CORALLANIDAE
400HM
Figure 76. Excorallana quadricornis: A, cephalon and pereonite 1 6; B, ventral cephalon; C, frontal lamina enlarged. Excorallana sexticornis: D, cephalon and pereonite 1 6; E, cephalon @; 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 intertidal coral rubble, in coarse sediments in Syringodium and Thalassia sea- grass beds, on brown alga Turbinaria, on Madracis sp. sponge, on Agaricia sp. coral; Gulf of Mexico.
Excorallana warmingii 167
Figure 77. Excorallana tricornis tricornis: A, cephalon and pereonite 1 6; B, ventral
cephalon; C, pleotelson and uropods.
REMARKS The subspecies Excorallana tricornis occidentalis Richardson, 1905a, from southern California, differs from the Gulf and Caribbean subspecies in lacking a gap between the margins of the pleotelsonic incision, and in having a relatively wider uropodal exopod which shows a distinctly asymmetrical apical notch.
Excorallana warmingii (Hansen, 1890) Figure 751, J
DIAGNOSIS 6 9.7 mm, ¢ 12.0 mm. Cephalon unornamented. Eyes con- tiguous, occupying most of dorsal surface of head. Posterior margins of pleonites very faintly tuberculate. Frontal lamina, length slightly more than twice basal width, tapering anteriorly to rounded apex. Pleotelson unorna- mented except for two faint submedian tubercles basally; lateral incisions lacking; apex broadly rounded, with five low but distinct marginal teeth.
168 FLABELLIFERA * CORALLANIDAE
Ra
—
“—
Figure 78. Nalicora rapax: A, 2; 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.
FLABELLIFERA * CYMOTHOIDAE 169
Nalicora rapax Moore, 1901 Figure 78
DIAGNOSIS 6 6.9 mm, ovigerous 2 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 d. Pleotelson wider than long; lateral margins faintly sinuous; apex rounded.
RECORDS Florida Keys, 55 m; Puerto Rico, 50-150 m; Gulf of Mexico off lorida, 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 * CYMOTHOIDAE
Key to genera of Cymothoidae
. Antennule broader and usually longer than antenna; cephalon very
weakly sunkiimto. Perconmite, Lo. 05 erect aut x «8-0 the tao ae Ste eee, eee 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: ioc os ane... oo eee 3
Bases of antennules comtieuwous: 2.6.4.0. san 4s. e. 7 see: Glossobius
. Body curved to one side; pleonite 1 extended laterally more on one side
thanon others ic. a.doa shee ee attic eee ee woe ote Mothocya Body rarely curved to one side; pleonite 1 extended equally on each
SIGE £0 loo can cs pour ne ens Gaetan kam Ae SR Nc mee heen ae Renocila
ee Le an CON ee Net eee eT ee eee Fae Agarna
. Cephalon not immersed in pereonite 1; posterior margin of cephalon {LISITMATE «0-2 Need nares: 2 ele etn Siete otochae ate eee ee ea aie ee eee 6 Cephalon to some degree immersed in pereonite 1; posterior margin of cephalon ‘not trsinMater 4 tary He = poland daca wie peel uel wise ed 7
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
PCREORULES. ofa are. cud acters aera) Oa at re EE Ose ea Nerocila
. Basal antennular articles expanded and contiguous ......... Ceratothoa Basal antennular articles expanded but not contiguous, or basal
antennular articles neither expanded nor contiguous ............. 8
Basal antennular articles expanded but not contiguous ......... Kuna
Basal antennular articles neither expanded nor contiguous ......... a
9. Pleonal 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 IMUINERSeC AN. PET CONME Jj cos iia eee haat ee ecie ee Cymothoa
FLABELLIFERA * CYMOTHOIDAE 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. CYMOTHOID PARASITES FROM THE CARIBBEAN AREA, LISTED BY FISH
HOST SPECIES
Fish host
Cymothoid parasite
Site of attachment
Abudefduf saxatilis
Acanthurus bahianus
Acanthurus chirurgus
Alutera schoepfi Anchoa lamprotaenia Apogon lachneri Apogon maculatus Apogon townsendi Arius felis Astrapogon stellatus Batrachoides surinamensis Caranx hippos Caranx latus
Caranx ruber
Caranx sp. Chaetodipterus faber Chaetodon capistratus Chaetodon ocellatus Chaetodon sedentarius Chaetodon striatus
Anilocra abudefdufi Kuna insularis Anilocra acanthuri
Anilocra acanthuri
Nerocila acuminata Lironeca tenuistylis Mothocya bohlkeorum Renocila colini Renocila colini Nerocila acuminata Mothocya bohlkeorum Nerocila acuminata Cymothoa oestrum Cymothoa oestrum Cymothoa oestrum Cymothoa oestrum Nerocila acuminata Anilocra chaetodontis Anilocra chaetodontis Anilocra chaetodontis Anilocra chaetodontis
beneath eye
gill chamber
? at base of pectoral fin; immature on or near pectoral or pelvic fin
? 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
next to dorsal fin
on or at base of fin
in gill chamber
on or at base of fin
inside mouth
inside mouth
inside mouth
inside mouth
on or at base of fin
beneath eye
beneath eye
beneath eye
beneath eye
(continued )
172
TABLE 3. (Continued)
Fish host
Cymothoid parasite
Site of attachment
Chilomycterus schoepfi Chromis cyaneus Chromis multilineatus Cynoscion nebulosus Cynoscion sp. Epinephelus cruentatus Epinephelus fulvus Epinephelus guttatus Epinephelus itajara Epinephelus sp. Exocoetus spp.
Gerres rhombeus Haemulon aurolineatum Haemulon carbonarium Haemulon chrysargyreum Haemulon flavolineatum Haemulon macrostomum Haemutlon plumieri Haemulon sciurus
Hemirhamphus brasiliensis
Hirundichthys speculifer Holacanthus tricolor Holocentrus ascensionis
Hyporhamphus unifasciatus
Leiostomus xanthurus
Lepiosteus spatula Lutjanus analis Lutjanus mahogoni Lutjanus synagris Megalops atlanticus Monacanthus ciliatus Mugil cephalus Mpyripristis jacobus
Ocyurus chrysurus Orthopristis chrysoptera
Nerocila acuminata Anilocra chromis Anilocra chromis Cymothoa excisa Cymothoa oestrum Anilocra haemuli Anilocra haemuli Anilocra haemuli Nerocila acuminata Cymothoa oestrum Glossobius impressus Lironeca redmanni Anilocra haemuli Anilocra haemuli Anilocra haemuli Anilocra haemuli Anilocra haemuli Anilocra haemuli Anilocra haemuli Glossobius hemirhamphi Glossobius impressus Anilocra holacanthi Anilocra holocentri
Mothocya nana Nerocila acuminata Cymothoa excisa Lironeca redmanni Nerocila acuminata Cymothoa excisa Cymothoa excisa Cymothoa excisa Cymothoa oestrum Nerocila acuminata Nerocila acuminata Anilocra myripristi
Cymothoa excisa Cymothoa excisa
on or at base of fin beneath eye beneath eye inside mouth inside mouth beneath eye beneath eye beneath eye on or at base of fin on or at base of fin inside mouth in gill chamber beneath eye beneath eye beneath eye beneath eye beneath eye beneath eye beneath eye inside mouth inside mouth beneath eye 2 between eyes, d and immature beneath eye in gill chamber on or at base of fin inside mouth in gill chamber on at base of fin inside mouth inside mouth inside mouth inside mouth on or at base of fin on or at base of fin 2 between eyes, imma- ture beneath eye inside mouth inside mouth
Agarna cumulus
173
Fish host
Cymothoid parasite
Orthopristis ruber
Anilocra haemuli
Site of attachment
beneath eye
Paranthias furcifer Anilocra haemuli beneath eye Phaeoptyx conklini Mothocya bohlkeorum in gill chamber Phaeoptyx pigmentaria Mothocya bohlkeorum in gill chamber
Pogonias cromis Pomacentrus partitus Priacanthus arenatus Scomberomorus cavalla Scomberomorus maculatus Scomberomorus regalis Selar crumenophthalmus Serranus tigrinus
Sphoeroides maculatus Synodus foetens
Nerocila acuminata Anilocra partiti Cymothoa oestrum Lironeca redmanni Lironeca redmanni Lironeca redmanni Cymothoa oestrum Renocila bowmani Renocila waldneri Nerocila acuminata Cymothoa excisa
on or at base of fin
beneath eye inside nouth
in gill chamber in gill chamber in gill chamber inside mouth next to dorsal fin next to dorsal fin
on or at base of fin
inside mouth
Agarna Schioedte and Meinert, 1883
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 ¢ 18 mm. Eyes present, indistinct. Pereon strongly “humped” dorsally. Uropod about '/3 length of pleotelson; uropodal exopod slightly longer, and twice width of endopod. Pleotelson triangular, length */s basal width, apex rounded.
RECORDS No host recorded: Key West, Florida.
174 FLABELLIFERA * CYMOTHOIDAE
Figure 79. Agarna cumulus: A, 2, dorsal view; B, °, 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 | 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.
Anilocra abudefdufi 175
Key to species of Anilocra
1. Pereopods 2—4 with swelling on outer margin of dactylus ........... 2 Pereopods 2—4 lacking swelling on outer margin of dactylus ........ 5 BeBody axis distorted by more than 10° .. 2222508 ea ae. holacanthi Paolveasisdistarned by lessthano vii sao: os eee 3 Be actylus of percopod 7 longer than propodus .:.......'......+.. partiti Mactylus of percopod.7 shorter than propodus~).0.5 2.5... <-- 4 4. Posteroventral angle of pereonite 7 overlapping pleonite | only a a ee Ce cyte Pr oe raid catia ede agora ae (epee abudefdufi Posteroventral angle of pereonite 7 overlapping pleonites 1 and 2 sna eer Sk ees teat a7o ese, uausiths ale a Giant chaetodontis a rosteroventral angle of pereonite 7 produced ........:). 0.0 c00.505+ 6 Eestcroventral angle of pereonite 7 not produced .............:.5... 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 ............. acanthun Uropod not reaching posterior margin of pleotelson ........... chromis
Anilocra abudefdufi Williams and Williams, 1981 Figure 80A—C
DIAGNOSIS Ovigerous 2 19.0-31.0 mm, ¢ 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 2 when attached to host is dark brown; rest of dorsal surface light brown to yellow. Attaching beneath eye of host.
RECORDS Sergeant major Abudefduf saxatilis: Panama; Colombia.
176 FLABELLIFERA * CYMOTHOIDAE
Figure 80. Anilocra abudefdufi: A, 2, 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 ¢ 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 2 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 haemuli 177
Anilocra chaetodontis Williams and Williams, 1981 Figure 80E
DIAGNOSIS Ovigerous 2? 18-28 mm, d 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 ? 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 2 16-28 mm, d 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 2 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 2? 21-40 mm, d 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 ¢ yellow to light brown. Attaching beneath eye of host.
178 FLABELLIFERA * CYMOTHOIDAE
Figure 81. Anilocra haemuli: A, 2, dorsal view; B, ¢, lateral view. Anilocra holacanthi: C, 2. Anilocra holocentri: D, 2°. Anilocra myripristis: E, pleotelson and uropods. Anilocra partiti: F, 2; G, pereopod 7.
RECORDS French grunt Haemulon flavolineatum: 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 fulvus: 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 furcifer: Dominican Republic; Puerto Rico; Colombia. No host recorded: Cuba; Jamaica; Dominica; Barbados; Venezuela; Brazil.
Anilocra holacanthi Williams and Williams, 1981 Figure 81C
DIAGNOSIS Ovigerous 2 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 2 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 2 32—46 mm, ¢ 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 2 dark brown, ventral surface light brown. @ attaching between eyes of host; d 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 2 29-40 mm, ¢ 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 2 light reddish brown, ventral surface yellow. 2 attaching between eyes of host; immature or transitional forms sometimes beneath eye.
RECORDS Blackbar soldierfish Myripristis jacobus: Bahamas; Dominican Re- public; Puerto Rico.
Anilocra partiti Williams and Williams, 1981 Figure 81F,G
DIAGNOSIS Ovigerous 2 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 partitus: 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 d of some species.
Ceratothoa deplanata Bovallius, 1885 Figure 82A
DIAGNOSIS 2 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 FLABELLIFERA * CYMOTHOIDAE
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
on lesseyesior tracest@l eyes present (ass 25. 52a oe, oem ete 2 Anterolateral angles of pereonite 1 broad, reaching to anterior margin
of cephalons-eyesvalsemt: ii. /e ca ate path cena ie ou sere Sofas 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 ¢ 17 mm, d 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 2 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 Synodus foetens: 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 2 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- racao. 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.
184 FLABELLIFERA * CYMOTHOIDAE
Key to species of Glossobius
1. Coxal plates of pereonites 1 and 2 anteroventrally protruding impressus Coxal plates of pereonites 1 and 2 close to body, not protruding a Nin a en tee aac apa er cae NOME So cat sr eer a, A aes hemirhamphi
Glossobius hemiramphi Williams and Williams, 1985a Figure 83A
DIAGNOSIS Ovigerous ¢ 27 mm. Eyes small but distinct. Cephalon pointed anteriorly. Fused coxa of pereonite | 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 ¢ 33 mm. Eyes small but distinct. Cephalon rounded anteriorly. Fused coxal plate of pereonite | 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 | not trisinuate. Number of articles in antennules and antennae
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 d. Pleonites dor- sally strongly convex, not immersed in pereonite 7.
186 FLABELLIFERA * CYMOTHOIDAE
Kuna insularis (Williams and Williams, 1985b) Figure 83C
DIAGNOSIS Ovigerous 2? 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 copulatory stylet present on pleopods 1-3 in 6. Pleotelson basally broader than long, pos- terior margin broadly rounded.
RECORDS’ Sergeant major Abudefduf saxatilis: Carrie Bow Cay, Belize; Cura- cao; 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 Gd, carinae present or absent in 2. 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
iinInIerSed Nin pErE OM: 2s eee << eee eles sibs ameter gel eae e redmannt Uropodal endopod about three times longer than wide; pleon barely iIMmMersed ul. PerEOn wt 5.4 oie aes one eee ee tenuistylis
Lironeca redmanni Leach, 1818 Figure 83D
DIAGNOSIS Ovigerous ¢ 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 regalis, Puerto Rico; king mackerel Scomberomorus cavalla, Colombia; Gerres rho- mbeus, Panama; spot Leiostomus xanthurus, Gulf of Mexico.
Brazil.
Lironeca tenuistylis (Richardson, 1912b) Figure 83E
DIAGNOSIS 2 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; plentelsonisulsrectamelan) 22 2. fe c.2' oe deo age os wns an bohlkeorum Cephalon anteriorly broad, deeply immersed in pereonite 1; pleotelson SIOE RIAN UIA ese cre he ee TGCS 2 ci Sige g hs ayers | waar @ see win nana
188 FLABELLIFERA * CYMOTHOIDAE
Mothocya bohlkeorum Williams and Williams, 1982 Figure 84B
DIAGNOSIS Ovigerous 2 7.6—8.5 mm, d¢ 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. ¢ 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 2 11.0-17.0 mm, d 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; ? lacking these spines. Pleon not immersed in pereonite 7. Pleonites subequal in length; pleonites 1 and 2 usually produced posterolaterally. Pleopods typically with small lamellar accessory gills; pleopods 3—5 folded into deep pockets or pleats. Uropods usually extending beyond pleotelsonic apex.
190 FLABELLIFERA * CYMOTHOIDAE
Nerocila acuminata Schioedte and Meinert, 1881
DIAGNOSIS Ovigerous 2 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 Arius felis: 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 colini 191
Renocila Miers, 1880
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 ...... Zs
2. Dorsal surface of body brown; posteroventral angle of pereonite 7
heaemine pleamite 6 Et chse a oc ed iaje a hint seopeete eta su0 wi/acas waldneri Dorsal surface of body black; posteroventral angle of pereonite 7 REACHING PICOMILEY Oph 2.56 or epee so Se es el Stn tonsa Bg enn edt a mies bowmani
Renocila bowmani Williams and Williams, 1980 Figure 85A
DIAGNOSIS 2 18.0 mm, d 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 3/1 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 2 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
192 FLABELLIFERA * CYMOTHOIDAE
Figure 85. A, Renocila bowmani. Renocila colini: B, 2; C, 3. D, Renocila waldneri.
of exopod. Pleotelson '/7 to '/2 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 ¢ LIMNORIIDAE 193
RECORDS Flamefish Apogon maculatus: Puerto Rico. Belted cardinalfish Ap- ogon townsendi: Puerto Rico.
Renocila waldneri Williams and Williams, 1980 Figure 85D
DIAGNOSIS Ovigerous 2 15.3-19.3 mm, d 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
PUropodal rami very unequal 7. 02022 22 22 tees Seek se oe aes 2 Mropodal rami-subedqtal fet 5 Sas Sh OS oa 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 3.0mm, ovigerous 2 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.
Limnoria insulae 195
Key to species of Limnoria
1. Dorsal surface of pleotelson lacking prominent tubercles, ridges, or carinae (L. simulata may appear to lack ornamentation; in this Species thie tubercles|are very smiall)ri owes ob 1s cbs ee onto om 2 2
Dorsal surface of pleotelson bearing tubercles, ridges, or carinae .... 3
2. Pleotelson flat; pleonite 5 with broadly rounded middorsal ridge
4 cpg Gatch a ie ROP Cue Ro RE RS a a platycauda Pleotelson cup shaped; pleonite 5 with strong narrowly rounded
TINIE EE O/RSE1| Fi) oF Sc oa care are ae aR insulae
3. Pleotelson with basal tubercles but lacking ridges ................. 4
Pleotelson with ridges but lacking freestanding tubercles ........... 7
4. ¢ pleotelson with single strong middorsal tubercle .......... unicornis
Pleotelson with more than one basal tubercle .......4...¢..22s%... 5
5 Pleoteisom with: three basal tubercles: ~ 4). s2 66s Obes sok tuberculata
Pleotelson with more than three basal tubercles ................... 6
6. Pleotelson with four basal tubercles in line (difficult to detect) simulata
aupleoteison with six, basal’ tubercles 2208 aieety 5 ee ee ake indica 7. Pleotelson with single middorsal longitudinal ridge ...... multipunctata Plecotelsonswithytwo rounded: basal ridges) 7.22 ye:-/..0 62. syed eis Seale 8 oe Pleonite) with strong. V-shaped ridge 23... scajed 6 c.c ae os 2 hens pfefferi Pleonite 5 with two posteriorly converging ridges .......... saseboensis
Limnoria insulae Menzies, 1957 Figure 86C
DIAGNOSIS’ d6 3.0 mm, ovigerous ¢ 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. Fiji; Guam; Palmyra Island; Caroline Islands.
196 FLABELLIFERA * LIMNORIIDAE
Figure 86. Limnoria indica: A, pleotelson, 3; B, pleotelson, 2. Limnoria insulae: C, pleotelson. Limnoria multipunctata: D, pleotelson in oblique-lateral view.
Limnoria multipunctata Menzies, 1957 Figures 86D; 87A
DIAGNOSIS 6 2.8 mm, ovigerous 2 3.0 mm. Pleonite 5 dorsally smooth. Pleotelson with middorsal longitudinal rounded ridge bearing several
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.
button-shaped tubercles in posterior half; posterior margin and lateral crests
tuberculate.
RECORDS Puerto Rico; Jamaica; Twin Cays, Belize. Japan; Kai Islands, South Pacific.
198 FLABELLIFERA * LIMNORIIDAE
Limnoria pfefferi Stebbing, 1904 Figure 87B
DIAGNOSIS d 3.8 mm, ovigerous ¢ 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 2 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 Curacao; Cozumel, Mexico; Twin Cays and Man o’War Cay, Belize. Aldabra Atoll, Indian Ocean.
Limnoria saseboensis Menzies, 1957 Figure 87D
DIAGNOSIS <6 3.5mm. 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 2 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.
Paralimnoria andrewsi 199
Limnoria tuberculata Sowinsky, 1884 Figure 87F
DIAGNOSIS <6 2.8 mm, ovigerous 2 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, 195la.
Limnoria unicornis Menzies, 1957 Figure 88A,B
DIAGNOSIS d 2.6 mm, ovigerous 2 2.6 mm. Mandibular palp of one arti- cle. Pleonite 5 with somewhat obscure Y-shaped ridge middorsally. Pleotelson in d 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 (Calman, 1910) Figure 88C,D
DIAGNOSIS 6 2.6 mm, 2 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.
200 FLABELLIFERA * LIMNORIIDAE
Figure 88. Limnoria unicornis: A, pleotelson, d; B, pleon, d, in lateral view. Paralimnoria andrewsi: C’, pleonite 5 and pleotelson; D, uropod. Phycolimnoria clarkae: E, pleonite 5 and pleotelson; F, uropod and pleotelson in lateral view.
RECORDS Florida Keys; Puerto Rico; Twin Cays, Belize; Curacao. Christmas Islands, Indian Ocean; Samoa; Hawaii; Japan.
REMARKS Menzies (1957) discusses three forms of this species: Forma typ- ica, which lacks a central depressed area dorsally on pleonite 5 and has a pair of submedian obscurely carinate tubercles on the pleotelson; Forma A, which has a triangular depressed area dorsally on pleonite 5 and a pair of subme-
FLABELLIFERA * SEROLIDAE 201
dian tubercles supported by strong carinae on the pleotelson; Forma B, 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 2 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 | in d and ¢ subchelate, pereopod 2 subchelate or ambulatory in 6, ambulatory in @. 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 d, ambulatory in 2. Pleopods 1—3, peduncles elon- gate, rami subelliptical. Pleopod 3, exopod uniarticulate.
Serolis mgrayi Menzies and Frankenberg, 1966 Figure 89
DIAGNOSIS 6 4.5 mm, ovigerous 2 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 2. 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, 3; B, pereopod 1; C, pereopod 2, d.
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 FLABELLIFERA * 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 Tecticipitinae, 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 ¢
SR o Be tiers MRNA aye elena SeaUd oe rene ck aeRO iy cichasy Pai a ascent Ancininae
Pereopods: | and 2 ambulatory, si 5.2 den os fa es ee eee 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
Sess. ctaNe, Seta ia Gita ede on aN Ratha tine 0 reo. ie ge eae Sphaeromatinae
Subfamily Ancininae Tattersall, 1905
DIAGNOSIS Body markedly dorsoventrally depressed. Cephalon fused me- dially with pereonite 1. Pereopod 1 prehensile in d and 2. Pereopod 2 pre- hensile in 6 only. Pleopods 4 and 5 similar, lacking branchial pleats. Uropods uniramous.
Ancinus belizensis 205
anterior pocket
oostegite posterior pocket
internal pouch
Figure 90. Diagrammatic representation of 2 sphaeromatid, showing marsupial structures (adapted from Harrison, 1984).
Ancinus H. Milne Edwards, 1840
DIAGNOSIS Eyes dorsal. 2 mouthparts not metamorphosed. Mandibular molar absent; palp of three articles. Maxilla | of single ramus, endite rudi- mentary. Maxilla 2 of two rami. 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 | 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 Bt ce OOo en ch soa Ba coca ila: alga int ke Ta a A One en belizensis
Ancinus belizensis Kensley and Schotte, 1987 Figure 91A—C
DIAGNOSIS 6 4.1 mm, ? 2.8 mm. Body oval, about twice longer than wide. Dorsal integument strongly pitted. Antennular flagellum of 12 articles; an-
206 FLABELLIFERA * SPHAEROMATIDAE
Figure 91. Ancinus belizensis: A, 2; B, pereopod 1 3; C, pereopod 2 ¢. Ancinus braziliensis: D, adult (from Glynn and Glynn, 1974).
tennal flagellum of 10 articles. d 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 6 7.0mm, 2 6.0 mm. Body about twice longer than wide. Dor- sal integument smooth. Antennular flagellum of 17 articles; antennal flagellum of 10 articles. d pereopod 2, dactylus strongly curved, reaching to midlength of posterior margin of carpus. Pleopod 2 almost three times longer than basal width.
Cassidinidea 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 | 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 of 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
208 FLABELLIFERA * SPHAEROMATIDAE
well developed, fused with sympod; exopod markedly reduced. Penial rami elongate, separate. 2 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
«Posterior mareginvef pleotelsoni truncate’ tic... use }. sees ae ovalis Postenior marginvef pleotelson rounded: . :.o21.2..24e sass. ap neer mosaica
Cassidinidea mosaica Kensley and Schotte, 1987 Figure 92A
DIAGNOSIS. 6 1.8 mm, ovigerous 2 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 2 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 1%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- eonites, pleonite 1, and uropods. Expanded basal articles of antennules con-
Paraleptosphaeroma 209
Figure 92. Cassidinidea mosaica: A, 3. Cassidinidea ovalis: B, 3; C, pereopod 1; D, pleopod 4; E, pleopod 5. Paraleptosphaeroma glynni: F, ¢.
tiguous in midline. Single articulated pleonite with short free lateral margin.
Uropodal sympod and endopod fused; exopod articulated, much shorter than fused endopod.
210 FLABELLIFERA * SPHAEROMATIDAE
Paraleptosphaeroma glynni Buss and Iverson, 1981 Figure 92F
DIAGNOSIS 6 2.58 mm, ovigerous ¢ 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 im ‘both sexes) 4. .c26.07 Gh oe yo ea 2
Pleotelson showing marked sexual dimorphism ..................-. 3
2. Cephalon and pleotelson smooth, lacking ridges .......... Ischyromene
Pleotelson and cephalon with ridges: 2.3... 0h ee oe ce Cerceis
3. JWrepods lamellar ‘both; sexes-4..4).\2)2.ta 10-4 pete seers <= or eee ee + Uropods lamellar in ¢, endopod reduced, exopod elongate-cylindrical
UNS Geo ese 4B esaia: Des oe lado eS eal Na, oS Ree ee 5
4. Ovigerous @ lacking oostegites; 6, pleopod 2 copulatory stylet basally broad, distally tapering, extending to or beyond ramus . . Dynamenella Ovigerous 2 with one pair of oostegites on pereonite 4; 6, pleopod 2 copulatory stylet narrow, extending well beyond ramus ... . Paradella
5. 6, strong median lobe in pleotelsonic notch reaching well beyond
margin; ovigerous 2 with three pairs of oostegites ........ Discerceis 3, pleotelsonic notch with short median lobe, if present; ovigerous 2 with three or four, pairs, of costegites’ .7. 7... .: 0-4 eee 6
6. Ovigerous ¢ with three pairs of oostegites; d pleotelsonic notch lacking “marginal teeth ‘or median lobes =.) <= ee ae Geocerceis Ovigerous ¢ with four pairs of oostegites; d pleotelsonic notch with marginal teeth and/or median lobe’..-% 002. see cet eee Paracerceis
Discerceis lal
Subfamily Dynameninae Bowman, 1981
DIAGNOSIS Cephalon not fused with pereonite 1. Pereopods | 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 d.
Uropods biramous.
Cerceis H. Milne Edwards, 1840
DIAGNOSIS Mouthparts metamorphosed in ¢. Broodpouch of four pairs of oostegites on pereonites 1—4, overlapping in midline. Brood held in four pairs of internal pouches. Pockets absent.
“Cerceis” carinata Glynn, 1970 Figure 93A
DIAGNOSIS 6 3.8 mm, @ 3.9 mm. Dorsal integument, especially posteri- orly, finely pustulose. Cephalon with three pairs of rounded dorsolateral carinae, one middorsal carina, not reaching posterior margin. Pereonite | with two ventrolateral carinae. Pleopod 3, exopod biarticulate. Pleotelson similar in d and @. Pleotelson with basal middorsal inflated area flanked by two smaller lateral swellings, with carina in midline almost reaching pos- terior margin; posterolateral margins converging to narrow, slightly concave posterior margin. d: Penes elongate, basally fused. Pleopod 2, copulatory stylet basally broad, tapering distally, articulating mediodistally on endo- pod; exopod with three enlarged distal plumose setae.
RECORDS’ Venezuela, 5—7 m.
REMARKS Several differences (in the male penial structure, copulatory stylet, antennular peduncle, and pleonal sutures) between Cerceis carinata and the definition of the genus (Harrison and Holdich, 1982; Harrison, 1984) indicate that this species has not been placed in the correct genus. Until fresh ovigerous females and mature males are available, the generic position must remain uncertain.
Discerceis Richardson, 1905
DIAGNOSIS Mouthparts in 2 metamorphosed. Broodpouch formed by four pairs of oostegites on pereonites 1—4, overlapping in midline. Brood held in
212 FLABELLIFERA * SPHAEROMATIDAE
Figure 93. “Cerceis” carinata: A, 3. Discerceis linguicauda: B, 3. Dynamenella acutitelson: C, 2; D, pleon 3. Dynamenella angulata: E, @.
internal pouches (number unknown). Pockets absent. d with uropodal endo- pod and sympod fused, very short; exopod elongate, cylindrical.
Dynamenella 213
Discerceis linguicauda (Richardson, 1901) Figure 93B
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 | 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. 2: 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
fers with pleotelsonic foramen) «)..2.. 0205.28.05 ene enn eee eee ees 2 3 lacking foramen but with notch, or appearing entire; 2 pleotelson UALR AIOLEM VISMONC 2 dioica iis soe are see ss s,s ewes se acutitelson
2. 3 with four strong pleotelsonic ridges; 2 with subcircular pleotelsonic fondimensiaen er iikimonise anes: bos cies Es bile ed ee quadrilirata 6 lacking pleotelsonic ridges; ¢ with posterior margin of pleotelson Suite teas Ar Bhs RRS te ies ad eek Ecards perforata
214 FLABELLIFERA *« SPHAEROMATIDAE
groove, or foramen. d: 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 2. 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 3.5mm, ? 2.3 mm. d: 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 either just 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 ¢ypica and their absence in glabrothorax. 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 Jschyromene. Again, until further mature males and ovigerous females are seen, the generic placement of this species must remain in doubt.
Geocerceis barbarae DS
Dynamenella perforata (Moore, 1901) Figure 94A,B
DIAGNOSIS 6 3.2mm, 2 2.6 mm. d: 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. : 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 granulata; Dominican Republic; Cuba.
Dynamenella quadrilirata Kensley, 1984 Figure 94C—H
DIAGNOSIS 6 2.6mm, 2 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. 2: 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 2 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. d 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 6 3.3mm, 2 2.5 mm. Pleopod 3 exopod of single article. Pleon with two elongate sutures reaching lateral pleon margin. d: Frontal lamina expanded into ventrally directed beaklike process. Penes separate, relatively
216 FLABELLIFERA * SPHAEROMATIDAE
Figure 94. Dynamenella perforata: A, 3; B, pleon 2. Dynamenella quadrilirata: C, 3; D, pleon ¢; E, pleon 2, lateral view; F, pleopod 2 3; G, pleopod 4; H, pleopod 5.
elongate. Pleonite 5 with three dorsal tubercles near posterior margin. Pleotelson with raised anterocentral area having two lateral longitudinal rounded ridges; apex notched. 2: Uropodal exopod and endopod subequal, lamellar. Pleotelson as in ¢. Frontal lamina not produced as in d.
RECORDS Puerto Rico, intertidal to 3 m, in coral rubble.
Ischyromene aT
Figure 95. Geocerceis barbarae: A, 3; B, pleon 2; C, 6, lateral view. Ischyromene barnardi: D, o.
Ischyromene Racovitza, 1908
DIAGNOSIS mouthparts not metamorphosed. Broodpouch of three pairs of oostegites on pereonites 2—4, overlapping in midline. Large posterior pocket covering posterior ventrum, opening anteriorly between fourth per- eopods. Brood housed in ventral body wall. Sexual dimorphism not pro-
218 FLABELLIFERA * SPHAEROMATIDAE
nounced. Uropodal rami lamellar. d 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 4.5mm, ¢ 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. ¢: 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 6, and with shallower median notch lacking teeth.
Key to species of Paracerceis (P. nuttingi not included)
1. d, pleotelsonic notch narrow, with median basal tooth; 2, pleotelson dorsally. unormmamented: 5 «fic canis idare nto ee ee ee ee 2
3, pleotelsonic notch wide, with lateral teeth; 2, pleotelson dorsally with: tubercles: 2.5.<25te vod.c Sal sicie oie elas Se eee 3
2. 3, median tooth of pleotelsonic notch almost as long as notch; subacute median tubercle on anterior pleotelson; 2, pleotelson with posterior margin faintly concave, not notched .............. edithae
3, median tooth less than half length of notch; pleotelson with blunt rounded median tubercle; 2, pleotelson with distinct posterior notch
88 DUE A Sie GEO a Se ee ee glynni
Paracerceis caudata 219
3. 3, pleotelsonic notch deep, margins usually with two teeth on each side; strong median tubercle on anterior pleotelson bluntly bifid; @, pleotelson with one or two rounded median tubercles and 2 smaller BUI RMCS EON Gd CI USTCMED Nyko) sete ire lais con dhe oan doesn 2 egahees caudata
3, pleotelsonic notch shallow, with tiny lateral denticles; median tubercle of pleotelson conical, acute; °, pleotelson with three large conical acute tubercles and several smaller scattered tubercles in CANA MACHTMEN AIL on tus eles ok con recM Maes oe so sees aks SACP See Ee cohenae
Paracerceis caudata (Say, 1818) Figure 96
DIAGNOSIS. 6 8.1 mm, 2 6.4 mm. ¢: 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. 2: 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.
220
FLABELLIFERA * SPHAEROMATIDAE
Ni
Al IME Ay, ly
Figure 96. Paracerceis caudata: A, 3; B, pleon 2; C, mandible 6; D, maxilla 1 6; E, maxilla 2 ¢d; F, maxilliped d; G, mandible 2; H, maxilla 1 2; J, maxilla 1 2; J,
maxilliped °.
Paracerceis cohenae Kensley, 1984
Figure 97A,B DIAGNOSIS 6 10.0 mm, 2 7.9mm. d: 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. @: Per- eon and pleon much as in 4, 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, 2 3.1 mm. d: 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. 2: Integument much less tuberculate-spinose than in d. Imma- ture 2, posterior margin of pleotelson with faintly rounded median lobe. In mature @, 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 6.4mm, 2 5.2 mm. d: 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. 2: Body far less setose and tuberculate than d. Pleotelson with strongly inflated anterior area having very faint mid- dorsal tubercle; notch well marked, formed by triangular lobes of
222 FLABELLIFERA * SPHAEROMATIDAE
Figure 97. Paracerceis cohenae: A, 3; B, pleotelson, 2. Paracerceis edithae: C, pleotelson, 2; D, mature 6; E, immature 6. Paracerceis glynni: F, 3; G, pleotelson,
2. Paracerceis nuttingi: H, &.
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
Paradella DS
m, from green alga Halimeda sp. on forereef, and from sponge Aphysina fistularis.
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 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 Both sexes lacking processes on pereon and pleon. Marked sex- 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 Dita distincelygoramulate os 203 3. aac tie can: aes es OR oes So OM Ohl eas Za PICGUN AGN PI EOLe ISON SIMOOUA. 2 2c ac. <n Spskus (rn ee Re, a Ss. - 3
2. 3d with pleotelsonic foramen distinctly heart shaped, with median point; four submedian tubercles of pleotelson in d and 2 somewhat elongate; 2 pleotelson posteriorly narrowed, slit visible dorsally
3 with pleotelsonic foramen wider than long, but lacking median point; four submedian tubercles of pleotelson in d small, rounded, obscure in 2; @ pleotelson posteriorly truncate, slit not visible (SPOT gS | | Pee ae UE OE ge ea ORR: Aieee 1 27 Sapo eta plicatura
3. Tubercles on pleotelson in d and ¢ small to obscure; d with plcotelsomic foramen subcircular 2.665.055.0605 00s oan quadripunctata Tubercles on pleotelson broadly rounded mounds in d and 2; 6 with pleotelsonic foramen wider than long ................. tumidicauda
224 FLABELLIFERA * SPHAEROMATIDAE
exopod with articulation. Uropodal rami lamellar. 2: 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. d: Penes long, basally fused. Copulatory stylet basally narrow, extending further be- yond endopod than in Dynamenella. Uropods broader than in &.
Paradella dianae (Menzies, 1962b) Figure 98A—C
DIAGNOSIS 6 3.4mm, ovigerous 2 3.7 mm. d: Pereonite 7, posterior mar- gin broadly bilobed. Pleonite 5 with two rounded submedian tubercles. Pleotelsonic foramen distinctly heart shaped with median point. Pleotelson with four submedian and two lateral tubercles in d and ¢, plus median tubercle at base of foramen in d; tubercles tending to be elongate and sub- carinate. Uropodal rami lamellar, margins finely crenulate, relatively broader than in @. @: Pleotelsonic slit wide, dorsally visible.
RECORDS Key West, Florida; Puerto Rico, intertidal. Baja California, intertidal.
Paradella plicatura (Glynn, 1970) Figure 98D,E
DIAGNOSIS 6 4.1 mm, ¢ 3.6 mm. Pleon and pleotelson with tiny scattered tubercles. 6: Pereonites 5—7 with posteriorly directed flanges, that of per- eonite 7 largest, bilobed. Pleotelson with four submedian and two lateral discreet rounded tubercles. Pleotelsonic foramen wider than long, with basal bulge but lacking median point. 2: Pleotelson with wide posterior slit not visible dorsally, posterior margin appearing truncate. Pleotelsonic tubercles less marked than in 6.
RECORDS Jamaica, under red mangroves; Margarita Island, Venezuela, shallow infratidal.
Paradella quadripunctata (Menzies and Glynn, 1968) Figure 98F,G
DIAGNOSIS 6 2.5mm, 2 2.5 mm. Pleonite 5 with two low rounded subme- dian tubercles. Pleotelson with four low submedian rounded tubercles and
Paradella quadripunctata 225
Figure 98. Paradella dianae: A, 3; B, pleopod 2 3; C, pleon 2. Paradella plicatura: D, 3; E, pleon 2. Paradella quadripunctata: F, 3; G, pleon 2. Paradella tumidicauda: H, pleon @ (from Glynn, 1970); J, 6.
four smaller lateral tubercles. ¢: Pleotelsonic foramen subcircular, ventral margins of foramenal tube barely touching. : Pleotelson posteriorly nar- rowly tapered, slit becoming tubelike, dorsally visible; four submedian tuber- cles less marked than in 6.
RECORDS Bermuda; Florida; Dominican Republic; Puerto Rico, intertidal; U.S. Virgin Islands, intertidal to 1 m.
226 FLABELLIFERA * SPHAEROMATIDAE
Paradella tumidicauda (Glynn, 1970) Figure 98H,I
DIAGNOSIS 6 6.7 mm, 2 6.5 mm. <d: 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. 2: 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 ... 25. se o2 Sphaeroma Uropodal exopod with outer margin entire or faintly crenulate ...... 2
2. 3, pleotelsonic notch with median lobe; 2, pleotelsonic apex barely notched with rounded median lobe; 2, mouthparts metamorphosed
alate Mate CIS ee NE EOE MIA SO Loe eee at eer ay eee 3
Pleotelson entire to very faintly notched in d and 2; 2, mouthparts not metamorphosed; with three pairs of oostegites ..... Exosphaeroma
3. Mature 6, uropodal exopod about twice length of endopod; 2 with three pairs OhOOSteEsItEs :. soca c ee tee nae ween eae Harrieta
3, uropodal rami subequal or exopod shorter than endopod; ? with {Our Pairs OL GOSTEPITES \...5-5.8 uc step oasian ce cee sortase Cymodoce
Cymodoce Leach, 1814
DIAGNOSIS Pleon with two elongate straight parallel incomplete sutures on each side. Pleotelsonic apex with marked notch bearing median tooth.
Cymodoce ruetzleri PT,
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. d: Maxillipedal palp articles 2—4 bearing setigerous lobes. Penial rami elongate, separate. Pleon usually more tuberculate than in 2. Uropodal exopod lamel- lar, shorter than endopod. 2: 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 2 7.5mm. 2: 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 d¢ 5.0 mm, 2 4.2 mm. ¢: 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. 2: Pleotelson with two conical apically acute tubercles, apex barely notched, with short rounded lobe slightly offset from posterior margin. Both
228 FLABELLIFERA * SPHAEROMATIDAE
Figure 99. “Cymodoce” barrerae: A, 2; B, frontal lamina. Cymodoce ruetzleri: C, 3; D, pleon in lateral view, 6; E, pleon @; F, 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.
Exosphaeroma alba 229
Exosphaeroma Stebbing, 1900
DIAGNOSIS Maxillipedal palp articles 2—4 produced medially into lobes. Pereonites 6 and 7 dorsally unarmed. Pleopod 3, exopod biarticulate. d: Penes short, separate. Copulatory stylet of pleopod 2 elongate, slender. Pleotelson lacking strong apical notch. 2: 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
See oe tea aeRO STING Me cc ane hn oP ake terest productatelson
3. Pleotelson with posterior margin faintly tnlobed, and with three low rounaged, tubercles anteriorly... 2. h.5. slosh sx teers soo yucatanum Pleotelson with posterior margin faintly notched .................. 4
4. Pleotelson posteriorly broadly notched; two rounded submedian tubercles;on' inflated midregion® = 2:..0. 2.062 22 Os Peed antillense Pleotelson with faint narrow notch posteriorly; lacking dorsal tubercles
OU A Sa 2 > RR a ER ORS oe A ME 7a ES OS LOU EE CS.» =. ee ROE alba
Exosphaeroma alba Menzies and Glynn, 1968 Figure 100A—C
DIAGNOSIS 6 2.0 mm, 2 2.3 mm. Frontal lamina anteriorly broadly rounded, basally slightly wider than midlength. Pleotelson similar in d and 2; 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.
230 FLABELLIFERA * SPHAEROMATIDAE
Figure 100. Exosphaeroma alba: A; B, frontal lamina; C, uropod. Exosphaeroma antillense: D; E, frontal lamina; F, uropod. Exosphaeroma diminuta: G; H, frontal lamina; J, uropod. Exosphaeroma productatelson: J; K, frontal lamina; L, uropod. Exosphaeroma yucatanum: M, pleon (from Richardson, 1905).
Exosphaeroma antillense Richardson, 1912d Figure 100D,F
DIAGNOSIS @ 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 <6 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 2.5mm, ¢ 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.
232 FLABELLIFERA * SPHAEROMATIDAE
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 ¢ 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 2, exopod twice length of endopod and oval in cross section in 6. 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 101A,B
DIAGNOSIS 6 6.0 mm, ¢ 6.5 mm. G: Frontal lamina with broad slightly convex anterior margin. T'wo 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. ¢: Essentially similar to d, 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 d and 2. Pleopod 3, exopod uniarticulate. Uropodal ex- opod with outer margin serrate. Able to conglobate. ¢d: Penes short, rounded, separate. Pleopod 2, copulatory stylet articulating basally on endo- pod, slender, reaching well beyond rami. 2: Mouthparts not meta-
Figure 101. Harrieta faxoni: A, 3 ; B, pleon 2. C, Sphaeroma quadridentata; D, Sphaeroma terebrans; E, Sphaeroma walkeri.
234 FLABELLIFERA * SPHAEROMATIDAE
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 Fo MS MSS Ae bos He ed re mae ecto rent eaatate ete nate, aaa ca eS terebrans Pleotelson: posteriorly broadly rounded .).2- 5a o5-. coe ie oe 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, ¢ 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 101D
DIAGNOSIS. 6 10.0 mm, @ 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 §. 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 9.5mm, 2? 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 ¢ than in 2. Frontal lamina narrow, pentagonal. Antennular 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 6 9.5 mm, ovigerous $ 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 ¢ greatly enlarged, projecting anteriorly from cephalon, not used in feeding. Mandibles lacking in 2. 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. 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, 2; B, pereopod 1; C, maxilla 1; D, mandible; E, 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 (d 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’ -teethi tr, 202 aig oe Bae ee Te, ee ee 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 ..............--- a
4. Lobe of outer margin of mandible notched; pereonite 5 twice wider
thanumiddorsal lengthy 26.2024 2s ots es oes tos sae ee care velosa
Lobe of outer margin of mandible rounded; pereonite 5 1.5 times wider
thanmiddorsallength On 2 snc... role ots Se ene oo ae 6
5. Anterior margin of cephalon with distinct medial process ..... virginalis Anterior margin of cephalon barely convex, lacking medial process
SSE Ate SR tO Se Ny Ein? Cee eS rathi
6. Inner proximal lobe of mandible distinct ................5---00: 7
Inner proximal lobe of mandible indistinct ............--- samariensis
7. Inner proximal lobe of mandible entire ..............+++--+s220+:: 8 Inner proximal lobe of mandible with rounded toothlike marginal
SEMUECLUROS. |) 50 Arta. 2 aie coy encire a eps heen ann eee se oer Kee og ren aaa johanna
$.!Pereonites.3—5 poorly defined 2.6). ).c ovata weine : poe puertoricensis
Pereonites.3—5 clearly detined rai Aisi, bas aise Sake ee magdalenensis
9. Anterior margin of cephalon concave, lacking projections ..... gonzalezi
Anterior margin of cephalon with four projections .......... beethoveni
Gnathia magdalenensis 239
Gnathia beethovenit Paul and Menzies, 1971 Figure 103A
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 Miller, 1988 Figure 103B
DIAGNOSIS. ¢ 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 6 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 Miller, 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, 5 (after Miller, 1988). Gnathia johanna: C, 3. Gnathia magdalenensis: D, 3 (after Miiller, 1988). Gnathia puertoricensis: E, 3; F, praniza larva; G, ovigerous °.
Gnathia triospathiona 241
Gnathia puertoricensis Menzies and Glynn, 1968 Figure 103E—G
DIAGNOSIS <6 3.0 mm, ovigerous 2 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 104A
DIAGNOSIS 6 1.9 mm, ovigerous 2 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 Miller, 1988 Figure 104B
DIAGNOSIS 6 2.0 mm. Anterior margin of cephalon with three tubercles, 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, 3 (after Miller, 1988); Gnathia triospathiona: C, 3; Gnathia virginalis: D, 3; Gnathia velosa: E, 3 (after Miiller, 1988).
MICROCERBERIDEA * MIGROCERBERIDAE 243
Gnathia velosa Miller, 1988 Figure 104E
DIAGNOSIS. ¢ 1.5 mm. Anterior margin of cephalon with three tubercles, 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
DIAGNOSIS. 6 2.2 mm. Anterior margin of cephalon with three tubercles, 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 | subchelate; pereopods 2—7 ambulatory. Pleon of two free pleonites plus pleotelson. Pleopod | in d 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 microcerberideans 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. Maxillipedal palp articles 2 and 3 enlarged; basis of pereopods lacking SPUMOUS: PLOCESS! s.5. 542 clases eee sto ans tare) wean Sad Yvesia Maxillipedal palp articles 2 and 3 not markedly enlarged; basis of pereopads: with spinous Process, fe. fees bs ee eee Microcerberus
Microcerberus Karaman, 1933b
DIAGNOSIS Maxillipedal 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. 6 1.1 mm, ¢ 1.1 mm. Tergal lobes of pereonites 2—4 rounded. Apical lobe of d 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, 6; B, pereopod 1; C, maxilliped; D, pereopod 2; E, pleopod 2 6. Yuvesia striata (from Coineau and Botosaneanu, 1973): F, maxilliped; G, pereopod 1; H, pereopod 2.
246 MICROCERBERIDEA * ONISCIDEA
Yvesia Coineau and Botosaneanu, 1973
Yvesia striata Coineau and Botosaneanu, 1973 Figure 105F—H
DIAGNOSIS ¢ 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 pereopod | 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; d with pleopod 2, and sometimes pleopod | 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.
ONISCIDEA 247
Key to genera and species of littoral Oniscidea
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 .....'2.5...... 250s. 3 (Tylos) Mropeds visible in dorsal view, 25...2% oo... 2. 2 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 JP: Seep Sa I Oe nO ee PY APR CIRC 2 CAR mee Rana Tylos latreillei
2. Uropodal rami both clongate, subequal .... 206. 66.6020. 6 (Ligia) Wropedal ram very unequal m length 22.2 2. dag ds ieee nt a ade oe 8 6. Propodus of d pereopod | with distal rounded lobe ...... Ligia exotica Propodus of 6 pereopod | lacking rounded lobe .................. ‘| imaAmexval i mleopod 2 club shaped 252.2... 4-0... S24 ee Ligia olfersii mpEXor oc pleapod.? (bile. 7.2. fh. ge shes wads «Te dds Ligia baudiniana eoeantennal fagelum of two articles 254. o¢2626:...--¢ Rhyscotus texensis Antennal flagellum of three articles 2.00.5 cos. 5.5252 he- 9 (Vandeloscia)
9. Endopod of ¢ pleopod | with large scalelike subapical process Ne er eRe href ec eto Ree ds he A Vandeloscia riedli
Endopod of d pleopod | with small scalelike subapical process BN da ig eka), 5) 2s g Pg ac sess eo) Vandeloscia culebrae
Armadilloniscus Ul yanin, 1875
Armadilloniscus ninae Schultz, 1984 Figure 106A
DIAGNOSIS. 6 3.2mm, 2 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, 3 pleopod 2 endopod. Ligia exotica: D, dactylus and propodus of pereopod 1; E, 6 pleopod 2 endopod. Ligia olfersii: F, 3 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 | endopod. N, Vandeloscia riedli.
Rhyscotus texensis 249
Ligia Fabricius, 1798 Ligia baudiniana H. Milne Edwards, 1840 Figure 106B,C
DIAGNOSIS. 6 and 2 up to 22 mm. Antennal flagellum elongate, multiar- ticulate. Apex of d 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 Liga, this species may be seen on rocks 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 2 32.0 mm. Propodus of d pereopod | with rounded lobe on inner distal surface. Apex of d pleopod 2 club shaped,
convoluted.
RECORDS New Jersey to Uruguay; Indo-Pacific.
Ligia olfersii Brandt, 1833 Figure 106F
DIAGNOSIS 6 20.0 mm, ovigerous 2 24.0 mm. Apex of d 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. ¢ and 2 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, @ 13.0 mm. Ventral extensions of pleonite 5 not meeting in midline.
RECORDS’ Bermuda; Cuba; Puerto Rico; Honduras. Mediterranean.
Tylos marcuzzu Soika, 1954 Figure 106]
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. 6 11.0 mm., 2 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; Curagao, 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, 2 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.
VALVIFERA 251
Vandeloscia Roman, 1977 Vandeloscia culebrae (Moore, 1901) Figure 106M
DIAGNOSIS 6 5.0 mm, 2 6.1 mm. Tiny lateral tubercles present on per- eonites. Endopod of pleopod | in d 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 6 5.9 mm, ¢ 6.0 mm. Tiny obsolete tubercles present on all pereonites. Endopod of d 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 OTE Se A, ne ee One a ee ee Arcturidae Body never geniculate; all pereopods ambulatory .......... Idoteidae p 755
252 VALVIFERA * ARCTURIDAE
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 | 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 Jen Sales obo dks ain a aden ee St Pe oe a he Ao Lae) MES Thermarcturus Pereonite 1 fused with cephalon; pleonites fused with pleotelson SN eR EAN SE Bae wa Blk Oma in Me a aati Aa ii acne desk RP Astacilla
Astacilla Cordiner, 1793
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 swith; spines or, tubercles..9. 656 citecye sears “sles 2
2. Pereonite 4 in d and 2 with strong middorsal tubercle; pairs of spines
lacking: On pereonitEs 280... seen oa eon ae oe eee regina Pereonite 4 lacking strong middorsal tubercle; pairs of spines on all
pereonites °./) (G0 cu Vita nn seer teraes oe ne kre ete eee eas lasallae
Astacilla regina 253
pleopod 1 d 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 ¢ 9.0 mm. Body cylindrical, ovigerous ? with pereonite 4 somewhat bulged, d with pereonite 4 elongate-cylindrical. Shallow groove marking fusion between cephalon and pereonite |. 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 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 2 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 * ARCTURIDAE
Figure 107. Astacilla cymodocea: A, 3; B, °. Astacilla lasallae: C, 3. Astacilla regina: D, 6; E, 2; F, pereopod 4; G, pereopod 1. Thermarcturus venezuelensis: H, 2 (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.
VALVIFERA * IDOTEIDAE 255
Thermarcturus Paul and Menzies, 1971
DIAGNOSIS Pereonite | 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 2 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.
RECORDS Off Venezuela, 95 m.
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
antennal Mapenunr mulliariculate: ic. ..6 . 65's cede fan Idotea Antennal flagetlumiclavate es 2... lees ol eee oh Dae Stic 2S 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 sss atk daha Riel APRESS A, WOR OER, OP ER dst Erichsonella
3. Pleon consisting of three complete and one incomplete pleonites plus TENTS Rel TCT CARRS gh Sea Ee en SE a MMC ene Cleantioides Pleon consisting of two complete and two incomplete pleonites plus pleotelsion. @ cj aganst So hiins deen lees ler Beds at 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 ? 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
Ll. .Pereonites with dotsal' spines q. 2o45 Sh chee ee fo Lh ceca soutien ae 2 Pereonitesdacking dorsal ‘Spies! We ule eS ele ite es ssi scan attenuata 2. Pereonites 1—4 with middorsal and lateral spines ............ floridana
Pereonites 1—4 with middorsal spines only .................. filiformis
Erichsonella 257
Figure 108. A, Cleantioides planicauda; B, Erichsonella attenuata 3; C, Erichsonella filiformis 3; D, Erichsonella floridana 2; E, Idotea balthica 3; F, Idotea metallica 2; G, Miratidotea bruscai 2.
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.
258 VALVIFERA * IDOTEIDAE
Erichsonella attenuata (Harger, 1873) Figure 108B
DIAGNOSIS 6 11.4 mm, ovigerous 2 12.0 mm. Body dorsally smooth. Cephalon lacking middorsal elevation. Antennule 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 filiformis (Say, 1818) Figure 108C
DIAGNOSIS 6 10.5 mm, ovigerous 2 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 floridana Richardson, 1901 Figure 108D
DIAGNOSIS Ovigerous ¢ 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.
Miratidotea 259
Idotea Fabricius, 1798
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
ie Postenormargin) of pleotelson, truncate, yo...) je a cuneis.s » (ores eae metallica Posterior margin of pleotelson with distinct median lobe ...... balthica
Idotea balthica (Pallas, 1772) Figure 108E
DIAGNOSIS. 6 24.5 mm, ovigerous 2 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 2? 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.
260. VALVIFERA °¢ IDOTEIDAE
Miratidotea bruscai Kensley, 1987a Figure 108G
DIAGNOSIS Ovigerous 2 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 dimeroceras (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 Tanirella caribbica Menzies, 1956b. South of Jamaica, 1244 m Tanirella 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 )
264 ZOOGEOGRAPHY
TABLE 4. (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 GNATHIIDEA Family Gnathiidae Akidognathia poteriophora Monod, 1926. Off U. S. Virgin Islands, 914 m. SUBORDER VALVIFERA Family Arcturidae Antarcturus annaoides Menzies, 1956b. South of Jamaica, 1244 m Arcturus caribbaeus Richardson, 1901. Off Aves 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
ZOOGEOGRAPHY 265
Car 74.8%
GoM/N/Car 7.2%
GoM/Car 5.4%
North 3.6% Out 1.8% Panam 4.2%
South 3.0% ==
Epicaridea excluded
Car 67.5%/
GoM/N/Car 9.6%
GoM/Car 6.4%
South 4.1% North 5.5% Panam 3.7% Out 3.2%
Epicaridea included
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
266 ZOOGEOGRAPHY
TABLE 5. SPECIES OF ISOPODS OCCURRING ON BOTH SIDES OF THE ISTHMUS OF PANAMA
*Aega deshaysiana (H. Milne Edwards, _Paradella dianae (Menzies, 1962b)
1840) Paraleptosphaeroma glynni Buss and Anopsilana browni (Van Name, 1936) Iverson, 1981 Cleantioides planicauda (Benedict, Probopyrus pandalicola (Packard, 1899) 1879) Excirolana braziliensis Richardson, *Rocinela oculata Harger, 1883 1912 *Rocinela signata Schioedte and Excorallana tricornis (Hansen, 1890) Meinert, 1879 *Nerocila acuminata Schioedte and Uromunna reynoldsi Frankenberg and Meinert, 1881 Menzies, 1966
* 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 Hoff (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 are “endemic,” three of these being interstitial microcerberideans.
ZOOGEOGRAPHY 267
TABLE 6. ISOPOD SPECIES OCCURRING IN THE GULF OF MEXICO
SUBORDER ANTHURIDEA
*Accalathura crenulata (Richardson, 1901)
*Amakusanthura magnifica (Menzies and Frankenberg, 1966)
Cyathura polita (Stimpson, 1855)
Horoloanthura irpex Menzies and Frankenberg, 1966
Kupellonura formosa (Menzies and Frankenberg, 1966)
* Mesanthura floridensis Menzies and Kruczynski, 1983
* Mesanthura hopkinsi Hooker, 1985
* Mesanthura pulchra Barnard, 1925
Paranthura floridensis Menzies and Kruczynski, 1983
Ptilanthura tricarina Menzies and Frankenberg, 1966
Skuphonura lindae Menzies and Kruczynski, 1983
*Xenanthura brevitelson Barnard, 1925
SUBORDER ASELLOTA
Carpias floridensis Menzies and Kruczynski, 1983
Gnathostenetrioides pugio Hooker, 1985
* Joeropsis coralicola Schultz and McCloskey, 1967
* Joeropsis rathbunae Richardson, 1902
Mexicope kensleyi Hooker, 1985
Munnogonium wilsoni Hooker, 1985
*Pleurocope floridensis 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 floridanus Markham, 1985
*Aporobopyrina anomala Markham, 1973
*Azygopleon schmitti (Pearse, 1932)
*Bopyrina abbreviata Richardson, 1904
*Bopyrione synalphei Bourdon and Markham, 1980
*Cancricepon choprae (Nierstrasz and Brender a Brandis, 1925)
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 and Brender a Brandis, 1929)
*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
*Furydice convexa Richardson, 1900b
Eurydice littoralis (Moore, 1901) *Eurydice piperata Menzies and Frankenberg, 1966
*Excirolana braziliensis Richardson, 1912a
*Excirolana mayana (Ives, 1891)
*Excorallana 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 Kruczynski, 1983
*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 Frankenberg, 1966
*Sphaeroma quadridentata Say, 1818
*Sphaeroma terebrans Bate, 1866
SUBORDER GNATHIIDEA
Gnathia floridensis Menzies and Kruczynski, 1983 SUBORDER MICROCERBERIDEA Microcerberus mexicanus Pennak, 1958
SUBORDER VALVIFERA
Antarcturus floridanus (Richardson, 1900b)
Arcturella bispinata Menzies and Kruczynski, 1983
Arcturella spinata Menzies and Kruczynski, 1983
Astacilla lauffi Menzies and Frankenberg, 1966
Chiridotea excavata Harper,
ZOOGEOGRAPHY 269
Edotea lyonsi (Menzies and Kruczynski, 1983)
Edotea montosa (Stimpson, 1853)
Edwinjoycea horologium Menzies and Kruczynski, 1983
*Erichsonella attenuata (Harger, 1873)
*Frichsonella filiformis (Say, 1818)
*Erichsonella floridana Benedict, 1901
1974 Erichsonella isabelensis Menzies, *Cleantioides planicauda (Benedict, 1951b 1899) *Idotea metallica Bosc, 1802
* species also occurring in the Caribbean Note: Records for the Gulf of Mexico have been assembled from published literature; in most cases, actual material has not been examined.
BERMUDA
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).
CAVE 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.
270 ZOOGEOGRAPHY
Alcirona krebsi Hansen, 1890
*Anthomuda stenotelson Schultz, 1979
*Apanthura harringtoniensis Wagele, 1981
*Arubolana aruboides (Bowman and Iliffe, 1983)
*Atlantasellus cavernicolus Sket, 1979
Bopyrissa wolffi Markham, 1978
Cancricepon choprae (Nierstrasz and Brender a Brandis, 1925)
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 personata Kensley, 1987b
TABLE 7. ISOPOD SPECIES OCCURRING AT 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
Parathelges 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
* recorded only from Bermuda
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 Curacao, 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. canariensis 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 during the Pleistocene probably served further to isolate these freshwater forms.
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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, Cirolanides, 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.
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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 collard, 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 Miller, 1988 Miller, H. G. 1988. Idoteidae aus N-Kolumbien mit Beschreibung von Edotea samariensis n. sp. (Crustacea: Isopoda: Valvifera). Senckenbergiana biologica 68(4/6):407—4 12.
Gnathia johanna Monod, 1926 Miller, 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- tigaciones 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.
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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.
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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(11):325a—442.
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Bate, C. S. 1866. Carcinological gleanings, 2. Annals and Magazine of Natural History (3)17:24— ale
Bate, C. S., and J. O. Westwood. 1868. A History of the British Sessile-eyed Crustacea. lvi + 536 pp. London: John van Voorst.
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Index
abbreviata, Bopyrina, 110, 267 Abudefduf saxatilis, 171, 175, 186 abudefdufi, Anilocra, 171, 175 Abyssianira dentifrons, 263 Acanthocope spinosissima, 263 acanthophora, Domecia, 111 Acanthopleura granulata, 215 acanthura, Anopsilana, 124, 125 acanthuri, Anilocra, 171, 175, 176 Acanthurus
bahianus, 171, 176
chirurgus, 171, 176 acanthurus, Macrobrachium, 112 Accalathura,64
crenulata, 64, 65, 267
setosa, 64, 65 Achelion occidentalis, 110 acuminata, Nerocila, 171, 172, 173, 190, 266,
268
forma acuminata, 190
forma aster, 190 acuta, Anilocra, 268 acutirostrata, Stenobermuda, 106, 270 acutirostris, Processa, 113 acutitelson, Dynamenella, 213, 214 Aega, 116, 117
antillensis, 117
incisa, 268 Aega (Aega), 116, 117
deshaysiana, 117, 266, 268
ecarinata, 117, 268 Aega (Rhamphion), 116, 117
dentata, 117, 119
tenuipes, 117, 119 Aegidae, 115, 116, 262 Aetobatus narinari, 166 affinis
Pseudione, 113, 270
Upogebia, 112 Agaricia, 88, 166 agaricicola, Metacirolana, 153, 154 Agarna, 170, 173
cumulus, 173
Agelas, 88 Akidognathia poteriophora, 264 alba, Exosphaeroma, 229 albidoida, Cirolana, 132, 133 Alcirona, 157, 158
insularis, 158
krebsii, 158, 268, 270 algicola, Carpias, 82, 83, 87 Allodardanus bredini, 112 Allodiplophryxus floridanus, 267 alphei, Probopyria, 112, 267 Alpheus
armillatus, 112
formosus, 111, 112
heterochaelis, 112
normanni, 112
viridari, 112 alphezemiotes, Ovobopyrus, 267 Alutera schoepfi, 171, 190 Amakusanthura, 17, 18
geminsula, 18
lathridia, 18, 20
magnifica, 18, 20, 267
signata, 18, 21
significa, 18, 23 amazonicum, Macrobrachium, 112 Ambidexter symmetricus, 113 americana, Dasyatis, 122, 166 americanus, Periclimenes, 111, 113 Amphiroa, 219 amyle, Licranthura, 43 analis, Lutjanus, 122, 172, 183 Anchoa lamprotaenia, 171, 187 Ancininae, 204 Ancinus, 205
belizensis, 205
brasiliensis, 205, 206
depressus, 268 andrewsi, Paralimnoria, 199 Angliera, 90, 91, 273
dubitans, 91
psamathus, 91
racovitzai, 91
293
294 INDEX
angulata
Dynamene, 214
Dynamenella, 214 angustifrons, Hexapanopeus, 11] Anilocra, 170, 174, 175
abudefdufi, 171, 175
acanthuri, 171, 175, 176
acuta, 268
chaetodontis, 171, 175, 177
chromis, 172, 175, 177
haemuli, 172, 173, 175, 177
holacanthi, 172, 175, 179
holocentri, 175, 179
laticauda, 268
myripristis, 172, 175, 180
partiti, 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
browni, 124, 125, 266
crenata, 124, 125
cubensis, 124, 126
Jonesi, 124, 127
radicicola, 124, 127 Antarcturus
annaoides, 264
Sloridanus, 269 Antennuloniscus dimeroceras, 263 Antheluridae, 275 Anthomuda, 17, 23, 291
stenotelson, 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
crucis, 25, 26
harringtoniensis, 25, 26, 270 Apanthuroides, 17, 26
millae, 27 Aphysina fistularis, 223
Apogon
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
Spinata, 252, 269 Arcturidae, 251, 252, 264 Arcturus
caribbaeus, 264
purpureus, 264 arenatus, Priacanthus, 173, 183 Argeia, 110
atlantica, 110 Argeiinae, 107 Arius felis, 171, 190 Armadilloniscus, 247
ninae, 247 armatus
Ischnomesus, 263
Petrolisthes, 110 armillatus, Alpheus, 112 arndti, Dies, 207
aruboides, Arubolana, 144, 145, 270
Arubolana, 144, 273
aruboides, 144, 145, 270
imula, 144, 145
Parvioculata, 144, 145 ascensionis, Holocentrus, 172, 179 Aselloidea, 75
Asellota, 15, 15, 73, 263, 267, 269
Astacilla, 252 cymodocea, 252, 253 lasallae, 252, 253 lauffi, 269 regina, 252, 253
Astalione, 110 cruciaria, 110
aster, forma, Nerocila acuminata, 190
Astrapogon stellatus, 171, 188 Asymmetrione, 110
clibanarii, 110
desultor, 110 Athelginae, 107 Atherinidae, 187
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 barnardi
Dies, 207
Ischyromene, 218
Paranthura, 69, 71 barracuda, Sphyraena, 122 barrerae, Cymodoce, 227 Bathynomus, 123, 129
giganteus, 131, 268 Batrachoides surinamensis, 171, 190 baudiniana, Ligia, 247, 249 beethoveni, Gnathia, 238, 239 belizensis, Ancinus, 205 Belonidae, 187 benjamensis, Paradynamene, 268 berbicensis, Excorallana, 161, 162 bermudensis
Carpias, 82, 83, 270
Curassanthura, 67, 270
Hemiramphus, 188 bicornutus, Microphrys, 110 bifasciatus, Joeropsis, 88 bifurcatus, Heteromesus, 263 bigelowiensis, cf., Prodajus, 268 bimini, Leidya, 111, 270 bispinata, Arcturella, 252, 269 bivittata, Mesanthura, 47 blackfordi, Lutjanus, 122 bohlkeorum, Mothocya, 171, 173, 187, 188 bonaci, Mycteroperca, 122
INDEX
bonairensis, Pagurus, 110, 113 bonelli, Macrobrachium, 112 bonnieri, Pandalus, 113 Booralana tricarinata, 275 Bopyrella, 110
harmopleon, 110 Bopyridae, 107, 114 Bopyrina, 110
abbreviata, 110, 267 Bopyrinae, 107 Bopyrinella, 110
thorii, 110 Bopyrione, 110
synalphei, 110, 267 Bopyrissa, 110
wolffi, 110, 270 Bopyroidea, 107 Bopyrophryxinae, 107 borealis, Cirolana, 268 Bothus lunatus, 122 bousfieldi, Synalpheus, 110 bowmani
Renocila, 173, 191
Stenetrium, 100 brachydactylus, Carpias, 82, 84 brasiliensis
Ancinus, 205, 206
Hemiramphus, 172, 184 braziliensis, Excirolana, 150, 266, 268 bredini, Allodardanus, 112 brevicarpus, Synalpheus, 110 brevidactylus, Pagurus, 112, 113 brevipes, var., Paracerceis caudata, 219 brevitelson, Xenanthura, 62, 267 brooksi, Synalpheus, 110, 111, 113 browni, Anopsilana, 124, 125, 266 bruscai, Miratidotea, 260 buccanella, Lutjanus, 122 Bythognathia yucatanensis, 275 Cabirops, 110 Calamus
bajonado, 122
calamus, 122
penna, 122 calamus, Calamus, 122 Callispongia plicifera, 221 Calyptolana, 124, 131
hancocki,132 camayae, Nannoniscus, 264 canaliculata, Processa, 113 canariensis, Curassanthura, 270, Cancricepon, 110
choprae, 110, 267, 270
295
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 caribbica
Tanirella, 263
Malacanthura, 45 caribbicus
Ischnomesus, 263
Macrostylis, 263 caribea, Uromunna, 94 carinata, ‘Cerceis,‘ 211, 268 carolii, Metaphrixus, 111 carolinense, Tozeuma, 112 carolinus, Cancrion, 111 Carpias, 82, 83
algicola, 82, 83, 87
bermudensis, 82, 83, 270
brachydactylus, 82, 84
floridensis, 267
harrietae, 82, 84
minutus, 82, 84, 270
punctatus, 82, 85
serricaudus, 82, 87
triton, 82, 87 Cassidinidea, 207, 208
mosaica, 208
ovalis, 207, 208, 268 Cassidininae, 204, 207 castrensis, Hyperphrixus, 267 caudata, Paracerceis, 219, 268, 270 Caulerpa, 219 cavalla, Scomberomorus, 173, 187 cavernicolus, Atlantasellus, 75, 269, 270 cephalus, Mugil, 172, 190 Ceratothoa, 170, 180
deplanata, 180
transversa, 268 Cerceis, 210, 211
carinata, 211, 268 Chaetodipterus faber, 171, 190 Chaetodon
capistratus, 171, 177
ocellatus, 171, 177
sedentarius, 171, 177 striatus, 171, 177
chaetodontis, Anilocra, 171, 175, 177
Chalixanthura, 17, 27
lewisi, 27, 29
scopulosa, 27, 29 Chilomycterus schoepfi, 172, 190 Chiridotea excavata, 269
chirurgus, Acanthurus, 171, 176
Chiton
marmoratus, 229
tuberculatus, 229 choprae
Cancricepon, 110, 267, 270
Synsynella, 113, 268, 270 Chromis
cyaneus, 172, 177
multilineatus, 172, 177 chromis, Anilocra, 172, 175, 177 chrysargyreum, Haemulon, 172, 178 chrysoptera, Orthopristis, 172, 183 chrysurus, Ocyurus, 172, 183 ciliatus, Monacanthus, 172, 190 circumsaltanus, Loki, 111 Cirolana, 123, 132
albidoida, 132, 133
borealis, 268
crenulitelson, 132, 133
minuta, 132, 135
obtruncata, 132, 135, 268
parva, 132, 135, 268 Cirolanidae, 1, 115, 122, 123, 139, 273 Cirolanides, 273 Cirolaninae, 123, 139 cirratum, Ginglymostoma, 122, 158 clarkae, Phycolimnoria, 201 Clastotoechus vanderhorsti, 110 Cleantioides, 255, 256
occidentalis, 256
planicauda, 256, 266, 269 clibanarii
Asymmetrione, 110
Stegias, 113, 270 Clibanarius
tricolor, 110, 112, 113
vittatus, 110, 112 coeca, Neoanthura, 263 cognata, Pseudione, 268 cohenae, Paracerceis, 219, 220 Colanthura, 64, 65
tenuis, 65, 270 colini, Renocila, 171, 191 collardi, Aporobopyrus, 275
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 coralicola, 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 crucis, Apanthura, 25, 26 cruentatus, Epinephelus, 172, 179 crumenophthalmus, Selar, 173, 183 Cryptoniscoidea, 107 Crytoniscidae, 107, 109 cubana, Cyathura (Cyathura), 33 cubense, Munidion, 111 cubensis
Anopsilana, 124, 126
Rocinela, 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
halma, 67, 68, 270 curassavica, Cyathura (Stygocyathura), 35 curri, Haliophasma, 41 curtatus, Aporobopyrus, 110 cuvieri, Galeocerdo, 122 cyaneus, Chromis, 172, 177 Cyathura, 17, 31, 270, 273
polita, 267
(Cyathura), 31
cubana, 33
INDEX
(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 excisa, 172, 173, 182, 268
oestrum, 171, 172, 173, 182, 183, 268
Cymothoidae, 115, 169, 170 Cynoscion, 172, 183 nebulosus, 172, 183 Dactylokepon caribaeus, 111 sulcipes, 267 Dajidae, 107 Dardanus fucosus, 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 dianae, Paradella, 224, 266, 268 Dicropleon
periclimenis, 111 Dictyota, 219 Dies, 207
arndti, 207
barnardi, 207
297
298 INDEX
dimeroceras, Antennuloniscus, 263 floridana, 256, 258, 269 diminuta, Exosphaeroma, 229, 231 isabelensis, 269 diogenes, Petrochirus, 110 Eriphia gonagra, 111 Diopatra cuprea, 256 Eubranchiatae, 203 Diplophryxus, 111 Eurycopidae, 263 Discerceis, 210, 211 Eurydice, 143, 147 linguicauda, 213 convexa, 147, 148, 149, 268 dispar, Paraliomera, 111 littoralis, 148, 149, 268 distorta, Leidya, 111 personata, 147, 149, 270 Domecia piperata, 147, 149, 268 acanthophora, 111 Eurydicinae, 123, 139, 143 hispida, 111 excavata, Chiridotea, 269 Dromidia antillensis, 111 Excirolana, 144, 149, 150 dubitans, Angliera, 91 braziliensis, 150, 266, 268 Dynamene angulata, 214 mayana, 150, 153, 268 Dynamenella, 210, 213, 214, 224 excisa, Cymothoa, 172, 173, 182, 268 acutitelson, 213, 214 Excorallana, 157, 159, 161 var. glabrothorax, 214 antillensis, 161, 162, 268 var. typica, 214 berbicensis, 161, 162 angulata, 214 delaneyi, 160, 161 perforata, 213, 215, 270 fissicauda, 161, 162 quadrilirata, 213, 215 mexicana, 160, 161, 268 Dynameninae, 204, 210, 211 oculata, 161, 163 ecarinata, Aega, 117, 268 quadricornis, 161, 165, 270 Echinothambema ophiuroides, 263 sexticornis, 161, 165 Echinothambematidae, 263 subtilis, 160 edithae, Paracerceis, 218, 221 tricornis, 266, 268 Edotea occidentalis, 167 lyonsi, 269 tricornis, 161, 165 montosa, 269 warmingti, 161, 167 samariensis, 276 exilipes, Palaemonetes, 113 edulis, Processa, 113 Exocoettus, 172, 184 edwardsi, Plesionika, 113 Exosphaeroma, 226, 229, 231 Edwinjoycea horologium, 252, 269 alba, 229 eglanteria, Raja, 122 antillense, 229, 230 Eisothistos, 16, 38, 39 diminuta, 229, 231 petrensis, 39 productatelson, 229, 231 teri, 39 yucatanum, 229, 231 ensis, Plesionika, 113 exotica, Ligia, 247, 249 Entoniscidae, 107, 109 Jaber, Chaetodipterus, 171, 190 Entophilinae, 107 fasciata, Mesanthura, 47, 49 Eophrixus subcaudalis, 111, 267 faustinum, Macrobrachium, 112 Epicaridea, 4, 14, 107, 267 Jaxoni, Harrieta, 232, 268 Epinephelus, 122, 172, 183 felis, Arius, 171, 190 cruentatus, 172, 179 filiforme, Syringodium, 253, 260 Julvus, 172, 179 filiformis, Erichsonella, 256, 258, 269 guttatus, 172, 179 fimbriata itajara, 122, 172, 190 Pleurocryptella, 112 morio, 122 Processa, 113 Erichsonella, 255, 256, 257 Jissicauda, Excorallana, 161, 162 attenuata, 256, 258, 269 fistularis, Aphysina, 223 filiformis, 256, 258, 269 Flabellifera, 2, 14, 114, 115, 268, 269
tropicalis, 257 flavolineatum, Haemulon, 122, 172, 178
flinti, Munida, 111 floridana
Erichsonella, 256, 258, 269
Pleurocrypta, 112 floridanus
Allodiplophryxus, 267
Antarcturus, 269
Thor, 110, 111 Sloridensis
Carpias, 267
Gnathia, 268
Mesanthura, 53, 267
Paranthura, 69, 71, 267
Pleurocope, 98, 267 foetens, Synodus, 173, 183 foliatus, Parathelges, 112 formosa, Kupellonura, 267 formosus, Alpheus, 111, 112 fritzmuelleri, Synalpheus, 110, 111 fucorum, Latreutes, 112 fucosus, Dardanus, 112 fulvus, Epinephelus, 172, 179 furcifer, Paranthias, 173, 179 Galathea rostrata, 112 galathinus, Petrolisthes, 110 Galeocerdo cuvieri, 122 geminsula, Amakusanthura, 18 Geocerceis, 210, 215
barbarae, 215 geracei, Bahalana, 128 Gerres 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
glynni Paracerceis, 218, 221 Paraleptosphaeroma, 210, 266
Gnathia, 238 beethoveni, 238, 239 floridensis, 268 gonzalezi, 238, 239 johanna, 238, 239, 276 magdalenensis, 238, 239 puertoricensis, 238, 239 rathi, 238, 241 samariensis, 238, 241
INDEX
triospathiona, 238, 241
velosa, 238, 243
virginalis, 238, 243 Gnathiidae, 237, 238 Gnathiidea, 14, 236, 238, 268 Gnathostenetroides, 77
laodicense, 78
pugio, 77, 267 Gnathostenetroididae, 77 Gnathostenetroidoidea, 77 gonagra, Eriphia, 111 gonzalezi, Gnathia, 238, 239 goodei, Synalpheus, 110, 111 gracilis, Natatolana, 140 granulata, Acanthopleura, 215 guttatus, Epinephelus, 172, 179 haemuli, Anilocra, 172, 173, 175, 177 Haemulon
aurolineatum, 172, 178
carbonarium, 172, 178
chrysargyreum, 172, 178
flavolineatum, 122, 172, 178
macrostomum, 172, 179
plumiert, 172, 179
sciurus, 172, 179
steindachneri, 122 halia, Metacirolana, 153, 154 Halimeda, 88, 90, 219, 223 Haliophasma, 17, 41
curri, 41
irmae, 43
valeriae, 41, 42 halma, Curassanthura, 67, 68, 270 Halodule, 232 hancocki, Calyptolana, 132 hanseni, Dendrotion, 263 Haplomesus tropicalis, 263 Haploniscidae, 263 Haploniscus unicornis, 263 Haptolana, 124, 137, 273
somala, 138
trichostoma, 138 harmopleon, Bopyrella, 110 Harrieta, 226, 232
faxoni, 232, 268 harrietae, Carpias, 82, 84 harringtoniensis, Apanthura, 25, 26, 270 harrisii, Rithropanopeus, 111 hayesi, Uromunna, 267 heardi, Probopyrinella, 267 Hemiarthrinae, 107 Hemiarthrus synalphei, 111, 267 Hemibranchiatae, 203
209
300 INDEX
hemiramphi, 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, Alpheus, 112 Heteromesus bifurcatus, 263 Hexapanopeus angustifrons, 111 Hippolyte
curacaoensis, 110
pleuracanthus, 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 Tanirella
caribbica, 263
vemae, 263 Idotea, 255, 259
balthica, 259
metallica, 259, 269 Idoteidae, 251, 252, 255 Idoteinae, 255, 256 Iliacantha
liodactyla, 111
subglobosa, 111 imbricata, Parapagurion, 112 impressa, Politolana, 140 impressus, Glossobius, 172, 184 imswe, Kupellonura, 60 imula, Arubolana, 144, 145 incisa, Aega, 268
indica, Limnoria, 194, 195
infundibulata, Paranthura, 69, 71, 270
insulae, Limnoria, 195 insularis
Alcirona, 158
Kuna, 170
Rocinela, 119, 120, 268 integra, Synsynella, 268 interger, Cyclograpsus, 111 intermedius, Palaemonetes, 113 Ioninae, 107 Iridopagurus, 112, 113
iris, 113 iris, Iridopagurus, 113 irmae, Haliophasma, 43 irpex, Horoloanthura, 267 irrasa, Munida, 111 irritans, Munidion, 111 isabelensis, Erichsonella, 269 Ischnomesidae, 263 Ischnomesus
armatus, 263
caribbicus, 263
multispinis, 263 Ischyromene, 210, 214, 217
barnardi, 218
itajara, 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 johanna, Gnathia, 238, 239, 292 jonesi, Anopsilana, 124, 127 kadiakensis, Palaemonetes, 113 kensleyi, Mexicope, 81, 267 kohleri, Mesosignum, 264 krebsi, Alcirona, 158, 268, 270 Kuna, 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 fucorum, 112
latreuticola, Probopyrinella, 112, 268, 270
latus, Caranx, 171, 183 lauffi, Astacilla, 269 Laurencia, 219 Leidya bimini, 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 Ligia, 247, 249 baudiniana, 247, 249 exotica, 247, 249 olfersii, 247, 249 Limnoria, 193, 194, 195, 235 indica, 194, 195 insulae, 195 multipunctata, 195, 196 pfefferi, 195, 198 platycauda, 195, 198 saseboensis, 195, 198 simulata, 195, 198 tripunctata, 199 tuberculata, 194, 195, 199, 268 unicornis, 195, 199 Limnoriidae, 114, 193 lindae, Skuphonura, 267 linguicauda, Discerceis, 213 liodactyla, Iliacantha, 111 Lironeca, 170, 172, 186 ovalis, 268 redmanni, 172, 173, 186, 268 tenuistylis, 171, 186, 187 texana, 268 tropicalis, 268 littoralis Eurydice, 148, 149, 268 Microcerberus, 244 Loki circumsaltanus, 111 longicarpus Pagurus, 110, 113 Synalpheus, 110, 111, 113
INDEX
longicaudatus, Periclimenes, 113 longipedis, Munidion, 112, 267 longipes, Munida, 112 looensis, Mesanthura, 47, 51 lunatus, Bothus, 122 Lutjanus
analis, 122, 172, 183
blackfordi, 122
buccanella, 122
mahogoni, 172, 183
synagris, 172, 183 lyonsi, Edotea, 269 Lysmata
rathbunae, 112
wurdemanni, 112 Macrobrachium
acanthurus, 112
amazonicum, 112
bonelli, 112
carcinus, 112
faustinum, 112
ohione, 112
olfersii, 112
surinamicum, 112 Macrocystis, 201 macrostomum, Haemulon, 172, 179 Macrostylidae, 263 Macrostylis
caribbicus, 263
minutus, 263
setifer, 264
vemae, 264 maculatus
Scomberomorus, 173, 187
Sphoeroides, 173, 190 Madracis, 29, 41, 88, 90, 166
mirabilis, 29, 41 magdalenensis, Gnathia, 238, 239 magnifica, Amakusanthura, 18, 20,
267
magnispina, Desmosoma, 263 mahogoni, Lutjanus, 172, 183 Malacanthura, 17, 43
caribbica, 45
cumanensis, 45 mangle, Rhizophora, 235 manningi, Thor, 111 marcuzzii, Tylos, 247, 250 margarita, Pontonia, 113 marginatus, Petrolisthes, 110 marmoratus, Chiton, 229 martia, Plesionika, 113 maximus, Lachnolaimus, 122
301
302 INDEX
mayana
Bahalana, 128
Excirolana, 150, 153, 268 mceclendoni, Synalpheus, 110, 111 Megalops atlantica, 172, 183 menziest, Metacirolana, 153, 154 Mesanthura, 17, 45
bivittata, 47
decorata, 53
fasciata, 47, 49
floridensis, 53, 267
hopkinsi, 47, 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
halia, 153, 154
menziesi, 153, 154
sphaeromiformis, 153, 154 metallica, Idotea, 259, 269 Metaphrixus carolii, 111 mexicana, Excorallana, 160, 161, 268 mexicanus, Microcerberus, 269 Mexicope, 80, 81
kensleyi, 81, 267 Mexilana, 273 mgrayi, Serolis, 202, 268 Microcerberidae, 243, 244 Microcerberidea, 14, 243, 269, 273 Microcerberus, 244, 273
littoralis, 244
mexicanus, 269
minutus, 244
mirabilis, 244
nunezi, 244
renaudi, 244
simplex, 244
syrticus, 244 Microcharon, 90, 91
herrerai, 93
phreaticus, 93
sabulum, 91 Micropanope barbadensis, 111 Microparasellidae, 80, 90, 273 Microphrys bicornutus, 110 miles, Munida, 112 millae, Apanthuroides, 27 milleri, Santia, 99, 267
minocule, Stenetrium, 100, 102 minus, Synalpheus, 110, 113 minuta, Cirolana, 132, 135 minutus Carpias, 82, 84, 270 Macrostylis, 263 Microcerberus, 244 Minyanthura, 16, 53 corallicola, 53 mirabilis Madracis, 29, 41 Microcerberus, 244 Miratidotea, 255, 259 bruscai, 260 Monacanthus ciliatus, 172, 190 montagui, Pandalus, 113 montosa, Edotea, 269 morio, Epinephelus, 122 mortenseni Gigantione, 111, 267 Parabopyrella, 112, 267
mosaica, Cassidinidea, 208
motasi, Cyathura (Stygocyathura), 35, 36
Mothocya, 170, 187 bohlkeorum, 171, 173, 187, 188 nana, 172, 187, 188
Mugil cephalus, 172, 190
multilineatus, Chromis, 172, 177
multipunctata, Limnoria, 195, 196
multispinis, Ischnomesus, 263
Munida constricta, 112 flinti, 111 irrasa, 111 longipes, 112 miles, 112 schroederi, 112 simplex, 110 stimpsoni, 111 valida, 110
Munidion cubense, 111 irritans, 111 longipedis, 112, 267
Munna, 93 petronastes, 94
Munnidae, 80, 93
Munnogonium, 96 wilsoni, 96
Moycteroperca bonaci, 122 venenosa, 122
Mpripristis 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 texana sayi, 111 Neostenetroides, 77, 78 stocki, 78 Nerocila, 170, 172, 188 acuminata, 171, 172, 173, 190, 266, 268 forma acuminata, 190 forma aster, 190 ninae, Armidilloniscus, 246, 247 niveus, Tylos, 247, 250 normanni, Alpheus, 112 northropi, Palaemon, 113 nunezi, Microcerberus, 244 nuttingi, Paracerceis, 218, 223 obtruncata, Cirolana, 132, 135, 268 occidentalis Achelion, 110 Cleantioides, 256 Excorallana tricornis, 167 Parathelges, 112 ocellatus, Chaetodon, 171, 177 | oculata Excorallana, 161, 163 Rocinela, 119, 120, 266, 268 Oculina arbuscula, 88 Ocyurus chrysurus, 172, 183 oestrum, Cymothoa, 171, 172, 173, 182, 183, 268 ohione, Macrobrachium, 112 Olencira praegustator, 268 olfersii Ligia, 247, 249 Macrobrachium, 112 Oncilorpheus, 124, 139 stebbingi, 139 Oniscidea, 1, 4, 14, 15, 246, 247, 262 ophiuroides, Echinothambema, 263 Orbioninae, 107 orghidani, Cyathura (Stygocyathura), 35, 36
INDEX
Orthopristis chrysoptera, 172, 183 ruber, 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 annulicornis, 113 bonnieri, 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 caudata, 219, 268, 270 var. brevipes, 219 cohenae, 219, 220 edithae, 218, 221 glynni, 218, 221 nuttingt, 218, 223
303
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 glynni, 210, 266 Paralimnoria, 193, 199 andrewsi, 199 forma A, 200 forma B, 201 forma typica, 200 Paraliomera dispar, \11 Paramunnidae, 80, 96 Paranthias furcifer, 173, 179 Paranthura, 64, 69 antillensis, 69 barnardi, 69, 71 floridensis, 69, 71, 267 infundibulata, 69, 71, 270 Paranthuridae, 16, 64, 263, 270 Parapagurion imbricata, 112 Parapagurus, 112 parapotamica, Cyathura (Stygocyathura), 35, 36 Parathelges foliatus, 112 occidentalis, 112 piriformis, 112, 270 tumidipes, 112, 270 partiti, Anilocra, 173, 175, 180 partitus, Pomacentrus, 173, 180 parva, Cirolana, 132, 135, 268 parvioculata, Arubolana, 144, 145 parviramus, Pseudione, 276 parvus, Colopisthus, 147, 270 patulipalma, Stenetrium, 100, 102 paucidens, Mesanthura, 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, Eisothistos, 39
Petrochirus diogenes, 110 Petrolisthes
armatus, 110
galathinus, 110
marginatus, 110 petronastes, Munna, 94 pfefferi, Limnoria, 195, 198 Phaeoptyx
conklini, 173, 188
pigmentaria, 173, 188 phreaticus, Microcharon, 93 Phreatocoidea, 14 Phycolimnoria, 193, 201
bacescui, 276
clarkae, 201 Phyllodurinae, 107 pigmentaria, 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
edwardsi, 113
ensis, 113
heterocarpus, 113
martia, 113 pleuracanthus, Hippolyte, 110, 111 Pleurocope, 97
floridensis, 98, 267 Pleurocopidae, 80, 81, 96 Pleurocrypta floridana, 112 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
polita, 140, 143, 268 Pomacentrus partitus, 173, 180 Pontonia margarita, 113 Porcellana sayana, 110 Porites, 88, 90 poteriophora, Akidognathia, 264 praegustator, Olencira, 268 Priacanthus arenatus, 173, 183 probatocephalus, Archosargus, 122 Probopyria alphei, 112, 267
Probopyrinella
heardi, 267
latreuticola, 112, 268, 270 Probopyrus pandalicola, 112, 266 Processa
acutirostris, 113
canaliculata, 113
edulis, 113
JSimbriata, 113
tenuipes, 113 processae, Urobopyrus, 113, 268 Prodajus cf. bigelowiensis, 268 productatelson, Exosphaeroma, 229, 231 provenzanoi, Pagurus, 110, 112, 113 psamathus, Angliera, 91 Pseudasymmetrione, 113 Pseudione
affinis, 113, 270
cognata, 268
parviramus, 276
upogebiae, 268 Pseudioninae, 107 Ptilanthura tricarina, 267 puertoricensis, Gnathia, 238, 241 pugilator, Uca, 111 pugio
Gnathostenetroides, 77, 267
Palaemonetes, 113 pulchra
caribbea, Storthyngura, 263
Mesanthura, 47, 52, 267 punctatus, Carpias, 82, 85 punctillata, Mesanthura, 47, 53 purpureus, Arcturus, 264 Pylopagurus, 110
corallinus, 112 quadricornis, Excorallana, 161, 165, 270 quadridentata, Sphaeroma, 234, 268 quadrifrons, Hydroniscus, 263 quadrilirata, Dynamenella, 213, 215 quadripunctata, Paradella, 223, 223 racovitzai, Angliera, 91 radicicola
Anopsilana, 124, 127
Xylolana, 157 Raja eglanteria, 122 rapax, Nalicora, 169, 268 rathbunae
Joeropsis, 88, 90, 267, 270
Lysmata, 112 rathi, Gnathia, 238, 241 redmanni, Lironeca, 172, 173, 186, 268 regalis, Scomberomorus, 173, 187
INDEX
regina, Astacilla, 252, 253 renaudi, Microcerberus, 244 Renocila, 170, 191
bowmani, 173, 191
colini, 171, 191
waldneri, 173, 191, 193 reticulata, Mesanthura, 47, 53 reynoldsi, Uromunna, 94, 95, 266, 267 Rhamphion, see Aega (Ramphion) Rhizophora mangle, 235 rhombeus, Gerres, 172, 187 Rhyscotus, 247, 249
texensis, 247, 249 richardsonae, Parabopyrella, 112, 267 ricordi, Sesarma, 111 riedli, Vandeloscia, 247, 251 Rithropanopeus harrisii, 111 Rocinela, 116, 119
cubensis, 119, 120
insularis, 119, 120, 268
oculata, 119, 120, 266, 268
signata, 119, 120, 266, 268 rostrata, Galathea, 112 ruber
Caranx, 171, 183
Orthopristis, 122, 173, 179 ruetzleri, Cymodoce, 227 sabulosa, Hypoconcha, 111 sabulum, Microcharon, 91
305
salpiscinalis, Cyathura (Stygocyathura), 35, 38
Samariensis
Edotea, 276
Gnathia, 238, 251 Santia, 98
milleri, 99, 267 Santiidae, 80, 98 Sargassum, 84, 201 saseboensis, Limnoria, 195, 198 Saxatilis, Abudefduf, 171, 175, 186 sayana, Porcellana, 110 sayi, Neopanope texana, 111 sbordonii, Cyathura (Stygocyathura), 35,
38
Schizobopyrina urocaridis, 113, 268 schmitti, Azygopleon, 110, 267 schoepfi
Alutera, 171, 190
Chilomycterus, 172, 190 schroederi, Munida, 112 sciurus, Haemulon, 172, 179 Sclerocrangon jacqueti, 110 Scomberomorus
cavalla, 173, 187
306 INDEX
Scomberomorus (cont.)
maculatus, 173, 187
regalis, 173, 187 scopulosa, Chalixanthura, 27, 29 sedentarius, Chaetodon, 171, 177 Selar crumenophthalmus, 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 sexticornis, Excorallana, 161, 165 signata
Amakusanthura, 18, 21
Rocinela, 119, 120, 266, 268 significa, Amakusanthura, 18, 23 simplex
Microcerberus, 244
Munida, 110 simulata, Limnoria, 195, 198 Skuphonura, 17, 58
laticeps, 58
lindae, 267 snanoi, Storthyngura, 263 somala, Haptolana, 138 Sparisoma viride, 122
spathulicarpus, 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 walkeri, 234, 235
Sphaeromatidae, 114, 115, 202, 204
Sphaeromatinae, 204, 226
sphaeromiformis, Metacirolana, 153, 154
Sphoeroides maculatus, 173, 190
Sphyraena barracuda, 122
Spinata, Arcturella, 252, 269
Spinianirella serrata, 263
spinosissima
Acanthocope, 263
Hypoconcha, 111 stebbingi
Oncilorpheus, 139
Stenetrium, 100, 104, 267, 270 Stegias clibanarii, 113, 270 Stegophryxus hyptius, 113, 268 steindachneri, Haemulon, 122 stellatus
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 Storthyngura
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, 110
brooksi, 110, 111, 113
Sritzmuelleri, 110, 111
goodei, 110, 111
hemphilli, 110, 111
longicarpus, 110, 111, 113 meclendoni, 110, 111 minus, 110, 113 pandionis, 111, 113 pectiniger, 110, 111, 113 Synodus foetens, 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 tenuipes Aega, 117, 119 Processa, 113 tenuis, Colanthura, 65, 270 tenuistylis, Lironeca, 171, 186, 187 terebrans, Sphaeroma, 234, 235, 268 teri, 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 Thor floridanus, 110, 111 manningi, 111 thorii, 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 tricolor Clibanarius, 110, 112, 113 Holacanthus, 172, 179 tricornis, Excorallana, 161, 165, 266, 268 occidentalis, 167 tricornis, 161, 165 Tridentella, 236 virginiana, 236
INDEX
Tridentellidae, 115, 235, 236 triospathiona, Gnathia, 238, 241 tripunctata, Limnoria, 199 triton, Carpias, 82, 87 Troglocirolana, 273 tropicalis
Erichsonella filiformis, 257
Haplomesus, 263
Lironeca, 268 tuberculata, Limnoria, 194, 195, 199, 268 tuberculatus, Chiton, 229 tumidicauda, Paradella, 223, 226 tumidipes, Parathelges, 112, 270 Turbinaria, 166, 219 Tylos, 247, 250
latreillei, 247, 250
marcuzzit, 247, 250
niveus, 247, 250
wegeneri, 247, 250 typica, var., Dynamenella acutitelson, 214 uberlackerae, Gigantione, 267 Uca, 111
pugilator, 111 unicornis
Haploniscus, 263
Limnoria, 195, 199 unifasciatus, Hyporhamphus, 172, 188 univam, Cyathura (Stygocyathura), 35,
38
Upogebia affinis, 112 upogebiae, Pseudione, 268 Urobopyrus processae, 113, 268 urocaridis, Schizobopyrina, 113, 268 Uromunna, 93, 94
caribea, 94
hayesi, 267
reynoldsi, 94, 95, 266, 267 valeriae, Haliophasma, 41, 42 valida, Munida, 110 Valvifera, 14, 251, 264, 269 Vandeloscia, 247, 251
culebrae, 247, 251
riedli, 247, 251 vanderhorsti, Clastotoechus, 110 velosa, Gnathia, 238, 243 vemae
Tanirella, 263
Macrostylis, 264 venenosa, Mycteroperca, 122 Venezanthura, 31 venezuelensis, Thermarcturus, 255 vetula, Balistes, 122
307
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 Xylolana, 123, 143, 144, 156 radicicola, 157 yucatanensis, Bythognathia, 275 yucatanum, Exosphaeroma, 229, 231 Yuesia, 246 Striata, 246 zostericola, Hippolyte, 110
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