av f ih 4f i on it 4 sng ma bis fai alia A ae ae - ti a - ie t oh a ria i F i f : ; © Th, ny it, : Cal he >i j : et), ; Ae, Fi Ay at | = 4 ' bu? Tes avy ne € a ee ein a a f Guide to the MARINE ISOPOD CRUSTACEANS of the Caribbean “cespot sae : ou nt ‘ 7 1m in a ery uf a a te an lia a ce 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 P mn 7 ; hie 4 g tJ j ; - =7 1 t i rr ae) J) rs i : 7 ? x . rian an t ei'a es o Tey >" cai fh ve ERR gets ‘ae ie ited oft s sal % eboigaed knee a ; " ys baat ab ANY Peel . ae : ran ae ee ia ' ra matt cys. is my Os | eerie ‘ i i ae i ie = : ri it spit " a bis ea a ha ao Th 4 Rou, - ‘et ia: 7 re Aa sennitko th scala, yi 4 ay sind 8 Th ‘a NS EM arnt et ihe | | at are a » ai ” At eotve , ‘ a eye hen ne i or ” intense. i ae” ir, Shee!) juacripeaveuontadhine 408 , My ; ; y \c it) ein nn cy iso S % i; - \ i ‘ 1 7 ‘ vf i le : i x en 1 oi . Pg? Pr. rn ir: ve ‘oe th aioas anem fied 7 : ; “ja? ao ce " e Ll os ane Mae: PTR ayy " ri ue — en . 0 : ie > 4h ) ie Pasi abe ih eit a : ; Ne 4 ‘ P . 7 i , aN on | ‘if ue at’ : uid Ae : f, A if a | ny 7 HC a ee 6 ; ae iv ; > j Ms “4 | Co Me SOE ong Tee o WARS pi? , in 7 oni : Lan a : ey im NS 4 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 ‘apis sty Aq pasaaoo varie jo deypy "| aansty BIQUO|OD ejenzeue, O) seinpuoy Binsuluad uejyeon, epuol4 og 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. +. antennule a A flage#um frontal lamina peduncle Ne ene Wane | Fat Yon in a clypeus oe e maxilla 2 ¥ _ maxilla 1 maxilliped pereopods ‘co. «6s dactylus ~, igh ae (Ss carpus rg ~merus ischium uropod Figure 2. Schematic representation of an isopod illustrating morphological terms. 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 + 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 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’. ae OP BASE ON, ey ZS 2 WIAA ALES IVE OSS ON ANS So — 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.. 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. 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. “SUBIPLINYIUS IAPD jo uonNnqtystp SuIMOYs dey ‘OTT aInsly BinyyeAyD e BINYJUSSEIND ¥ eurjojdey o e||auesealD w eurgjeueg w euejoqniy a CURIISCOUY @ ‘spluvjosIO 2Avd JO UONNQIsIp Surmoys dey TL] ans ZOOGEOGRAPHY DAs 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. a ’ oe far Ps bl ; Sar ar hoary Meee st A ‘Meth andnaey tie siete Bedale wollte a) | 4nantecs oasramd aye afy eo Gwot os aan yelie: “ie } ifp