6 - Sphaeromatidae::“Cute As Buttons”
6 - Sphaeromatidae::“Cute As Buttons”
6 - Sphaeromatidae::“Cute As Buttons”
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Guide to the<br />
MARINE ISOPOD<br />
CRUSTACEANS<br />
of the Caribbean
Guide to the<br />
MARINE ISOPOD<br />
CRUSTACEANS<br />
of the Caribbean<br />
Brian Kensley and<br />
Marilyn Schotte<br />
SMITHSONIAN INSTITUTION PRESS<br />
WASHINGTON, D.C., AND LONDON
© 1989 by the Smithsonian Institution<br />
All rights reserved<br />
Designer: Linda McKnight<br />
Editor: Nancy Dutro<br />
Library of Congress Cataloging-in-Publication Data<br />
Kensley, Brian Frederick.<br />
Guide to the marine isopod crustaceans of the<br />
Caribbean / Brian Kensley<br />
and Marilyn Schotte.<br />
p. cm.<br />
Bibliography: p.<br />
Includes index.<br />
ISBN 0-87474-724-4 (alk. paper)<br />
1. Isopoda—Caribbean Sea—<br />
Classification. 2. Crustacea—Caribbean Sea—<br />
Classification. I. Schotte, Marilyn. II. Title.<br />
QL444.M34K434 1989<br />
595.3'7209153'35—dcl9 88-38647<br />
British Library Cataloging-in-Publication Data<br />
available<br />
Manufactured in the United States of America<br />
10 9 8 7 6 5 4 3 2 1<br />
98 97 96 95 94 93 92 91 90 89<br />
°° The paper used in this publication meets the<br />
CIP<br />
minimum requirements of the American National<br />
Standard for Performance of Paper for Printed<br />
Library Materials Z39.48-1984
Contents<br />
1 Introduction<br />
1 HISTORIC BACKGROUND<br />
3 GEOGRAPHIC AREA COVERED IN THIS GUIDE<br />
4 ARRANGEMENT OF THE GUIDE AND HOW TO USE IT<br />
5 ACKNOWLEDGMENTS<br />
7 Glossary of Technical Terms<br />
13 Marine Isopods of the Caribbean<br />
13 ORDER ISOPODA<br />
15 SUBORDER ANTHURIDEA<br />
73 SUBORDER ASELLOTA<br />
107 SUBORDER EPICARIDEA<br />
114 SUBORDER FLABELLIFERA<br />
236 SUBORDER GNATHIIDEA<br />
243 SUBORDER MICROCERBERIDEA<br />
246 SUBORDER ONISCIDEA<br />
251 SUBORDER VALVIFERA<br />
261 Zoogeography<br />
261 FAUNAL PROVINCES<br />
262 ANALYSIS OF THE ISOPOD FAUNA<br />
266 THE BAHAMAS<br />
269 BERMUDA<br />
269 CAVE ISOPODS<br />
275 Appendix<br />
277 Literature Cited<br />
293 Index
Introduction<br />
The title of this work will no doubt raise several questions in many readers'<br />
minds: why the Caribbean? why not the Caribbean and the Gulf of Mexico?<br />
why only the marine isopods? just what is the "Caribbean area"? We hope<br />
that the answers to some of these (and other) questions will become<br />
apparent.<br />
There are several works that already deal with the isopods of the Carib<br />
bean, as part of a wider treatment of North American isopods (e.g.,<br />
Richardson, 1905; Schultz, 1969). Why then this "Isopods of the Carib<br />
bean"? <strong>As</strong> partial answer, the following: many new records of isopods from<br />
the Caribbean region (in its broadest sense) have appeared in scattered pub<br />
lications in the last few decades. The time has come to pull these together in a<br />
single work. The number of marine laboratories in the area has increased,<br />
with more and more students exploring especially the shallow marine en<br />
vironment. A single work on a relatively speciose and abundant group of<br />
invertebrates would be useful to such investigators, as they build up a com<br />
prehensive view of the biology of the region. Concepts of the taxonomy of<br />
several isopod groups have changed radically over the last few years; again,<br />
there is obvious utility in having these changes summarized in a single<br />
source. New species and records are continuously being found. Having a<br />
single baseline work decreases the time needed for investigating and estab<br />
lishing the validity of such records.<br />
HISTORIC BACKGROUND<br />
The history of isopod taxonomic research in the Caribbean really starts with<br />
a worldwide monographic work on the Cirolanidae by Hansen in 1890. In<br />
cluded here were about 12 species from the Danish West Indies, now the<br />
U.S. Virgin Islands. Since then a few major works on Caribbean isopods<br />
have appeared, such as Moore's report on the isopods of Puerto Rico (1901)<br />
and Menzies and Glynn's report on the same area (1968). Some areas have<br />
received considerable attention, such as the aforementioned Puerto Rico and,<br />
more recently, Belize. A list of 116 species of isopods from Cuba (including<br />
Oniscidea) has been published (Ortiz, Lalana, and Gomez, 1987). At the<br />
other extreme, there are no records from a number of localities, especially the<br />
1
Figure 1. Map of area covered by this guide.
INTRODUCTION 3<br />
islands of the southeastern chain of the Lesser Antilles. In total there are<br />
about 40 publications, varying from descriptions of single species to longer<br />
works, that deal with isopods from the Caribbean. These publications will be<br />
encountered in the following guide, under the specific taxa.<br />
GEOGRAPHIC AREA COVERED IN THIS GUIDE<br />
The accompanying map (Figure 1) shows the area for which records are<br />
included in this guide.<br />
While it may seem logical to include the Gulf of Mexico, and while there<br />
are several isopod species common to both areas, this has not been done.<br />
There are relatively few isopod records from the Gulf; undoubtedly a great<br />
deal of taxonomic work awaits the careful collector in this area. Also, from a<br />
zoogeographic point of view, separation of the Gulf may be justified.<br />
Bermuda, on the other hand, situated in the northwestern Atlantic several<br />
hundred miles off the coast of the United States, is included. This island,<br />
although remote from the Caribbean, is swept by waters that earlier have<br />
passed through the Caribbean. Zoogeographically, the shallow-water Ber-<br />
mudan and Caribbean faunas have much in common.<br />
While perhaps not strictly in the Caribbean Sea, the Bahamas and the<br />
Florida Keys are included here, their shallow-water marine faunas being<br />
overwhelmingly Caribbean in nature.<br />
Turning to depths limits, within the area under discussion, species from<br />
the intertidal to 200 meters have been dealt with in some detail. This arbi<br />
trary cutoff depth was selected because most Caribbean isopod species in<br />
habit relatively shallow depths. About 30 species have been recorded from<br />
below 200 meters in the Caribbean, many of these known only from the type<br />
material. A list of species of this very poorly known deeper fauna is included<br />
here. Without doubt, many species in the deeper waters of the Caribbean<br />
await discovery.<br />
A fascinating group of isopods, while not strictly shallow-water marine<br />
forms, is included. These are the true cave forms, found mainly in the sub<br />
orders Anthuridea and Flabellifera. Given the history of the Caribbean from<br />
the Quaternary to the present, it is not surprising that caves are common<br />
throughout the region. These caves may be well inland and contain only<br />
freshwater, but are more commonly anchialine, that is, having some (fre<br />
quently subterranean) link to the sea. Less common, and of lesser interest<br />
from an isopod taxonomist's point of view, are the fully marine caves in<br />
direct communication with the sea or, indeed, under the surface of the sea<br />
itself.
4 INTRODUCTION<br />
ARRANGEMENT OF THE GUIDE AND HOW TO USE IT<br />
A short introduction to the Crustacea Isopoda is provided, followed by a<br />
glossary of descriptive terms and morphological features used throughout the<br />
guide (see Figure 2).<br />
Keys and diagnoses to the suborders and all lower taxa follow. For ease of<br />
usage, except in the keys, all taxa are presented in alphabetical order, regard<br />
less of their phylogenetic relationships.<br />
Diagnoses are provided for all suborders, families, genera, and species.<br />
The only exceptions to this are in the suborders Epicaridea and Oniscidea.<br />
Within each suborder, a key to the families occurring in the Caribbean is<br />
provided. Similarly, within each family and genus, keys are provided to the<br />
relevant genera and species, respectively.<br />
In whatever context, where an author and date appear, a reference to these<br />
is provided in the Literature Cited section. In some cases, reference is made<br />
to useful publications such as revisions of families or genera.<br />
<strong>As</strong> this is not a textbook on the Isopoda, biological information is generally<br />
kept to a minimum. In the case of individual species, however, what little<br />
ecological information is available, is provided. For general texts on biology,<br />
internal anatomy, physiology, and reproduction, the reader is referred to<br />
works such as Kaestner (1967), Waterman (1960), Bliss (1982-1985), and<br />
Schram (1986).<br />
Within each species discussion, a diagnosis is given, along with maximum<br />
(total) middorsal lengths for males and females, where known. The diag<br />
noses are not exhaustive, but provide only the information needed to distin<br />
guish the species. Diagnoses thus vary in length from the statement of a<br />
single feature to a paragraph concerning several features, depending on the<br />
understanding and complexity of the taxonomy of the group. In the longer<br />
diagnoses, morphological features are dealt with in order from anterior to<br />
posterior on the animal's body. Records are given, rather than geographical<br />
distribution, as our knowledge of many species is woefully incomplete. These<br />
records are given in a roughly north-to-south order; records outside of the<br />
Caribbean region, as here defined, are given on a separate line. A few species<br />
not yet recorded from the Caribbean are included, in the strong likelihood<br />
that they will eventually be found here. The records include depth distribu<br />
tion information in meters, where known. Records were taken from pub<br />
lished papers; in addition, the collections of the United States National<br />
Museum of Natural History, Smithsonian Institution, were scoured, and<br />
many unpublished records from this source are also included. In the "Re<br />
marks" section, ecological information such as substrate preferences is given.<br />
Hosts of parasitic species are given. Formal synonymies are not provided,
INTRODUCTION 5<br />
but nomenclatural comments are included in the few cases where a species<br />
may be known under a more commonly used name. Usually, a figure of the<br />
entire animal of each species is given. Diagnostic features are usually illus<br />
trated. Unless otherwise stated, all illustrations are original and by the au<br />
thors, and were made from actual specimens.<br />
Common and scientific names of fishes that are hosts to parasitic isopods<br />
are taken from the American Fisheries Society special publication no. 12<br />
(Robins et al., 1980).<br />
Finally, a word of warning. Difficulties may be experienced in using the<br />
keys, for which there may be any of several reasons: characters seen in the<br />
animal may not clearly conform to those in the key (in which case refer to the<br />
figures, as well as to good recent descriptions or diagnoses); your material<br />
may be a new record for the region; or you may have an undescribed species<br />
(in which case refer to more comprehensive treatments of the group).<br />
ACKNOWLEDGMENTS<br />
Much of the material covered in this work comes from the many collectors<br />
who have deposited specimens from several regions of the Caribbean in the<br />
collections of the National Museum of Natural History, Smithsonian Institu<br />
tion. Rather than risk the unwitting omission of a name, we thank all of these<br />
individuals collectively. Without their efforts, our knowledge of the Carib<br />
bean fauna would be the poorer.<br />
Material was borrowed from several institutions. We thank the following<br />
scientists for their assistance in this connection: Jan Stock and Dirk Platvoet<br />
of the Instituut voor Taxonomische Zoologie, University of Amsterdam;<br />
Jean Just and Torben Wolff of the Zoological Museum, University of<br />
Copenhagen; Jacques Forest of the Museum National d'Histoire Naturelle,<br />
Paris; Richard Heard of the Gulf Coast Research Laboratory, Ocean<br />
Springs, Mississippi; Willard Hartman of the Peabody Museum of Natural<br />
History, Yale University; Herbert W. Levi of the Museum of Comparative<br />
Zoology, Harvard University; Harold S. Feinberg of the American Museum<br />
of Natural History, New York; John E. Miller of the Harbor Branch Founda<br />
tion, Florida; and Paula M. Mikkelsen of the Indian River Coastal Museum<br />
at Fort Pierce, Florida.<br />
Bruce Collette of the National Marine Fisheries Laboratory at the<br />
Smithsonian Institution assisted with fish names used in this work, for which<br />
we are grateful.<br />
We thank the staff of the Scanning Electron Microscope Laboratory of the<br />
Smithsonian Institution, and especially Susann Braden, who produced the<br />
electron micrographs used here.
6 INTRODUCTION<br />
We are grateful to the Smithsonian Research Opportunities fund, admin<br />
istered by David Challinor, then <strong>As</strong>sistant Secretary for Research, and to the<br />
Smithsonian's Caribbean Coral Reef Ecosystems program administered by<br />
Klaus Riitzler of the Department of Invertebrate Zoology, for the funding of<br />
several fieldtrips to the Caribbean. The second author acknowledges a finan<br />
cial award from the Smithsonian's Women's Committee for a grant to facili<br />
tate fieldwork.<br />
Several individuals have provided encouragement, advice, suggestions,<br />
critical comments, and missed references, all of which have vastly improved<br />
this work. In this regard we are especially grateful to Thomas E. Bowman, C.<br />
W. Hart, Jr., Horton H. Hobbs, Jr., and Molly K. Ryan, all of the Depart<br />
ment of Invertebrate Zoology, National Museum of Natural History,<br />
Smithsonian Institution; Dan Adkison, Bass Harbor, Maine (who provided<br />
considerable assistance with the Epicaridea); Richard Heard of the Gulf<br />
Coast Research Laboratory, Mississippi; and Paul Delaney of the Los An<br />
geles County Museum of Natural History.<br />
This is Contribution Number 248 of the Caribbean Coral Reef Ecosystems<br />
(CCRE) program, Smithsonian Institution, supported in part by the Exxon<br />
Corporation.
Glossary of Technical Terms<br />
AESTHETASC. Thin-walled sensory seta usually found on flagellum of<br />
antennule<br />
AMBULATORY (as applied to pereopods). Used for walking.<br />
ANCHIALINE. An aqueous habitat near the sea; referring to saltwater or brackish<br />
pools fluctuating with the tides, but with no surface connection to the sea.<br />
ANGULATE. Having an angle or an angular shape.<br />
ANTENNA. Paired appendage of the third cephalon segment; sometimes referred<br />
to as antenna 2.<br />
ANTENNULE. Paired appendage of the second cephalon segment; sometimes<br />
referred to as antenna 1.<br />
APICAL. Relating to the apex or tip.<br />
APPENDAGE. An articulated structure used for feeding, locomotion, sensory<br />
reception, e.g., mouthparts, antennae, pereopods, pleopods, uropods.<br />
ARTICLE. A single section of an appendage, with an articulation at one or both<br />
ends.<br />
BASIS. Article of appendage adjoining coxa proximally, and carrying endopod<br />
distally, i.e., article 2 of pereopod.<br />
BIARTICULATE. Composed of two articles.<br />
BIDENTATE. Having two teeth.<br />
BIFID. Divided into two lobes or parts by a cleft.<br />
BILOBED. Composed of two lobes.<br />
BIRAMOUS. Composed of two rami or branches.<br />
BIUNGUICULATE. Having two claws, as in a bifid dactylus.<br />
CARINA. A keel, or an acute ridge.<br />
CARINATE. Having one or more carinae or acute ridges.<br />
CARPOCHELATE. Having a chela or pincerlike structure formed by the seventh<br />
(dactylus) and fifth (carpus) articles of an appendage.<br />
CARPUS. Article 5 of pereopod.<br />
CEPHALON. Anterior region of body or head; more correctly the cephalothorax<br />
in isopods, as the first pereonal segment is usually fused with the head.<br />
CHELA. Distal pincerlike part of appendage, often formed by a mobile and an<br />
immobile finger.<br />
CHELATE. Having a chela; modified to form a pincer.<br />
CLAVATE. Club shaped; having one end thickened.<br />
CLYPEUS. Platelike structure of cephalon, anterior to upper lip or labrum,<br />
sometimes fused with frontal lamina.<br />
CONGLOBATE. Able to roll up into a ball, as in some sphaeromatid and<br />
oniscidean isopods.<br />
CONSPECIFIC. Belonging to the same species.<br />
CONTIGUOUS. Touching.
frontal lamina<br />
clypeus<br />
labrum<br />
peduncle<br />
uropod<br />
flageHum<br />
I<br />
rrjn<br />
antennule<br />
14<br />
> * antenna<br />
maxi<br />
maxilla 2<br />
pereopods<br />
ischium<br />
Vy —^ carpus<br />
merus<br />
pleopods<br />
propodus<br />
maxilliped<br />
dactylus<br />
F i g u r e 2 . S c h e m a t i c r e p r e s e n t a t i o n o f a n i s o p o d i l l u s t r a t i n g m o r p h o l o g i c a l t e r m s .
GLOSSARY 9<br />
COPULATORY STYLET. Structure situated on endopod of pleopod 2 in males,<br />
used for transfer of spermatophore in some species; also referred to as appendix<br />
masculina.<br />
CORDATE. Heart shaped in outline.<br />
COXA. Basal article of an appendage, attached to sternite, sometimes expanded<br />
into a lateral coxal plate.<br />
CRENULATE. Having a scalloped edge with rounded teeth, usually used to refer<br />
to the margin of a structure.<br />
DACTYLUS. Terminal (7th) article of a pereopod or thoracic appendage.<br />
DENTATE. Edged with teeth.<br />
DENTICLE. A small tooth.<br />
DENTICULATE. Having fine teeth.<br />
DIGITIFORM. Fingerlike.<br />
DISTAL. Situated away from the base or point of origin or attachment.<br />
ECDYSIS. Molting of the integument.<br />
EMARGINATE. Having the margin concave.<br />
ENDITE. Medially directed lobe of coxa or basis of an appendage, especially the<br />
maxilliped.<br />
ENDOPOD. Inner ramus of a biramous appendage.<br />
ENTIRE. Complete; usually referring to the margin of a structure that is smooth.<br />
EPIMERON. Lateral part of a somite.<br />
EPIPOD. Lateral extension of a protopodite.<br />
EXCAVATE. Hollowed out.<br />
EXOPOD. Outer ramus of a paired appendage.<br />
FALCATE. Sickle shaped; curved and tapering to a point.<br />
FLAGELLUM. Distal part of antenna or antennule, usually multiarticulate,<br />
occasionally reduced to one or a few articles.<br />
FRONTAL LAMINA. Platelike structure of the cephalon immediately anterior to,<br />
and sometimes fused with, clypeus.<br />
GENICULATE. Bent at an abrupt angle, as in the body of many arcturid isopods.<br />
GRANULATE. Having the appearance of bearing beadlike or grainlike<br />
protuberances; usually applied to the description of a surface.<br />
HIRSUTE. Bearing hairs (elongate hairs in the case of most isopods).<br />
HYPOGEAN. Underground.<br />
IMMERSED. Sunken into, as with one structure into another.<br />
INCISOR. Cutting process of the mandible, usually dentate, sometimes modified<br />
for piercing.<br />
INDURATE. Hardened, usually by calcium carbonate or sclerotized protein.<br />
INTEGUMENT. Outer covering, e.g., the exoskeleton.<br />
INTERSTITIAL. Relating to interstices; living in the interstices of sand grains,<br />
gravel, or rubble.<br />
ISCHIUM. Article 3 of pereopod.<br />
LABIUM. Lower lip; usually consisting of a pair of lobes posterior to the mouth.<br />
LABRUM. Unpaired projection anterior to mouth, attached to the clypeus; upper<br />
lip.<br />
LACINIA MOBILIS. Small, usually toothed, process articulating at base of incisor<br />
in left or both mandibles.<br />
LAMELLAR. In the shape or structure of a thin plate or lamella.
10 GLOSSARY<br />
LAMINA DENTATA. Serrate platelike structure in the mandible of anthurideans,<br />
formed by the fusion of spines of the spine-row.<br />
LANCEOLATE. Lance shaped; narrow and tapering to a point.<br />
LINGUIFORM. Tongue shaped.<br />
MANGA. Young of some peracaridean crustaceans (including isopods), lacking<br />
last thoracic appendage at time of release from broodpouch.<br />
MANDIBLE. First pair of mouthparts, functioning as jaws, often sclerotized.<br />
MARSUPIUM. Structure in which eggs are retained by female; the broodpouch.<br />
MAXILLA (1 and 2). Two sets of paired mouthpart appendages immediately<br />
posterior to mandible.<br />
MAXILLIPED. First paired appendage of the thorax; usually incorporated into the<br />
mouthparts.<br />
MEDIAN. At, near, or directed toward the middle or midline.<br />
MERUS. Article 4 of pereopod.<br />
METAMORPHOSED. Transformed; changed in appearance, structure, or function.<br />
MESIAL. Near or toward the middle or midline.<br />
MOLAR. Grinding, and sometimes piercing, structure of the mandible.<br />
MULTIARTIGULATE. Composed of many articles.<br />
NATATORY. Adapted for swimming.<br />
OBSOLETE. Becoming vestigial, and losing original function.<br />
OMMATIDIA. Individual visual components of the compound eye.<br />
OOSTEGITE. Medially directed lamellar structure arising from coxa of pereopod<br />
in the female, forming part of the broodpouch or marsupium.<br />
OPERCULIFORM. In the form of a cover or lid.<br />
OVATE. Egg shaped or oval.<br />
PALM. Cutting edge of the propodus, often defined proximally by a spine, in a<br />
subchelate appendage.<br />
PALP. Articulated ramus consisting of one to three articles in mandible, of up to<br />
five articles in the maxilliped.<br />
PECTINATE. Having teeth like a comb.<br />
PEDUNCLE. Stalk or proximal part of an appendage, as in antennae.<br />
PENIAL RAMI. Paired submedian process on sternite 7 of male.<br />
PEREON. Middle or thoracic region of the body, consisting of seven segments or<br />
pereonites, first fused with cephalon in isopods.<br />
PEREONITE. Segment of the pereon.<br />
PEREOPOD. Paired appendage of the pereon, consisting of seven articles when<br />
unmodified.<br />
PILOSE. Covered with short hairs or setae.<br />
PLEON. Posterior or abdominal region of the body, primitively consisting of six<br />
segments or pleonites, and bearing paired pleopod and uropod appendages.<br />
PLEONITE. Segment of the pleon.<br />
PLEOPOD. Paired appendage of the pleon, five pairs being present in the<br />
primitive condition.<br />
PLEOTELSON. Structure resulting from the fusion of the telson and one or more<br />
pleonal segments.<br />
PLICATE. Pleated or folded.<br />
PRANIZA. Juvenile, immature stage of gnathiideans.<br />
PREHENSILE. Adapted for holding or clinging.
PRODUCED. Extended or lengthened.<br />
PROPODUS. Article 6 of pereopod.<br />
GLOSSARY 11<br />
PROTANDROUS. In hermaphroditic forms, becoming a functional male producing<br />
spermatozoa before becoming a functional female producing eggs.<br />
PROTOGYNOUS. In hermaphroditic forms, becoming a functional female<br />
producing eggs before becoming a functional male producing spermatozoa.<br />
PROTOPODITE. Proximal part of an appendage, consisting of the coxa and basis.<br />
PROXIMAL. Situated near the point of attachment.<br />
PYLOPOD. First pereopod of the Gnathiidea, modified to form part of the<br />
mouthparts.<br />
RAMUS. Branch of an appendage.<br />
RENIFORM. Kidney shaped.<br />
RETICULATE. Resembling or forming a network.<br />
RETINACULAE. Small hooks on an appendage, used to link the left and right<br />
members of a pair of appendages.<br />
ROSTRUM. Anterior middorsal projection of cephalon.<br />
SAGITTATE. Arrow shaped.<br />
SGLEROTIZED. Hardened, usually with chitin.<br />
SERRATE. Edged with toothlike projections as in a saw.<br />
SETIFEROUS. Bearing setae.<br />
SETOSE. Bearing setae.<br />
SINUATE. Having a wavy margin.<br />
SINUOUS. Having curves.<br />
SOMITE. Body segment, usually having a pair of appendages.<br />
SPATULATE. Shaped like a spatula.<br />
SPICATE. Shaped like a spike.<br />
SPINE-ROW. Row of spines situated between the incisor and molar processes of<br />
the mandible.<br />
SPINOSE. Bearing spines.<br />
STATOCYST. Small saclike sensory organ, often containing granules, used to<br />
indicate to the animal its orientation.<br />
STYGOBIONT. Cave organism.<br />
STYLIFORM. Having a long, slender, stilettolike shape.<br />
SUB-. A prefix indicating ''almost" or "just less than," e.g., submarginal—almost<br />
on the margin.<br />
SUBGHELATE. Having a subchela, forming a pincerlike structure, especially by<br />
the dactylus folding back on the propodus.<br />
SUTURE. A line indicating an area of articulation, or of incomplete fusion.<br />
SYMPOD. Proximal part of an appendage, often formed by the fusion of the coxa<br />
and basis.<br />
TELSON. Terminal part of the body, usually bearing the anus.<br />
THORAX. Tagma or body region between the cephalon and the abdomen.<br />
TRAGHEATE. Bearing tubular respiratory trachea (more correctly pseudotrachea)<br />
on pleopods, as in Oniscidea.<br />
TRICUSPID. Bearing three cusps or points.<br />
TRIDENTATE. Having three teeth.<br />
TRIFID. Divided into three parts or lobes.<br />
TRILOBED. Divided into three lobes.
12 GLOSSARY<br />
TRISINUATE. Having three curves.<br />
TRIUNGUIGULATE. Bearing three claws, as in a trifid dactylus.<br />
TRUNCATE. Having the appearance of having been abruptly cut off.<br />
TUBERCULATE. Bearing knoblike or wartlike prominences or tubercles.<br />
UNIARTIGULATE. Composed of one article.<br />
UNIRAMOUS. Having one ramus or branch.<br />
UNIUNGUICULATE. Having a single claw, as in a dactylus.<br />
UROPOD. Paired pleonal appendage of the last pleonite, usually situated at the<br />
base of the telson.
Marine Isopods of the Caribbean<br />
Phylum Arthropoda<br />
Superclass Crustacea Pennant, 1777<br />
Class Malacostraca Latreille, 1806<br />
Subclass Eumalacostraca Grobben, 1892<br />
Superorder Peracarida Caiman, 1904<br />
Order Isopoda Latreille, 1817<br />
DIAGNOSIS Body usually dorsoventrally depressed, occasionally sub-<br />
cylindrical, rarely bilaterally compressed. Carapace lacking. Antennules and<br />
antennae uniramous (scale on antenna in some asellotes may represent rudi<br />
mentary second ramus). Eyes sessile (although situated on nonmobile stalks<br />
in some asellotes). Mouthparts consisting of one pair of mandibles, two pairs<br />
of maxillae, one pair of maxillipeds; latter appendages of first thoracic seg<br />
ment fused with cephalon. Mandible usually with palp consisting of one to<br />
three articles; incisor, lacinia mobilis, and molar usually present; lacinia mo-<br />
bilis often differing on left and right sides, sometimes absent from right man<br />
dible; molar variable. Maxilliped usually consisting of palp of no more than<br />
five articles, lamellar endite often with coupling hooks, lamellar epipod. Per-<br />
eonites usually separate, although pereonite 1 sometimes fused with<br />
cephalon. Coxae of pereopods variously fused with, and forming expanded<br />
lateral processes of, pereonites. Pereopod 1 forming additional mouthpart<br />
(pylopod) only in Gnathiidea. Pereopods generally similar, ambulatory;<br />
pereopods 1-3 secondarily variously modified and becoming subchelate or<br />
prehensile; pereopods 4-7 occasionally modified, becoming natatory or pre<br />
hensile. Pereopod 7 occasionally not developed (neotenous condition).<br />
Broodpouch or marsupium formed by varying number of oostegites attached<br />
ventrally and medially to coxae of pereopods; eggs held in anterior or pos<br />
terior pockets or internal pouches in gnathiids and some sphaeromatids.<br />
Pleon consisting of six pleonites, free or variously fused, plus telson; if one or<br />
more pleonites fused with telson, resulting structure referred to as pleotelson.<br />
13
14 1SOPODA<br />
Key to suborders of Isopoda<br />
1. Parasitic on crustaceans; body of 9 nearly always asymmetrical<br />
Epicaridea<br />
Free-living or parasitic on fishes; body of 9 bilaterally symmetrical, or<br />
if parasitic, 9 somewhat distorted 2<br />
2. Body more or less bilaterally compressed Phreatocoidea<br />
Body more or less dorsoventrally depressed or subcylindrical 3<br />
3. With six pereonites and five pairs of pereopods Gnathiidea<br />
With seven pereonites and six or seven pairs of pereopods 4<br />
4. Body usually more than six times longer than wide, subcylindrical,<br />
uropods never operculiform 5<br />
Body usually less than six times longer than wide, usually<br />
dorsoventrally depressed; if subcylindrical, uropods operculiform . . 6<br />
5. Uropodal exopod often folding dorsally over pleotelson; rarely<br />
interstitial forms : Anthuridea ^IC<br />
Uropods terminal, exopod lacking; minute interstitial forms<br />
6. Antennules minute; terrestrial forms, with pleopods tracheate<br />
Microcerberidea<br />
Oniscidea<br />
Antennules rarely minute; aquatic forms, pleopods never tracheate . . 7<br />
7. Uropods ventral, operculiform, covering pleopods Valvifera r'<br />
Uropods never operculiform over pleopods 8<br />
8. Uropods lateral or ventrolateral, forming tailfan with pleotelson;<br />
pleopods 1 and 2 rarely operculiform Flabellifera(? I v<br />
Uropods terminal or subterminal; pleopods 1 and 2 variously<br />
operculiform <strong>As</strong>ellota ,•;••-<br />
The suborder Phreatocoidea contains freshwater forms, and has a<br />
Gondwanian distribution, primarily in the Southern Hemisphere.<br />
Pleopods (on pleonites 1—5) biramous, lamellar, primarily for respiration;<br />
anterior pleopods occasionally operculiform. Pleopod 2 in male (and occa-
ANTHURIDEA 15<br />
sionally also pleopod 1 in Oniscidea and <strong>As</strong>cllota) with endopod bearing<br />
copulatory stylet. One pair of uropods on pleonite 6. Young leave brood-<br />
pouch as manca, i.e., resembling adult but lacking pereopod 7; in Epi-<br />
caridea, manca stage represented by epicaridium stage; latter transforms<br />
into microniscium and then cryptoniscium stage, before becoming adult.<br />
Suborder Anthuridea Leach, 1814<br />
DIAGNOSIS Body generally elongate and subcylindrical. Eyes absent in<br />
some genera. Antennular peduncle of three articles; antennal peduncle of five<br />
articles. Mandible with palp of one to three articles, or absent; body of mand<br />
ible either styliform and lacking molar and lacinia mobilis, or with molar<br />
variously specialized or reduced, lacinia mobilis absent, and spine-row modi<br />
fied to form platelike lamina dentata. Maxilla 1 with inner ramus reduced,<br />
outer ramus slender. Maxilla 2 rudimentary. Maxilliped variable, with palp<br />
of one to five articles, endite present, modified, reduced, or absent. Pereonite<br />
1 free. Pereopod 1, or pereopods 1-3 subchelate; pereopods 4-7 generally<br />
ambulatory. Pleonites 1-5 free or fused, pleonite 6 partly or completely fused<br />
with telson. Pleopods 1-5 similar, or pleopod 1 variously modified to form<br />
operculum. Uropodal exopod often folded dorsally over pleotelson.<br />
Pleotelson with pair of statocysts, with single statocyst, or lacking statocysts.<br />
REMARKS Protogyny has been demonstrated in several species of An<br />
thuridea. The order of development in these cases is: egg, manca (both in the<br />
broodpouch), immature subadult, ovigerous female, premale, male, with<br />
varying numbers of molts between each stage. At least one molt takes place<br />
between ovigerous female and premale, the latter being distinguished by the<br />
loss of the oostegites and by the elongation of, and acquisition of more flagel<br />
lar articles in, the antennule. One or two molts take place between premale<br />
and sexually mature male, the latter being characterized by the possession of<br />
elongate antennular flagella bearing dense whorls of aesthetascs, a more set<br />
ose and/or spinose pereopod 1, and sometimes by an elongation of the pleon<br />
and uropods. In some genera, the males have somewhat atrophied<br />
mouthparts, suggesting that they do not feed at this stage. <strong>As</strong> a result of this<br />
seemingly widespread protogyny, sex ratios are strongly biased toward<br />
females, and in several species males are not yet known.<br />
The number of families in the suborder Anthuridea has not been settled.<br />
At present, three families are recognized. Doubtless, further families will be<br />
defined and the genera reshuffled.
16 ANTHURIDEA • ANTHUR1DAE<br />
Key to families of Anthuridea<br />
1. Mouthparts adapted for piercing and sucking, together forming<br />
conelike structure Paranthuridae p£i<br />
Mouthparts adapted for cutting, lamina dentata and molar usually<br />
present 2<br />
2. Pereopod 1 subchelate, with propodus expanded; pleonites generally<br />
fused; if free, much shorter than wide Anthuridae A<br />
Pereopods 1—3 subchelate, subsimilar; pleonites free, often as long as<br />
wide Hyssuridae pC&<br />
Family Anthuridae Leach, 1814<br />
DIAGNOSIS Mouthparts adapted for cutting. Pereopod 1 usually markedly<br />
different from remaining pereopods, subchelate with propodus more or less<br />
inflated. Exopod of pleopod 1 operculiform, covering remaining pleopods.<br />
Pleonites 1—5 fused, with fusion marked ventrolateral^ by short slits, occa<br />
sionally with dorsal grooves marking lines of fusion, or free; if free, length of<br />
each pleonite much less than width. Pleotelson with pair of statocysts, or<br />
single medial statocyst, or lacking statocysts.<br />
Key to genera of Anthuridae<br />
1. Pleopod 1, both rami contributing to operculum 2<br />
Pleopod 1, only exopod operculiform 6<br />
2. Antennal peduncle bearing serrate process Licranthura M><br />
Antennal peduncle lacking any serrate process 3<br />
3. Pleopod 1 in 9, rami to some degree fused Eisothistos ?Z<br />
Pleopod 1 in 9, rami free 4<br />
4. Pereopods 1 and 2 subchelate, of similar size, propodi not noticeably<br />
inflated 5<br />
Pereopod 1 much larger and propodus more expanded, than<br />
pereopod 2 Minyanthura
Amakusanthura 17<br />
5. Integument noticeably pitted; mandible on one side lacking molar, on<br />
other side with spicate molar Apanthuroides P- 2°<br />
Integument not pitted; mandibles similar on both sides Chalixanthura p.
18 ANTHURIDEA • ANTHURIDAE<br />
illipedal palp of three articles; endite small or absent. Pereopod 1, propodal<br />
palm with step or tubercle. Pereopods 4-7, carpus triangular. Pleonites<br />
short, pleonites 1-4 fused, boundaries demarked by complete dorsal folds,<br />
pleonites 4 and 5 separated only by lateral fold, not demarked dorsally;<br />
pleonite 6 dorsally demarked from telson. Pleopod 1 exopod operculiform.<br />
Uropodal exopod often notched or excavate distally. Pleotelson with two<br />
basal statocysts.<br />
Key to species of Amakusanthura<br />
1. Telson thickened, with raised area at midlength, widening and sloping<br />
posteroventrally; uropodal exopod distally shallowly excavate<br />
magnified<br />
Telson dorsally flat, not thickened; uropodal exopod distinctly notched<br />
or barely excavate distally 2<br />
2. Integument pigmented; uropodal endopod length 1.5 times basal width<br />
signata<br />
Integument not pigmented 3<br />
3. Uropodal exopod distinctly notched 4<br />
Uropodal exopod, outer margin weakly excavate; mandibular palp<br />
article 3 bearing three spines lathridia<br />
4. Mandibular palp article 3 bearing three spines signified<br />
Mandibular palp article 3 bearing two spines geminsula<br />
Amakusanthura geminsula (Kensley, 1982)<br />
Figure 3A-E<br />
DIAGNOSIS 9 : 8.1 mm. Shallow middorsal pit on pereonites 4-6, appearing<br />
chalky white in life. Antennular flagellum of three articles. Antennal<br />
flagellum of two articles. Mandibular palp article 3 bearing two spines. Max-<br />
illiped with short slender endite; terminal palp article set obliquely at outer<br />
distal angle of penultimate article. Pereopod 1, propodal palm with step;<br />
carpus distally a rounded lobe. Uropodal exopod with notch. Pleotelson<br />
tapering in posterior half to subtruncate apex. 6: 4.8 mm. Eyes larger than<br />
9. Antennular flagellum of 8-9 articles. Pereopod 1, carpus with distal lobe
Amakusanthura geminsula 19<br />
Figure 3. Amakusanthura geminsula: Ay 9; By pereopod 1; Cy telson; Dy uropodal<br />
sympod and endopod; E> uropodal exopod. Amakusanthura lathridia: F, pereopod 1;<br />
Gy telson; H, uropodal exopod; /, uropodal sympod and endopod.<br />
narrowly rounded; propodal palm with rounded tubercle; irregular band of<br />
setae on mesial surface.<br />
RECORDS Carrie Bow Cay and Twin Cays, Belize, intertidal to 1.5 m;<br />
Jamaica.
20 ANTHURIDEA • ANTHUR1DAE<br />
Amakusanthura lathridia (Wagele, 1982)<br />
Figure 3F-I<br />
DIAGNOSIS 9 2.7 mm. Antennular flagellum of three articles. Antennal<br />
flagellum of three articles. Mandibular palp article 3 bearing three spines.<br />
Maxilliped lacking endite. Pereopod 1, propodal palm with distal step;<br />
carpus with triangular posterodistal lobe. Uropodal exopod apically acute,<br />
outer margin slightly excavate, sinuate; endopod length slightly less than<br />
twice basal width. Pleotelson elliptical, widest at about midlength, margins<br />
weakly serrate.<br />
RECORDS Cuba, interstitial beach sand just above water line.<br />
Amakusanthura magnifica (Menzies and Frankenberg, 1966)<br />
Figures 4, 5<br />
DIAGNOSIS 9: 13.9 mm. Maxilliped with short endite, terminal article of<br />
palp short, set obliquely at outer distal angle of penultimate article. Pereopod<br />
1, propodal palm with strong tubercle; carpus with low distal sclerotized<br />
lobe. Uropodal exopod ovate, with shallow distal notch; endopod twice<br />
longer than wide. Pleotelson with posterior margin evenly rounded to sub-<br />
truncate; dorsally with raised area at midlength, broadening and sloping<br />
B<br />
D<br />
C<br />
Figure 4. Amakusanthura magnifica: A><br />
pereopod 1; B, telson; C, uropodal sympod<br />
and endopod (setae omitted); D> uropodal<br />
exopod (setae omitted)
Figure 5. Amakusanthura magnijica: A,<br />
p l e o n i t e s i n l a t e r a l v i e w ; B } p l e o n i n d o r s a l<br />
v i e w .<br />
Amakusanthura signata 21<br />
a w a y p o s t e r i o r l y . 6 : 1 0 . 3 m m . A n t e n n u l a r f l a g e l l u m o f 2 4 a r t i c l e s . P e r e o p o d<br />
1 , c a r p u s w i t h s c l e r o t i z e d d i s t a l r o u n d e d l o b e ; p r o p o d a l p a l m w i t h s t r o n g<br />
s c l e r o t i z e d t u b e r c l e l o n g e r t h a n i n 2 ; n u m e r o u s s e t a e o n m e s i a l s u r f a c e .<br />
P l e o p o d 2 , c o p u l a t o r y s t y l e t o f e n d o p o d n o t r e a c h i n g b e y o n d r a m u s .<br />
R E C O R D S O f f G e o r g i a , 1 7 - 1 3 7 m ; o f f F l o r i d a , 7 - 1 1 m ; C u b a ; G u l f o f<br />
M e x i c o .<br />
A m a k u s a n t h u r a s i g n a t a ( M e n z i e s a n d G l y n n , 1 9 6 8 )<br />
F i g u r e 6 A - E<br />
D I A G N O S I S 2 : 4 . 9 m m . I n t e g u m e n t w i t h s t r o n g p a t c h e s o f p i g m e n t o n<br />
c e p h a l o n ; p i g m e n t s p a r s e o n p e r e o n i t e s ; t w o b a r s o n p l e o n i t e 6 . A n t e n n u l a r
22 ANTHURIDEA • ANTHURIDAE<br />
Figure 6. Amakusanthura signata: A, 9; B, telson; C, pereopod 1; D, uropodal<br />
sympod and endopod; E> uropodal exopod. Amakusanthura signijica: F, telson; G,<br />
uropodal sympod and endopod; H, uropodal exopod; I, pereopod 1.<br />
flagellum of three articles. Antennal flagellum of three articles. Mandibular<br />
palp article 3 bearing three spines. Maxilliped with short endite not reaching<br />
base of palp article 2. Pereopod 1, carpus with strong subacute sclerotized<br />
lobe distally; propodal palm with transparent convex flange and step.
Anthomuda stenotelson 23<br />
Uropodal exopod with strong distal notch; endopod length about 1.5 times<br />
basal width. Pleotelson widest in posterior half, tapering posteriorly to shal-<br />
lowly notched apex. 6: 4.5 mm. Eyes larger than in 9 . Antennular flagellum<br />
of nine articles. Pereopod 1, carpus with strong distal lobe, propodal palm<br />
with step, convex transparent flange, and numerous setae on mesial surface.<br />
Pleopod 2, copulatory stylet of endopod reaching well beyond ramus.<br />
RECORDS Cuba; Puerto Rico, intertidal to 1.5 m; Carrie Bow Cay, Belize,<br />
intertidal to 24 m.<br />
Amakusanthura significa (Paul and Menzies, 1971)<br />
Figure 6F—I<br />
DIAGNOSIS 9 5.0 mm. Antennular flagellum of three articles. Antennal<br />
flagellum of four or five articles. Mandibular palp article 3 bearing three<br />
spines. Maxilliped lacking endite. Pereopod 1, propodal palm with step;<br />
carpus distally rounded. Uropodal exopod ovate, with distal notch; endopod<br />
length slightly more than twice basal width. Telson elliptical, posterior mar<br />
gin narrowly rounded.<br />
RECORDS Off Venezuela, 95 m.<br />
Anthomuda Schultz, 1979<br />
DIAGNOSIS Eyes present. Mandibular palp of three articles. Maxillipedal<br />
palp of four articles; endite reaching well beyond base of palp article 2. Per-<br />
eopods 1 and 2 similar, subchelate. Pleonites 1-6 short, free; pleonite 6 dor-<br />
sally demarked. Telson with single semiovate hollow representing statocyst.<br />
Anthomuda stenotelson Schultz, 1979<br />
Figure 7A<br />
DIAGNOSIS 9 8.8 mm. Antennular flagellum of three or four articles. An<br />
tennal flagellum of six articles. Pereopods 1 and 2 similar, propodi somewhat<br />
inflated; pereopod 1, propodal palm straight, unarmed; pereopod 2 propodal<br />
palm with three sensory spines. Pereopods 4—7 with rectangular carpi.<br />
Pleonite 6 free, posterior margin broadly bilobed. Uropodal exopod elongate-<br />
elliptical, distally narrowly rounded; endopod length 2.5 times basal width,<br />
distally broadly rounded. Telson narrowly lanceolate, posteriorly narrowly<br />
rounded, with single open hollow statocyst.<br />
RECORDS Off Bermuda, 90 m.
Figure 7. Anthomuda stenotelson: A, 9. Apanthura cracenta: B, 9; C, telson; D,<br />
uropodal endopod and sympod; E, uropodal exopod. Apanthura crucis: F, uropodal<br />
exopod; G, uropodal endopod and sympod. Apanthura harringtoniensis (from Wagele,<br />
1981): H, uropodal endopod and sympod; /, uropodal exopod.
Apanthura Stebbing, 1900<br />
Apanthura cracenta 25<br />
DIAGNOSIS Integument sometimes pigmented. Eyes present. 9: Antennu<br />
lar flagellum of three articles. Antennal flagellum of two to four articles.<br />
Mandibular palp of three articles, terminal article bearing distal spines.<br />
Maxillipedal palp of three articles; endite small, or lacking. Pereopod 1, pro-<br />
podal palm usually with step or tubercle; propodus inflated. Pereopods 4-7,<br />
carpus triangular. Pleonites 1-5 fused; pleonite 6 dorsally demarked.<br />
Pleopod 1, exopod operculiform. Uropodal exopod ovate, sometimes distally<br />
notched or excavate. Pleotelson with two basal statocysts. 6 antennular<br />
flagellum of about 10 articles.<br />
REMARKS Differentiation of species, especially if long preserved and the<br />
pigmentation is lost, depends on subtle features of the mandible, maxilliped,<br />
pereopods, and pleotelson.<br />
Key to species of Apanthura<br />
1. Uropodal exopod distally notched or excavate 2<br />
Uropodal exopod distally entire; uropodal endopod length subequal to<br />
width cruris<br />
2. Uropodal exopod distally notched; uropodal endopod length 2.4 times<br />
basal width cracenta<br />
Uropodal exopod faintly excavate or sinuate; uropodal endopod length<br />
about twice basal width harringtoniensis<br />
Apanthura cracenta Kensley, 1984<br />
Figure 7B-E<br />
DIAGNOSIS 9: 4.6 mm. Antennular flagellum of three articles. Antennal<br />
flagellum of two articles. Mandibular palp article 3 bearing four spines.<br />
Maxilliped with short rounded endite; terminal palp article set at oblique<br />
angle on, and less than half length of penultimate article. Pereopod 1, carpus<br />
triangular with acute sclerotized tip overlapping base of palm; propodal<br />
palm with rounded tubercle near midlength. Uropodal exopod deeply<br />
notched; endopod ovate, length 273 basal length. Pleotelson lanceolate. 6:<br />
3.8 mm. Antennular flagellum of six articles. Antennal flagellum of two arti<br />
cles. Pleopod 2, copulatory stylet of endopod reaching by half its length be<br />
yond ramus.
26 ANTHUR1DEA • ANTHURIDAE<br />
RECORDS Turks and Caicos Islands, 1 m; Carrie Bow Cay, Belize, intcrti-<br />
dal on reef crest, 2 m.<br />
Apanthura crucis (Barnard, 1925)<br />
Figure 7F,G<br />
DIAGNOSIS 9: 5.9 mm. Antennular flagellum of four articles. Antennal<br />
flagellum of four articles. Mandibular palp article 3 with eight spines. Max-<br />
illiped lacking endite; terminal palp article more than half length of penulti<br />
mate article. Pereopod 1, propodal palm with low rounded tubercle at mid-<br />
length. Uropodal exopod ovate, outer margin evenly convex; endopod<br />
subcircular. Pleotelson anteriorly narrow, widest at midlength, posterior<br />
margin broadly rounded. 6: 6.2 mm. Antennular flagellum of 12 or 13 arti<br />
cles. Antennal flagellum of four articles. Pereopod 1, propodal palm with low<br />
rounded tubercle; dense band of setae on mesial surface. Copulatory stylet on<br />
endopod of pleopod 2 reaching a third of its length beyond ramus.<br />
RECORDS Turks and Caicos Islands, 1 m; St. Croix, U.S. Virgin Islands, 8<br />
m.<br />
Apanthura harringtoniensis Wagele, 1981<br />
Figure 7H,I<br />
DIAGNOSIS 9 6.0 mm. Antennular flagellum of four articles. Antennal<br />
flagellum of four articles. Mandibular palp article 3 bearing two spines.<br />
Maxillipedal endite short, not reaching base of palp article 2. Pereopod 1,<br />
propodal palm stepped; carpus distally bluntly triangular. Uropodal exopod<br />
apically acute, outer margin distally sinuate; endopod length about twice<br />
basal width. Pleotelson with posterolateral margins faintly denticulate; apex<br />
subtruncate and bearing several setae.<br />
RECORDS Harrington Sound, Bermuda.<br />
Apanthuroides Menzies and Glynn, 1968<br />
DIAGNOSIS Eyes present. Mandibular palp of three articles; body of mand<br />
ible anteriorly produced, molar absent on left side, present as narrow<br />
spikelike process on right. Maxillipedal palp of three articles; endite present,<br />
reaching palp article 2. Pereopod 1, propodus barely expanded, palm un-
Chalixanthura 27<br />
armed. Pereonite 7 less than half length of pereonite 6. Pereopods 4-7, carpi<br />
with free anterior margin shorter than posterior margin. Pleonites 1—5 fused,<br />
6 fused with telson. Pleopod 1, both rami forming operculum.<br />
Apanthuroides millae Menzies and Glynn, 1968<br />
Figure 8<br />
DIAGNOSIS 9: 4.5 mm. Integument with diffuse brown pigmentation and<br />
numerous shallow pits. Antennular flagellum of four articles. Antennal<br />
flagellum of seven articles. Pleotelson with strong middorsal ridge. 6: 5.0<br />
mm. Integumental pigment stronger than in 9, in irregular brown patches.<br />
Eyes larger than in 9 . Antennular flagellum of six or seven articles. Antennal<br />
flagellum of seven articles. Pereopod 1, propodal palm armed with five<br />
fringed spines.<br />
RECORDS Carrie Bow Cay, intertidal to 30 m; Puerto Rico, intertidal.<br />
REMARKS The highly modified mandible probably indicates some spe<br />
cialized form of feeding, but this has yet to be discovered.<br />
Chalixanthura Kensley, 1984<br />
DIAGNOSIS Eyes present, enormously enlarged, especially ventrally in
Figure 8. Apanthuroides millae: A, $; B, right mandible; C, left<br />
maxilliped; E, uropodal exopod; F, pleopod 1; G, pereopod 1.
Chalixanthura lewisi Kensley and Snelgrove, 1987<br />
Figure 9A-D<br />
Cortezura 29<br />
DIAGNOSIS 9: 3.1 mm. Integument lacking pigment. Antennular flagellum<br />
of three articles. Antennal flagellum of seven articles. Maxillipedal endite<br />
large, apically acute. Pereopod 1 propodus slightly expanded, palm with few<br />
(3) simple setae. Pereopod 7 propodus with two elongate anterodistal fringed<br />
spines. Uropodal and telsonic margins serrate. 6: 2.2 mm. Antennular<br />
flagellum of 11 articles. Antennal flagellum of six articles. Maxilliped re<br />
duced, lacking endite. Pereopod 1, propodal palm with two spines plus seven<br />
spines on mesial surface. Pereopod 7 as in 9.<br />
RECORDS Barbados, in Madracis mirabilis coral, 9—15 m.<br />
Chalixanthura scopulosa Kensley, 1984<br />
Figure 9E-J<br />
DIAGNOSIS 9: 2.5 mm. Antennular flagellum of three articles. Antennal<br />
flagellum of seven articles. Maxillipedal endite short, not reaching base of<br />
palp article 4. Pereopod 1, propodal palm unarmed. Pereopod 7, propodus<br />
with single elongate anterodistal fringed spine. Uropodal exopod margin de<br />
eply incised; endopod ovate, margins serrate. Telson elongate-ovate, pos<br />
terior margin serrate. 6: 2.6 mm. Eyes considerably larger and with more<br />
ommatidia than in 9. Antennular flagellum of seven articles. Antennal<br />
flagellum of four articles. Pereopod 1 propodal palm with three sensory<br />
spines. Uropodal exopod deeply incised.<br />
RECORDS Carrie Bow Cay, Belize, 0.1 m.<br />
Cortezura Schultz, 1977<br />
DIAGNOSIS Eyes small, weakly pigmented. Antennular flagellum of two<br />
short articles. Antennal flagellum of single article. Mandibular palp of two<br />
articles. Maxillipedal palp of two articles, terminal article small; short endite<br />
present. Pereopod 1, propodus inflated. Pereopods 4-7 with carpus having<br />
anterior margin shorter than posterior. Pleonites 1—5 fused; pleonite 6 dor-<br />
sally demarked. Pleopod 1, exopod operculiform. Pleotelson with two basal<br />
statocysts.<br />
*
Figure 9. Chalixanthura lewisi: A, 6; By 9; C, uropodal exopod; D, pleopod 1.<br />
Chalixanthura scopulosa: E, 6] F, 9; G, maxilliped; H, pereopod 1; 7, pleopod \\J,<br />
uropodal exopod.
Cortezura confixa (Kensley, 1978)<br />
Figure 10<br />
Cyathura (Cyathura) 31<br />
DIAGNOSIS 9 13.4 mm. Antennular peduncle articles 1 and 2 each with<br />
clump of ventrally directed setae. Antennal peduncle article 2 and antennu<br />
lar peduncle article 1 locked together. Mandibular palp of two articles, distal<br />
article twice length of proximal. Pereopod 1, propodus expanded, especially<br />
posteriorly; palm concave, with strong tubercle and irregular band of setae<br />
on mesial surface. Uropodal exopod ovate, apically rounded; endopod length<br />
1.5 times greatest width. Telson ovate, posteriorly faintly narrowed, prox-<br />
imally thickened, with faint ridges diverging posteriorly.<br />
RECORDS Gubagua Island, Venezuela, 4-10 m.<br />
REMARKS This species was described under the generic name Venezanthura<br />
Kensley (1978). The only other known species of Cortezura is the type of the<br />
genus, C. penascoensis Schultz, 1977, from California.<br />
Cyathura Norman and Stebbing, 1886<br />
DIAGNOSIS Eyes present or absent. Antennular flagellum of one to three<br />
articles in 9 . Antennal flagellum of one to three articles. Mandibular palp of<br />
three articles. Maxillipedal palp of two articles; endite reduced or absent.<br />
Pereopod 1, propodus inflated. Pereopods 4-7, carpi triangular. Pleopod 1<br />
exopod operculiform. Pleonites 1-5 short, fused; pleonite 6 fused with telson,<br />
sometimes dorsally demarked. Pleotelson with two basal statocysts.<br />
Key to subgenera of Cyathura<br />
1. Pleonite 6 not dorsally demarked from telson; articulation of uropodal<br />
exopod very short; exopod not adpressed to telson dorsally<br />
Stygocyathura<br />
Pleonite 6 dorsally demarked from telson; articulation of uropodal<br />
exopod relatively elongate; exopod adpressed dorsally to telson<br />
Cyathura (Cyathura) Norman and Stebbing, 1886<br />
Cyathura<br />
DIAGNOSIS Integument usually strongly pilose or setose; eye and body pig-
Figure 10. Cortezura confixa: A, 9; B, cephalon and pereonite 1, lateral view<br />
(pereopod 1 removed); C, antennule; D, pleon, lateral view; E, mandible; F,<br />
maxilliped; G, pereopod 1; H, pereopod 7.
Cyathura (Stygocyathura) 33<br />
mentation usually present. Pereopod 1, propodal palm armed with tubercle.<br />
Pleopod 1, protopod with retinaculae. Uropodal exopod articulation relative<br />
elongate along lateral margin of sympod; exopod well developed, ovate.<br />
Pleonite 6 dorsally demarked from telson. Marine or estuarine forms.<br />
Cyathura (Cyathura) cubana Negoescu, 1979<br />
Figure 11A,B<br />
DIAGNOSIS Ovigerous 9: 7.0 mm. Antennular flagellum of two articles. An-<br />
tennal flagellum of one article. Pereopod 1, propodal palm with rounded lobe<br />
in proximal half Maxillipedal palp with distal article 0.34 times length of<br />
proximal article; small rounded endite present. Dorsal pigmentation consist<br />
ing of irregular brown mottling. 6: 5.5 mm. Antennular flagellum of four<br />
t<br />
articles. Antennal flagellum of three articles. Pereopod 1, propodal palm with<br />
rounded lobe in proximal half. Copulatory stylet elongate-cylindrical, api-<br />
cally narrowed and flexed.<br />
RECORDS Cuba, in mangroves, 2.5-7.0 m; Salt Creek, Belize, in man<br />
groves, 1.5 m.<br />
Cyathura (Stygocyathura) Botosaneanu and Stock, 1982<br />
DIAGNOSIS Eye and body pigmentation absent. Body sparsely pilose or set<br />
ose. Tendency toward elongation of some appendages, especially propodus of<br />
pereopods 2—7. Pereopod 1, propodal palm lacking strong tubercle. Pleopod<br />
1, protopod lacking retinaculae. Pleonite 6 fused with telson, not dorsally<br />
demarked. Uropodal exopod with very short articulation on sympod, not<br />
adpressed dorsally to telson. Cave or hypogean forms.<br />
REMARKS The ten species of Stygocyathura from the area covered in this work<br />
are morphologically very similar, with specific differences, although real,<br />
being very subtle. A dichotomous key would be cumbersome and require<br />
considerable dissection of mouthparts. The copulatory stylet of the male pro<br />
vides a valuable specific feature but males are not always available. Instead<br />
of a key, we have provided a list of species with their total lengths and lo<br />
calities (Table 1). Given the very restricted distribution of these cave species,<br />
material from localities not listed here should be treated as potentially un-<br />
described, and the material compared with descriptions, especially those of<br />
Botosaneanu and Stock (1982).
34 ANTHURIDEA • ANTHURIDAE<br />
Figure 11. Cyathura (Cyathura) cubana: A, 9; By pereopod 1,9. Cyathura<br />
(Stygocyathura) cuborientalis: C, pereopod 1, 9; D, telson. Cyathura (Stygocyathura)<br />
curassavica: E, pereopod 1, 9; F9 telson. Cyathura (Stygocyathura) hummetincki: G,<br />
pereopod 1, 9; H, telson. Cyathura (Stygocyathura) motasi: I, telson; J, pereopod 1,<br />
9.<br />
Cyathura (Stygocyathura) cuborientalis Botosaneanu and Stock, 1982<br />
Figure 11C,D<br />
DIAGNOSIS 9 6.8 mm. Pereopod 1, propodal palm straight, bearing about
Cyathura (Stygocyathura) hummelincki 35<br />
TABLE l. CARIBBEAN SPEGIES OF Cyathura (Stygocyathura), THEIR TOTAL LENGTHS<br />
(MM) AND LOCALITIES<br />
C. (S.) cuborientalis<br />
C. (S.) curassavica<br />
C. (S.) hummelincki<br />
C. (S.) motasi<br />
C. (S.) orghidani<br />
C. (S.) parapotamica<br />
C. (S.) salpiscinalis<br />
C. (S.) sbordonii<br />
C. (S.) specus<br />
C. (S.) univam<br />
9 6.8<br />
6 7.0, 9 9.2<br />
6 4.75, 9 8.5<br />
6 6.8, 9 10.0<br />
9 8.0<br />
ovig. 9 3.6, 9 4.1<br />
6 5.6, 9 7.3<br />
6, 9 9.0<br />
6 18.0, 9 19.8<br />
9 10.0<br />
Cuba<br />
Curasao<br />
Aruba<br />
Haiti<br />
Cuba<br />
Jamaica<br />
Haiti<br />
Vera Cruz, Mexico<br />
Cuba<br />
Venezuela<br />
10 pectinate marginal spines; low triangular ridge present. Pleotelson evenly<br />
tapering to notched apex; angle of apex about 90°.<br />
RECORDS Oriente Province, Cuba, interstitial in river alluvia.<br />
Cyathura (Stygocyathura) curassavica Stork, 1940<br />
Figures 11E,F; 12<br />
DIAGNOSIS 6 7.0 mm, 9 9.2 mm. Pereopod 1, propodal palm gently con<br />
vex, bearing about 10 pectinate spines. Pleotelsonic margins in anterior two-<br />
thirds subparallel, tapering gently to finely notched apex; angle of apex less<br />
than 90°.<br />
RECORDS Curagao, from pits and wells.<br />
Cyathura (Stygocyathura) hummelincki Botosaneanu and Stock, 1982<br />
Figure 11G,H<br />
DIAGNOSIS 6 4.75 mm, 9 8.5 mm. Pereopod 1, propodal palm gently sinu<br />
ate, bearing 11-23 marginal pectinate spines. Pleotelsonic margins faintly<br />
concave in midregion, posterior margin evenly convex, apex with slight<br />
notch.<br />
RECORDS Aruba, in pits, wells, and temporary water sources.
36 ANTHURIDEA • ANTHURIDAE<br />
Figure 12. Cyathura<br />
(Stygocyathura) curassavica:<br />
pleon.<br />
Cyathura (Stygocyathura) motasi Botosaneanu and Stock<br />
Figure 111 J<br />
?<br />
1982<br />
DIAGNOSIS 6 . 8 m m , 9 1 0 . 0 m m . P e r e o p o d 1 , p r o p o d a l p a l m g e n t l y s i n u <br />
a t e , b e a r i n g 1 1 - 1 8 m a r g i n a l p e c t i n a t e s e t a e . P l e o t e l s o n i c m a r g i n s t a p e r i n g<br />
e v e n l y t o s l i g h t l y n o t c h e d a p e x ; a n g l e o f a p e x l e s s t h a n 9 0 ° .<br />
RECORDS Haiti, from wells.<br />
Cyathura (Stygocyathura) orghidani Negoescu Vladescu, 1983<br />
Figure 13A,B<br />
DIAGNOSIS 8.0 mm. Pereopod 1, propodal palm slightly convex, bearing<br />
about 11 marginal pectinate setae. Pleotelson, angle of apex obtuse, with<br />
small notch.<br />
RECORDS Pinar del Rio Province, Cuba, from freshwater lake in cave.<br />
C y a t h u r a ( S t y g o c y a t h u r a ) p a r a p o t a m i c a B o t o s a n e a n u a n d S t o c k , 1 9 8 2<br />
Figure 13D,E<br />
DIAGNOSIS m m ( o v i g . 9 3 . 6 m m ) . P e r e o p o d 1 , p r o p o d a l p a l m<br />
a t e , b e a r i n g e i g h t m a r g i n a l p e c t i n a t e s e t a e . P l e o t e l s o n g e n t l y t a p e r i n g t o<br />
n o t c h e d a p e x ; a n g l e o f a p e x a b o u t 9 0 ° .<br />
R E C O R D S J a m a i c a , f r o m r i v e r a l l u v i a .
Figure 13. Cyathura (Stygocyathura) orghidani (from Negoescu, 1983): A, telson; B,<br />
pereopod 1,9. Cyathura (Stygocyathura) sbordonii: C, pereopod 1, 9 (from Argano,<br />
1971). Cyathura (Stygocyathura) parapotamica: D> telson; E, pereopod 1, 9. Cyathura<br />
(Stygocyathura) salpicinalis: F, pereopod 1, 9; G, telson. Cyathura (Stygocyathura)<br />
specus: H, 9; I, telson and uropods; J, pereopod 1, 9. Cyathura (Stygocyathura)<br />
univam: K, telson.
38 ANTHURIDEA • ANTHURIDAE<br />
Cyathura (Stygocyathura) salpiscinalis Botosaneanu and Stock, 1982<br />
Figure 13F,G<br />
DIAGNOSIS 6 5.6 mm, 9 7.3 mm. Pereopod 1, propodal palm sinuate,<br />
bearing up to 15 marginal pectinate setae, and with distinct triangular ridge.<br />
Pleotelson gently tapering, with slight apical eminence.<br />
RECORDS Haiti, from alluvia of lake.<br />
Cyathura (Stygocyathura) sbordonii Argano, 1971<br />
Figure 13C<br />
DIAGNOSIS 6 and 9 9.0 mm. Pereopod 1, propodal palm convex, bearing<br />
up to 16 marginal pectinate setae. Pleotelson with angle of apex obtuse.<br />
RECORDS Vera Cruz, Mexico, from freshwater in cave.<br />
Cyathura (Stygocyathura) specus Bowman, 1965<br />
Figure 13H-J<br />
DIAGNOSIS 6 18.0 mm, 9 19.8 mm. Pereopod 1, propodal palm almost<br />
straight, bearing up to 15 marginal pectinate setae. Pleotelson with angle of<br />
apex obtuse.<br />
RECORDS Las Villas Province, Cuba, from freshwater lake in cave.<br />
Cyathura (Stygocyathura) univam Botosaneanu, 1983<br />
Figure 13K<br />
DIAGNOSIS 9 10.0 mm. Pereopod 1, propodal palm gently convex, bearing<br />
22 marginal pectinate setae. Pleotelson posteriorly broadly rounded, apex<br />
emarginate.<br />
RECORDS Peninsula de Morocoy, Venezuela, from phreatic water in cave.<br />
Eisothistos Haswell, 1884<br />
DIAGNOSIS Mouthparts forwardly produced, tailfan spiny, indurate. Eyes<br />
present, larger and with more ommatodia in 6 than in 9. Pereonites some<br />
times elongate. Mandible with strong incisor, reduced lamina dentata, palp<br />
and molar lacking. Maxilliped lacking endite, palp slender, of three to five
Eisothistos teri 39<br />
articles. Pereopods 1-3 not subchelate, propodi relatively elongate, mini<br />
mally expanded. Pleopod 1 9, rami fused, together forming operculum; 6<br />
rami separate. Pleonites free, short, longer in 6 than in 9. Telson lacking<br />
statocysts.<br />
REMARKS Wagele (1979) first recorded species of Eisothistos preying on ser-<br />
pulid polychaete worms in their tubes.<br />
The genus contains about 12 species in the Pacific, Indian Ocean, Carib<br />
bean, Antarctic, and Mediterranean.<br />
Key to species of Eisothistos<br />
1. 9 telson with middorsal spines; 6 pereopod 1 propodal palm with 19<br />
or 20 spines teri<br />
9 telson lacking middorsal spines; 6 pereopod 1 propodal palm with<br />
11 spines petrensis<br />
Eisothistos petrensis Kensley, 1984<br />
Figure 14A-E<br />
DIAGNOSIS 9: 4.0 mm. Antennular flagellum of six articles. Antennal<br />
flagellum of six articles. Telson posteriorly faintly bilobed, margin strongly<br />
serrate; faint anterior middorsal ridge; middorsally unarmed. Pereopod 1,<br />
propodal palm unarmed. Pleopod 1 rami fused for V12 of length. 6: 2.0 mm.<br />
Antennular flagellum of eight articles. Antennal flagellum of six articles. Per<br />
eopod 1 propodal palm with 11 fringed spines. Telson narrower than in 9,<br />
with middorsal ridge running almost entire length.<br />
RECORDS Carrie Bow Cay, Belize, 0.1-36 m; Looe Key, Florida, 5-6 m;<br />
Turks and Caicos Islands, 1.0 m; St.Thomas, U.S. Virgin Islands, 7-10 m.<br />
Eisothistos teri Kensley and Snelgrove, 1987<br />
Figure 14F-H<br />
DIAGNOSIS 9: 3.2 mm. Basal antennular peduncle article bearing triangu<br />
lar apically rounded laminate process; flagellum of seven articles. Antennal<br />
flagellum of six articles. Mandible with biserrate lamina dentata. Pereopod 1<br />
propodal palm unarmed. Pleopod 1 rami fused for 3 A of length. Uropodal<br />
exopod with one or two strong slightly recurved spines on dorsal surface.
40 ANTHURIDEA • ANTHURIDAE<br />
Figure 14. Eisothistos petrensis: Ay 9; By uropodal sympod and endopod; Cy<br />
uropodal exopod; D, pleopod 1; Ey pereopod 1, 6. Eisothistos ten: Fy telson and<br />
uropod; Gy pleopod 1; Hy pereopod 1, 6.<br />
Telson with eight or nine slightly recurved middorsal teeth becoming longer<br />
posteriorly. 6: 2.0 mm. Antennular flagellum of eight articles. Antennal
Haliophasma curri 41<br />
flagellum of six articles. Pereopod 1 propodal palm with 19 or 20 fringed<br />
spines. Uropodal exopod lacking dorsal teeth. Telson lacking middorsal<br />
teeth; posterior margin incised into 12 acute or narrowly rounded teeth.<br />
RECORDS Barbados, in Madracis mirabilis coral, 9—15 m.<br />
Haliophasma Haswell, 1881<br />
DIAGNOSIS Eyes present. Integument often indurate, with scattered pitting.<br />
Antennular flagellum usually of two articles. Antennal flagellum of 4—7 arti<br />
cles. Mandibular palp of three articles. Maxillipedal palp of two articles,<br />
article 2 smaller than article 1. Pereopod 1, propodus expanded. Pereopods<br />
4-7, carpi roughly rectangular. Pleopod 1 exopod operculiform. Pleonites 1 —<br />
5 short, fused; pleonite 6 usually demarked from telson. Latter with two<br />
basal statocysts. 6 often with more elongate form than 9. Antennular<br />
flagellum multiarticulate. Eyes larger.<br />
Key to species of Haliophasma<br />
1. Telson posteriorly narrowly rounded; dactylus of pereopod 1 dentate<br />
Telson posteriorly broadly rounded; dactylus of pereopod 1 entire<br />
Haliophasma curri Paul and Menzies, 1971<br />
Figure 15A-C<br />
valeriae<br />
DIAGNOSIS 9 7.0 mm. Antennular flagellum of three articles. Antennal<br />
flagellum of five articles. Mandibular palp article 3 with five spines. Max<br />
curri<br />
illipedal palp, terminal article small, set obliquely at distolateral angle of<br />
article 1. Pereopod 1, carpus triangular, posterodistal margin crenulate, dis-<br />
tally rounded; propodal palm crenulate, with low rounded proximal lobe.<br />
Pleonite 6 dorsally demarked, narrow, with middorsal point in posterior<br />
margin. Uropodal exopod elongate, outer margin sinuate, dentate; endopod<br />
ovate, distally narrowed, length twice greatest width, outer margin dentate.<br />
Telson parallel sided, posterior margin broadly rounded.<br />
RECORDS Off Venezuela, 95 m; Culebra Island, Bay of Panama, intertidal.
42 ANTHURIDEA • ANTHURIDAE<br />
Figure 15. Haliophasma curri: A, 9; B, pereopod 1; C, uropod. Haliophasma valeriae:<br />
D, 9; E, pereopod 1; F, uropod.<br />
Haliophasma valeriae Paul and Menzies, 1971<br />
Figure 15D-F<br />
DIAGNOSIS 9 6.5 mm. Body long and slender. Antennular flagellum of
Malacanthura 43<br />
three articles. Antennal flagellum of six articles. Mandibular palp article 3<br />
with four spines. Pereopod 1, carpus triangular, tipped with acute tooth;<br />
propodus elongate, palm with about seven teeth, five fringed spines on mesial<br />
surface; unguis of dactylus strongly flexed; margin of dactylus with three<br />
strong triangular teeth. Uropodal exopod elongate, apically acute, margins<br />
serrate; endopod length little more than twice greatest width, outer margin<br />
serrate, apex acute. Telson elongate-elliptical, apically narrowly rounded;<br />
strong middorsal longitudinal rounded ridge running almost entire length.<br />
RECORDS Off Venezuela, 95 m.<br />
REMARKS Wagele (1981) made this species the type of his new genus Ne-<br />
manthura, based primarily on the elongate form of the body and appendages.<br />
Haliophasma irmae Paul and Menzies, 1971, from the same locality as the<br />
above species, is probably the same species.<br />
Licranthura Kensley and Schotte, 1987<br />
DIAGNOSIS Serrate process on antennal peduncle article 3. Mandibular<br />
palp of three articles; molar lacking. Maxillipedal endite short. Pereopod 1<br />
larger than pereopods 2 and 3. Pereopods 4—7, carpi triangular. Pleonites<br />
short, free. Pleopod 1, both rami forming operculum. Pleotelson lacking<br />
statocysts.<br />
Licranthura amyle Kensley and Schotte, 1987<br />
Figure 16<br />
DIAGNOSIS 9 3.8 mm. Eyes small, pigmented. Antennular flagellum of<br />
three articles. Antennal peduncle article 3 with lamellar expanded process,<br />
serrate on mesial margin; flagellum of six articles. Maxillipedal palp of five<br />
articles; very short endite. Pereopod 1, propodal palm unarmed. Uropodal<br />
and telsonic margins serrate.<br />
RECORDS Carrie Bow Cay, Belize, 0-25 m, in coral rubble.<br />
Malacanthura Barnard, 1925<br />
DIAGNOSIS Eyes present. Mandibular palp of three articles. Maxillipedal<br />
palp of three articles, terminal article usually broadly ovate. Pereopod 1,<br />
propodus expanded. Pereopods 4-7 with carpi roughly rectangular. Pleopod
44 ANTHURIDEA • ANTHURIDAE<br />
Figure 16. Licranthura amyle: A, 9; B, antennule and antenna; C, telson and<br />
uropod.<br />
1, exopod operculiform. Pleonites 1—5 short, fused; pleonite 6 dorsally de-<br />
marked. Pleotelson with two basal statocysts.
Malacanthura caribbica Paul and Menzies, 1971<br />
Figure 17<br />
Mesanthura 45<br />
DIAGNOSIS 9 27.1 mm. Integument moderately indurate. Antennular<br />
flagellum of seven articles. Antennal flagellum of four articles. Mandibular<br />
palp, article 3 with comb of 11 spines. Maxillipedal palp, terminal article<br />
broadly ovate, penultimate article with row of seven spines on mesial margin.<br />
Pereopod 1 propodus expanded, palm straight, with few spines on mesial<br />
margin. Uropodal exopod barely reaching base of endopod, narrow, apically<br />
acute, outer margin sinuate, serrate; endopod set obliquely on sympod, mar<br />
gin serrate, apically acute. Telson lanceolate, apically narrowly rounded,<br />
with strong longitudinal middorsal carina.<br />
RECORDS Off Venezuela, 95 m; off Colombia, 42-44 m.<br />
REMARKS Malacanthura cumanensis Paul and Menzies, 1971, described from<br />
the same locality as M. caribbica, was shown to be the latter species (Kensley,<br />
1980).<br />
Mesanthura Barnard, 1914<br />
DIAGNOSIS 9: Dorsal integument with (usually) species-specific pigment<br />
pattern; pigment persistent in alcohol. Mandibular palp of three articles,<br />
terminal article with row of spines, number of which specific for species.<br />
Maxilliped with endite either very reduced or absent; palp of three articles,<br />
with terminal article usually about half length of penultimate article, suture<br />
transverse. Pereopod 1, propodus expanded, palm often with step. Pereopods<br />
2 and 3, propodi not expanded. Pereopods 4—7, carpi roughly triangular,<br />
with anterior margin shorter than posterior margin. Pleonites 1—5 fused,<br />
pleonite 6 dorsally demarked. Pleopod 1, exopod operculiform. Telson with<br />
two basal statocysts. 6: Eyes larger than in 9. Antennular flagellum of sel<br />
dom more than 10 articles bearing numerous aesthetascs. Mouthparts, espe<br />
cially body of mandible, reduced. Pereopod 1, propodus bearing dense band<br />
of spines on mesial surface near palm. Pigment pattern more diffuse than in<br />
9, extending onto ventral surface.<br />
REMARKS Mesanthura is a relatively large genus of about 30 species, re<br />
corded from most tropical and temperate seas, in shallow habitats. The<br />
males of few species have been recorded; by themselves, males are difficult to<br />
identify as the dorsal pigment pattern characteristic of the female breaks<br />
down and spreads onto the ventrum.
Figure 17. Malacanthura caribbica: A, 9;<br />
maxilliped; E, uropod; F, pleopod 1.
Key to species of Mesanthura (9 only)<br />
Mesanthura bivittata 47<br />
1. Pigment in tiny evenly scattered chromatophores over body; cephalon<br />
with solid patch of pigment; mandibular palp article 3 with seven<br />
spines punctillata<br />
Pigment not evenly scattered over body 2<br />
2. Pereonites 4 and 5 with patch of pigment, sometimes with unpigmented<br />
area in middle of patch 3<br />
Pereonites 4 and 5 lacking fairly solid patch of pigment 6<br />
3. Pereonites 4 and 5 lacking unpigmented area in pigment patch; five<br />
transverse lines on pigment on pleon; mandibular palp article 3 with<br />
six spines paucidens<br />
Pereonites 4 and 5 with unpigmented area in pigment patch; pleon<br />
lacking transverse pigment lines 4<br />
4. Pigment in obvious double longitudinal bands on pereon and pleon;<br />
mandibular palp article 3 with nine spines bivittata<br />
Pigment not in obvious double bands 5<br />
5. Unpigmented area in middle of pigment patch of pereonites 1-3;<br />
mandibular palp article 3 with 10 spines pulchra<br />
No unpigmented area in middle of pigment of pereonites 1-3;<br />
mandibular palp article 3 with eight spines looensis<br />
6. Pigment of pereon in fine reticulate lines; mandibular palp article 3<br />
with six spines reticulata<br />
Pigment of pereon not in fine reticulate lines; mandibular palp article 3<br />
with four spines 7<br />
7. Pigment in more or less complete rings on pereonites 1—6 hopkinsi<br />
Pigment in strong transverse posterior bars on pereonites 4-7 . .fasciata<br />
Mesanthura bivittata Kensley, 1987a<br />
Figures 18A, 20A-D<br />
DIAGNOSIS 6 5.2 mm, ovigerous $ 7.8 mm. Pigment in obvious double<br />
longitudinal bands on pereon and pleon. Mandibular palp article 3 with nine<br />
spines. Maxilliped lacking endite. Pereopod 1, propodal palm with rounded<br />
lobe.<br />
RECORDS Twin Cays, Belize, under red mangroves, 1-2 m.
Mesanthura<br />
W®<br />
A-O<br />
I,<br />
• •<br />
V > : < <<br />
^<br />
A<br />
rr-<br />
/v\'<br />
a *<br />
Mesanthura bivittata; B, Mesanthura fasciata<br />
^<br />
«-tf<br />
4<br />
-1'<br />
V<br />
C
Mesanthura fasciata 49<br />
Figure 19. A, Mesanthura paucidens; B, Mesanthura pulchra; C, Mesanthura punctillata;<br />
D, Mesanthura reticulata.<br />
Mesanthura fasciata Kensley, 1982<br />
Figures 18B, 20E-H<br />
DIAGNOSIS 9 4.5 mm. Pigment in triangular patch on head, open irregular<br />
rings on pereonites 1-3; pereonites 4—7 with strong transverse posterior bar;
Figure 20. Mesanthura bivittata: A, mandible; By maxilliped; C, pereopod 1; D,<br />
uropodal exopod. Mesanthura fasciata: E, mandible; F, maxilliped; G, pereopod 1;<br />
H, uropodal exopod. Mesanthura hopkinsi: I, mandible; J, maxilliped; K, pereopod 1;<br />
L, uropodal exopod.
Mesanthura paucidens 51<br />
five bars on pleon. Mandibular palp article 3 with four spines. Maxilliped<br />
with very reduced endite. Pereopod 1, propodal palm with step.<br />
RECORDS Looe Key, Florida, 5-6 m; Cozumel, Mexico; Jamaica; Carrie<br />
Bow Cay, Belize, 0.2-6 m.<br />
Mesanthura hopkinsi Hooker, 1985<br />
Figures 18C, 20I-L<br />
DIAGNOSIS 9 2.4 mm. Pigment in triangle on cephalon; in irregular rings<br />
on pereonites 1-6; four transverse bars on pleon. Mandibular palp article 3<br />
with four spines. Maxilliped lacking endite. Pereopod 1, palm of propodus<br />
lacking step.<br />
RECORDS Looe Key, Florida, 0.5 m; Florida Middlegrounds, Gulf of Mex<br />
ico, 55 m.<br />
Mesanthura looensis Kensley and Schotte, 1987<br />
Figures 18D, 21A-D<br />
DIAGNOSIS 9 10.0 mm. Pigment in solid patches on cephalon, pereonites,<br />
pleon, uropods and telson; pereonites 4—6 with open central area in patch.<br />
Mandibular palp article 3 with eight spines. Maxilliped lacking endite. Per<br />
eopod 1, propodal palm with step.<br />
RECORDS Looe Key, Florida, 1 m.<br />
Mesanthura paucidens Menzies and Glynn, 1968<br />
Figures 19A, 2 IE-1<br />
DIAGNOSIS 9: 6.6 mm. Pigment in roughly rectangular to ovate patches on<br />
cephalon and pereonites; in five transverse lines connected laterally on pleon.<br />
Mandibular palp article 3 with six spines. Maxilliped with short narrow en<br />
dite. Pereopod 1 propodal palm with step. 6: 6.4 mm. Antennular flagellum<br />
of seven articles. Pigment more diffuse than in 9 but retaining five pleonal<br />
bars.<br />
RECORDS Looe Key, Florida, 5-6 m; Carrie Bow Cay, Belize, 15.2 m; Pu<br />
erto Rico, intertidal; Jamaica.
52 ANTHURIDEA • ANTHURIDAE<br />
Figure 21. Mesanthura looensis: A, mandible; B, maxilliped; C, pereopod 1; D,<br />
uropodal exopod. Mesanthura paucidens: Ey mandible; F9 maxilliped; G, pereopod 1,<br />
9; H, pereopod 1, 8; I> uropodal exopod. Mesanthura pulchra: J, mandible; K,<br />
maxilliped; L, pereopod 1, 6; M, pereopod 1, 9; JV, uropodal exopod.<br />
Mesanthura pulchra Barnard, 1925<br />
Figures 19B, 21J-N<br />
DIAGNOSIS 9: 9.3 mm. Pigment pattern in roughly rectangular patches on<br />
cephalon and pereonites 1-6, with open oval middorsal area in patch. Man<br />
dibular palp article 3 with 10 spines. Maxilliped lacking endite. Pereopod 1,<br />
propodal palm with step. 6: 5.4 mm. Antennular flagellum of eight articles.
Minyanthura corallicola 53<br />
RECORDS Egmont Key, Florida, 18.3-36.6 m; Looe Key, Florida, 0.5-12<br />
m; Dry Tortugas; Turks and Caicos Islands, 1 m; Puerto Rico, intcrtidal to<br />
1.5 m; Carrie Bow Cay, Belize, intertidal to 1.5 m; St.Thomas and St.John's,<br />
U.S. Virgin Islands; Cozumel, Mexico.<br />
REMARKS Menzies and Glynn (1968) recorded this species as M. decorata<br />
from Puerto Rico, while Menzies and Kruczynski (1983) recorded it as M.<br />
Jloridensis.<br />
Mesanthura punctillata Kensley, 1982<br />
Figures 19C, 22A-F<br />
DIAGNOSIS 9: 6.4 mm. Pigment in solid patch between eyes, rest of pig<br />
ment on pereon and pleon with chromatophores scattered, diffuse, not in any<br />
regular pattern. Mandibular palp article 3 with seven spines. Maxilliped<br />
lacking endite; terminal palp article semicircular. Pereopod 1 propodal palm<br />
with step. 6: 4.5 mm. Antennular flagellum of 10 articles.<br />
RECORDS Turks and Caicos Islands, 1 m; Carrie Bow Cay, 0.2-20 m.<br />
Mesanthura reticulata Kensley, 1982<br />
Figures 19D, 22G-J<br />
DIAGNOSIS 9 6.1 mm. Dorsal pigment pattern a network of<br />
chromatophores arranged in fine lines. Mandibular palp article 3 with six<br />
spines. Maxilliped lacking endite. Pereopod 1 propodal palm with step.<br />
RECORDS Carrie Bow Cay, 10-24 m.<br />
Minyanthura Kensley, 1982<br />
DIAGNOSIS Mandible lacking molar and palp. Maxillipedal palp of five ar<br />
ticles; large apically acute endite present. Pereopods 4-7, carpi rectangular.<br />
Pleopod 1, both rami forming operculum. Pleonites 1-5 fused; pleonite 6<br />
fused with telson, not dorsally demarked. Telson with two basal statocysts.<br />
Minyanthura corallicola Kensley, 1982<br />
Figure 23<br />
DIAGNOSIS 9: 1.7 mm. Antennular flagellum of one article. Antennal
Figure 22. Mesanthura punctillata: A, mandible, 9; B, mandible 8; C, maxilliped; D,<br />
pereopod I, 9; E, pereopod 1, 6\ F9 uropodal exopod. Mesanthura reticulata: G,<br />
mandible; H, maxilliped; /, pereopod \\ J, uropodal exopod.
Minyanthura corallicola 55<br />
Figure 23. Minyanthura corallicola: A, 9; B, telson and uropod; C, pleopod 1; D,<br />
mandible; E, maxilliped; F, pereopod 1.<br />
flagellum of four articles. Pereopod 1, propodus somewhat inflated. Uropo-<br />
dal rami and posterior telsonic margin serrate. 6: 1.3 mm. Eyes larger than<br />
in 9. Antennular flagellum of three articles. Antennai flagellum of two<br />
articles.<br />
RECORDS Carrie Bow Cay, Belize, 6-24 m; Barbados, 9-15 m; Jamaica.
56 ANTHURIDEA • ANTHURIDAE<br />
Pendanthura Menzies and Glynn, 1968<br />
DIAGNOSIS 9 : Integument with some red-brown dorsal pigmentation. Eyes<br />
small, pigmented. Antennular flagellum of two articles. Antennal flagellum<br />
of one article. Mandibular palp of single reduced article; incisor, molar, and<br />
lamina dentata present. Maxillipcdal palp of single broad article; small tri<br />
angular endite present. Pereopod 1, propodus expanded. Pereopods 4-7,<br />
carpi short, triangular, lacking free anterior margin. Pleonites 1-5 very<br />
short, fused, 6 fused with telson. Pleopod 1, exopod operculiform. Telson<br />
basally broad, with two statocysts at about midlength. 6: Antennular ped<br />
uncle of four articles. Antennal flagellum of one article. Pereopod 1, pro<br />
podus with dense clump of spines on mesial surface. Eyes only slightly larger<br />
than in 9.<br />
REMARKS The genus comprises three species, two from the Caribbean and<br />
one from the Pacific, all of which have been taken from shallow coral reefs.<br />
Key to species of Pendanthura<br />
1. Dorsal pigmentation over entire body; pereopod 1, propodal palm with<br />
rounded lobe tanaiformis<br />
Dorsal pigmentation on cephalon, pereonite 2, and pleon; pereopod 1,<br />
propodal palm lacking rounded lobe hendleri<br />
Pendanthura hendleri Kensley, 1984<br />
Figure 24A-E<br />
DIAGNOSIS 9: 3.3 mm. Dorsal pigmentation limited to cephalon, pereonite<br />
2, and very short pleon. Reduced mandibular palp with two setae. Pereopod<br />
1, propodus expanded, palm gently convex, lacking rounded lobe, with four<br />
spines on mesial surface near palmar margin. 6: 2.8 mm. Pigmentation as in<br />
9. Pereopod 1, propodal palm gently convex, lacking rounded lobe.<br />
RECORDS Carrie Bow Cay, Belize, 9-23 m; Twin Cays, Belize, 0-2 m; Pan<br />
ama, 30 m.<br />
Pendanthura tanaiformis Menzies and Glynn, 1968<br />
Figure 24F-H<br />
DIAGNOSIS Ovigerous 9: 2.9 mm. Dorsal pigmentation a dense red-brown
Figure 24. Pendanthura hendleri: A, 2; B, mandible; C, maxilliped; D, pereopod 1,<br />
6\ Ey pereopod 1, 2. Pendanthura tanaiformis: Fy pereopod 1, 2; G, 2; H> mandible.
58 ANTHURIDEA • HYSSURIDAE<br />
reticulation over entire dorsum. Mandibular palp with one seta. Pereopod 1,<br />
propodus expanded, palm with rounded lobe, mesial surface with six spines<br />
near palmar margin. 6: 2.8 mm. Pigmentation as in 9. Propodal palm with<br />
rounded lobe.<br />
RECORDS Bermuda; Carrie Bow Cay, Belize, intertidal to 1 m; Puerto Rico,<br />
intertidal; Cozumel, Mexico.<br />
REMARKS Kensley (1984) characterized this common species of the interti<br />
dal of the reef crest as stress-tolerant, breeding throughout the year.<br />
Skuphonura Barnard, 1925<br />
DIAGNOSIS Antennal flagellum of single article. Cephalon with midventral<br />
toothlike process at base of mouthparts. Mandibular palp of three articles.<br />
Maxillipedal palp of three articles; endite lacking. Pereopod 1, propodus ex<br />
panded. Pereopods 4—7, carpi triangular. Pleopod 1, exopod operculiform.<br />
Pleonites 1—5 fused; pleonite 6 dorsally demarked. Pleotelson with two basal<br />
statocysts.<br />
Skuphonura laticeps Barnard, 1925<br />
Figure 25<br />
DIAGNOSIS 6 6.0 mm. Cephalon wider anteriorly than posteriorly, with<br />
anterolateral lobes extending well beyond rostrum. Antennular flagellum of<br />
two articles. Pereonite 1 with strong midventral forwardly directed tooth.<br />
Pereopod 1, carpus with posterodistal angle produced into triangular lobe;<br />
propodus expanded, palmar margin proximally convex, numerous setae on<br />
mesial surface. Pleopod 2, copulatory stylet of endopod reaching beyond<br />
rami. Pleonite 6 dorsally demarked, posterior margin middorsally incised.<br />
Uropodal exopod ovate, with distal notch; endopod length slightly more than<br />
basal width. Pleotelson widest at midlength, posteriorly narrowly rounded,<br />
with broadly rounded longitudinal raised area.<br />
RECORDS St. Thomas, U.S. Virgin Islands, 8-40 m.<br />
Family Hyssuridae Wagele, 1981<br />
DIAGNOSIS Maxillipedal palp usually of five articles; endite present. Per<br />
eopods 1-3 subsimilar, often all three pairs subchelate. Pleopod 1 similar to
Figure 25. Skuphonura laticeps: A, 6; B, cephalon and pereonite<br />
antennule, 8\ D, pereopod 1, 6; E, antenna; F, maxilliped.
60 ANTHURIDEA • HYSSURIDAE<br />
following pleopods, not operculiform. Pleonites 1-5 freely articulating, rela<br />
tively elongate. Pleotclson lacking statocysts.<br />
Key to genera of Hyssuridae<br />
1. Pleotelson not covered by uropodal exopods; maxillipedal rami basally<br />
free Kupellonura<br />
Pleotelson covered completely by uropodal exopods; maxillipedal rami<br />
basally fused Xenanthura<br />
Kupellonura Barnard, 1925<br />
DIAGNOSIS Mandibular palp of three articles. Maxillipedal palp of five arti<br />
cles; large endite present. Pereopods 1-3 similar, propodi somewhat ex<br />
panded. Pereopods 4-7, carpi triangular. Pleonites 1—5 elongate, free, sube-<br />
qual. Pleopods 1—5 similar.<br />
Kupellonura imswe (Kensley, 1982)<br />
Figures 26, 27<br />
DIAGNOSIS 9: 3.4 mm. Eyes present. Antennular flagellum of four articles.<br />
Antennal flagellum of seven articles. Maxillipedal endite reaching to base of<br />
article 3. Pereopods 1-3 similar; pereopod 1, carpus posterodistally acute;<br />
propodus expanded, palm straight, unarmed; carpus triangular, free anterior<br />
margin shorter than posterior. Pleonite 6 with middorsal incision in posterior<br />
margin. Uropodal exopod, outer margin serrate, distally narrowly rounded;<br />
endopod length twice greatest width. Telson widest at midlength,<br />
posterolateral margin serrate, apically broadly rounded. 6: 2.7 mm. Eyes<br />
larger than in 9. Mouthparts reduced. Pereopod 1, propodal palm armed<br />
with row of eight fringed spines.<br />
RECORDS Carrie Bow Cay, Twin Cays, Glover's Reef, Belize, 0-6.0 m;<br />
Montego Bay, Jamaica, 1 m.<br />
Xenanthura Barnard, 1925<br />
DIAGNOSIS Mandibular palp of single article. Maxillipedal rami fused; ar<br />
ticulations of palp articles obscure. Pereopods 1—3 similar. Pereopods 4-7,<br />
carpi triangular.
Figure 26. Kupellonura imswe: A, 9; B, cephalon and pereonite 1, 6; C, maxilliped;<br />
Dy pereopod 1, 9; E, pereopod 1, 6.
62 ANTHURIDEA • HYSSURIDAE<br />
Figure 27. Kupellonura imswe: A phalon, dorsal view; B, posterior pleon, d<br />
view; C, pleon, ventral view; D> cephalon and pereonites 1 and 2, lateral view<br />
Xenanthura brevitelson Barnard, 1925<br />
Figure 28<br />
D I A G N O S I S 9 : 4 . 0 m m . O m m a t i d i a o f e y e s i n l o n g i t u d i n a l r o w o f t h r e e o r<br />
f o u r g r o u p s . A n t e n n u l a r flagellum o f t h r e e a r t i c l e s . A n t e n n a l f l a g e l l u m o f<br />
t h r e e a r t i c l e s . M a n d i b u l a r p a l p o f s i n g l e s h o r t a r t i c l e b e a r i n g s i n g l e s e t a .<br />
M a x i l l i p e d a l r a m i b a s a l l y f u s e d , p a l p o f t h r e e o r f o u r o b s c u r e l y s e p a r a t e d<br />
a r t i c l e s . P e r e o p o d s 1 - 3 s i m i l a r , c a r p u s o f p e r e o p o d 2 t r i a n g u l a r , w i t h s t r o n g<br />
t r i a n g u l a r p r o j e c t i n g l o b e p o s t e r o d i s t a l l y ; p r o p o d i e x p a n d e d , p a l m i n c i s e d<br />
i n t o s e v e r a l r o u n d e d l o b e s . U r o p o d a l e x o p o d s c i r c u l a r , o v e r l a p p i n g d o r s a l l y<br />
a n d c o v e r i n g t e l s o n ; e n d o p o d p r o j e c t i n g b e y o n d e x o p o d s v e n t r a l l y , m e s i a l<br />
m a r g i n w i t h s t e p , d i s t a l l y r o u n d e d . T e l s o n s h o r t e r t h a n u r o p o d a l e x o p o d ,<br />
p o s t e r i o r m a r g i n t r u n c a t e t o f a i n t l y c o n c a v e . 6 : 3 . 5 m m . A n t e n n u l a r<br />
flagellum o f s e v e n a r t i c l e s .<br />
R E C O R D S O f f G e o r g i a , 2 0 - 1 4 5 m ; o f f F l o r i d a , 8 - 1 0 m ; T u r k s a n d C a i c o s<br />
I s l a n d m ; S t . T h o m a s , U . S . V i r g i n I s l a n d s , 5 0 - 6 0 m ; G u l f o f M
Figure 28. Xenanthura brevitelson: A, 9;B, pereopod 1; C, uropodal endopod<br />
telson; E, uropodal exopod.
64 ANTHURIDEA • PARANTHURIDAE<br />
Family Paranthuridae Menzies and Glynn, 1968<br />
DIAGNOSIS Mouthparts together forming somewhat elongate cone adapted<br />
for piercing and sucking. Mandible with styliform incisor, lacking lamina<br />
dentata and molar. Maxilla slender, styliform, distally serrate. Maxilliped<br />
elongate; number of palp articles usually reduced. Pereopod 1, or pereopods<br />
1-3 subchelate. Pleopod 1 exopod operculiform. Pleonites short, free or<br />
Key to genera of Paranthuridae<br />
1. Pereopod 7 lacking in adult 2<br />
Pereopod 7 present in adult 3<br />
2. Eyes present Colanthura<br />
Eyes absent Curassanthura<br />
3. Antennular and antennal flagellum of more than 10<br />
articles Accalathura<br />
Antennular and antennal flagella with fewer than 10 articles 4<br />
4. Antennal flagellum a single (rarely two or 3) flattened article;<br />
maxillipedal palp of one or two articles Paranthura<br />
Antennal flagellum of seven articles; maxillipedal palp of three articles<br />
Accalathura Barnard, 1925<br />
Virganthura<br />
DIAGNOSIS Eyes present. Antennular and antennal flagella multiarticulate,<br />
each of more than 10 articles. Mandibular palp of three articles. Maxillipedal<br />
Key to species of Accalathura<br />
1. Uropodal exopod elongate-narrow; endopod length twice basal width<br />
crenulata<br />
Uropodal exopod ovate, apically subacute; endopod length 1.5 times<br />
basal width setosa
Colanthura tenuis 65<br />
palp of two articles, endite almost reaching end of palp. Pereopod 1 subche-<br />
late, propodus inflated, larger than pereopods 2 and 3. Pereopods 4-7, carpi<br />
with anterior and posterior margin subequal. Pleonites free, short. Pleopod 1,<br />
exopod operculiform. Telson with single statocyst.<br />
Accalathura crenulata (Richardson, 1901)<br />
Figure 29A-D<br />
DIAGNOSIS 9: 16.0 mm. Antennular flagellum of about 26 articles. Anten<br />
nal flagellum of about 18 articles. Uropodal exopod narrow, parallel sided.<br />
Telson apically subacute. 6: 15.0 mm. Pleopod 2, copulatory stylet of endo-<br />
pod apically acute, with subapical "heel."<br />
RECORDS OfFNorth Carolina, 30 m; off Georgia, 20 m; Cuba; Puerto Rico,<br />
intertidal; Cozumel, Mexico; Carrie Bow Cay, Twin Cays, Belize, intertidal<br />
to 6 m; west coast of Florida, Gulf of Mexico, 55 m.<br />
Accalathura setosa Kensley, 1984<br />
Figure 29E-H<br />
DIAGNOSIS 9: 8.5 mm. Antennular flagellum of 11 articles. Antennal<br />
flagellum of 13 articles. Uropodal rami, margins closely setose; exopod ovate,<br />
outer margin sinuate, apex subacute; endopod ovate, length 1.25 times great<br />
est width. Telson apically rounded. 6: 7.0 mm. Pleopod 2, copulatory stylet<br />
of endopod apically strongly bifid.<br />
RECORDS Carrie Bow Cay, Belize, intertidal to 0.5 m.<br />
Colanthura Richardson, 1902<br />
DIAGNOSIS Integument with minute squamae. Mandible lacking palp.<br />
Maxillipedal palp articles fused except for minute terminal article. Pereopod<br />
1 subchelate, propodus expanded. Pereopods 2 and 3 subchelate but smaller<br />
than pereopod 1. Pereopods 4-6, carpi rectangular. Pereonite 7 very short,<br />
pereopod 7 lacking. Pleotelson lacking statocyst.<br />
Colanthura tenuis Richardson, 1902<br />
Figure 30A-C<br />
DIAGNOSIS 9: 3.5 mm. Eyes present. Integument diffusely brown in color.<br />
Antennular flagellum of four articles. Antennal flagellum of single article.
Figure 29. Accalathura crenulata: A, Q,B, uropodal sympod and endopod; C,<br />
uropodal exopod; D, telson. Accalathura setosa: E, 9; F, uropodal sympod and<br />
endopod; G, uropodal exopod; H, telson.
Curassanthura bermudensis 67<br />
Pereopod 1, propodus with mesial surface bearing proximal row of six spines.<br />
Pleonites 1—5 short, fused, boundaries marked dorsally by folds. Telson pos<br />
teriorly broadly rounded. 6: 3.5 mm. Antennular flagellum of five articles.<br />
RECORDS Bermuda, intertidal to 0.5 m.<br />
Curassanthura Kensley, 1981<br />
DIAGNOSIS Eyes lacking. Mandibular palp of three articles. Maxillipedal<br />
palp of five articles; short endite present. Pereopod 1 subchelate. Pereopods<br />
2-6 similar, carpi rectangular. Pereopod 7 lacking. Pleonites 1-5 free;<br />
pleonite 6 dorsally demarked. Pleopod 1, exopod operculiform. Pleotelson<br />
with single statocyst. Interstitial littoral forms.<br />
REMARKS Three species of this interstitial genus are known, two from the<br />
Caribbean, and one from the upper sublittoral gravels of a lava tunnel on<br />
Lanzarote, Canary Islands.<br />
Key to species of Curassanthura<br />
1. Telson with posterior fourth abruptly narrowed; uropodal exopod<br />
length 4.5 times greatest width bermudensis<br />
Telson tapering, but posterior fourth not abruptly narrowed; uropodal<br />
exopod length 2.5 times greatest width halma<br />
Curassanthura bermudensis Wagele, 1985<br />
Figure 30G,H<br />
DIAGNOSIS 9 3.0 mm. Pereopod 1 propodal length 2.5 times proximal<br />
width, palm with proximal strongly recurved hooklike tooth. Uropodal ex<br />
opod slender, parallel sided, 4.5 times longer than greatest width. Telson<br />
constricted in posterior fourth.<br />
RECORDS Church Cave, Bermuda, in shore sediments.
68 ANTHURIDEA • PARANTHURIDAE<br />
Figure 30. Colanthura tenuis: A, 9; B3 uropod; C, pereopod 1. Curassanthura halma:<br />
D, 9; E, telson; F} uropodal exopod. Curassanthura bermudensis: G3 telson; H3<br />
uropodal exopod (from Wagele, 1985).<br />
Curassanthura halma Kensley, 1981<br />
Figure 30D-F<br />
DIAGNOSIS 9 2.3 mm. Pereopod 1, propodal length about 1.7 times proxi<br />
mal width, palm with eight fringed spines and basal tridentate lobe. Uropo-
Paranthura antillensis 69<br />
dal exopod triangular, length 2.5 times greatest width, shorter than sympod.<br />
Telson tapering but not abruptly narrowed in posterior fourth.<br />
RECORDS Curagao, in shore sediments 1.5 m above tide line; Bonaire, in<br />
shore sediments above tide line.<br />
Paranthura Bate and Westwood, 1868<br />
DIAGNOSIS Eyes present. Antennular flagellum shorter than peduncle. An-<br />
tennal flagellum usually of single flattened setose article. Mandibular palp of<br />
three articles, article 3 with comb of spines. Maxillipedal palp of one or two<br />
articles; endite small to absent. Pereopod 1, propodus inflated, larger than<br />
that of pereopods 2 and 3. Pereopods 4-7, carpi rectangular. Pleonites short,<br />
more or less distinct. Pleopod 1, exopod operculiform. Telson lacking<br />
statocyst.<br />
REMARKS This is the largest of the paranthurid genera, with over 50 names<br />
in the literature. Many of these are poorly described. Species of Paranthura are<br />
common in the shallow waters of the temperate and tropical seas.<br />
Key to species of Paranthura<br />
1. Telson posteriorly truncate; uropodal exopod rectangular, margins<br />
serrate infundibulata<br />
Telson posteriorly rounded; uropodal exopox ovate, margins entire . . 2<br />
2. Uropodal endopod longer than wide 3<br />
Uropodal endopod as long as wide antillensis<br />
3. Uropodal exopod elongate-elliptical; telson parallel sided for half length<br />
jloridensis<br />
Uropodal exopod ovate, outer margin sinuate; telson evenly elliptical<br />
Paranthura antillensis Barnard, 1925<br />
Figure 31A-F<br />
barnardi<br />
DIAGNOSIS 9 5.1 mm. Antennular flagellum of four articles. Mandibular<br />
palp, article 3 with five spines. Maxillipedal palp of single article three times<br />
longer than basal width; short endite present. Pereopod 1, propodus ex-
Figure 31. Paranthura antillensis: Ay 9; B, telson; Cy uropodal exopod; D, uropodal<br />
sympod and endopod; Ey pereopod 1, 9; F, pereopod 2, 9. Paranthura barnardi: Gy<br />
9; H, telson; /, pereopod 1, 9;Jy uropodal exopod; K, uropodal sympod and<br />
endopod.
Paanthura infundibulata 71<br />
panded, palm bearing row of setae, mesial surface near palmar margin with<br />
convex flange and row of about 10 spines. Pereopod 2, propodal palm with<br />
five stout sensory setae. Pleonite 6 free, posterior margin bilobed. Uropodal<br />
exopod ovate, outer margin sinuous; endopod almost circular, as wide as<br />
long. Telson posteriorly rounded.<br />
RECORDS St. Johns, St. James, U.S. Virgin Islands, 32 m; Carrie Bow Cay,<br />
Belize, intertidal to 1.5 m.<br />
Paranthura barnardi Paul and Menzies, 1971<br />
Figure 31G-K<br />
DIAGNOSIS 9 6.0 mm. Mandibular palp article 3 with eight spines. Max-<br />
illipedal palp of single article, length four times basal width. Pereopod 1,<br />
propodal palm concave, with convex flange and row of about 17 spines on<br />
mesial surface. Uropodal exopod broadly ovate, apically subacute; endopod<br />
ovate, length about 1.5 times basal width. Telson evenly elliptical, apex<br />
evenly rounded.<br />
RECORDS Off Venezuela, 95 m.<br />
Paranthura Jloridensis Menzies and Kruczynski, 1983<br />
Figure 32A-E<br />
DIAGNOSIS 9 6.3 mm. Integument with sparse irregular pigmentation.<br />
Mandibular palp article 3 with eight spines. Maxillipedal palp of single arti<br />
cle, length 3.5 times basal width. Pereopod 1, propodal palm with trans<br />
parent flange and row of 10 setae on mesial surface. Uropodal exopod<br />
elongate-elliptical; endopod ovate, length 1.5 times greatest width. Telson<br />
posteriorly broadly rounded, parallel sided for about half its length.<br />
RECORDS OfTSanibel Island, Florida, Gulf of Mexico, 73 m.<br />
Paranthura infundibulata Richardson, 1902<br />
Figure 32F-J<br />
DIAGNOSIS 9 8.2 mm, 6 6.0 mm. Integument with red-brown pigmenta<br />
tion; broad irregular patch between eyes running onto bases of antennules;<br />
pereonites 1 and 2 with anterior patches, remainder of pereonites with pos<br />
terior patches; strong patch on uropodal exopod, endopod, and telson. Man<br />
dibular palp article 3 with 11 or 12 spines. Maxillipedal palp of single article,
Figure 32. Paranthura Jloridensis: A, 9; B, pereopod 1, 9; C, uropodal sympod and<br />
endopod; D, uropodal exopod; Ey telson. Paranthura infundibulata: F> 9; G, telson;<br />
Hy pereopod 1, 9; /, uropodal exopod; J, uropodal sympod and endopod.
ASELLOTA 73<br />
length 2.5 times basal width. Pereopod 1, propodus with convex flange and<br />
row of more than 20 setae. Uropodal exopod elongate-rectangular, mesial<br />
and distal margins serrate; endopod roughly square, margins serrate. Telson<br />
parallel sided, posterior margin truncate.<br />
RECORDS Bermuda, 11-12 m; Carrie Bow Cay, Belize, intertidal to 1 m;<br />
Cozumel, Mexico; Venezuela.<br />
Virganthura Kensley, 1987b<br />
DIAGNOSIS Eyes present. Mandibular palp of three articles. Maxillipedal<br />
palp of three articles; endite present. Pereopods 1-3 subchelate, pereopod 1<br />
larger than pereopods 2 and 3; pereopods 4-7, carpi rectangular. Pleopod 1,<br />
exopod operculiform. Pleonites 1-5 short, distinct; pleonite 6 dorsally de-<br />
marked. Telson with single statocyst.<br />
Virganthura crassa (Barnard, 1925)<br />
Figure 33<br />
DIAGNOSIS 9 6.8 mm. Antennular flagellum of three articles. Antennal<br />
flagellum of seven articles. Maxillipedal endite reaching distal margin of first<br />
palp article. Pereopod 1, propodal palm slightly concave, bearing seven<br />
spines. Uropodal exopod ovate, outer margin sinuate; endopod distally<br />
rounded, articulating obliquely on sympod.<br />
RECORDS U.S. Virgin Islands, 30 m.<br />
Suborder <strong>As</strong>ellota Latreille, 1803<br />
DIAGNOSIS Antennules uniramous. Antennae uniramous, with scale in<br />
some families. Mandible usually with palp, but palp lacking in some groups.<br />
Pereopod 1 usually subchelate, sexually dimorphic in some groups. Coxae<br />
small, sometimes not all visible in dorsal view. Pleon seldom of more than<br />
two free pleonites plus pleotelson. Pleopod 1 absent in 9. One pair of<br />
pleopods in 9, and one or two pairs of pleopods in 6 forming operculum<br />
over remaining respiratory pleopods. Pleopod 2 in 6 usually adapted for<br />
copulation. Uropods usually pedunculate, but peduncle may be reduced,<br />
biramous or uniramous, terminal or subterminal.<br />
REMARKS The suborder <strong>As</strong>ellota is usually divided into four superfamilies,<br />
based on pleopodal arrangement. The great majority of families, however,
A<br />
Figure 33. Virganthura crassa: A, 9; B, maxilliped; C, uropodal endopod and<br />
sympod; D, telson; E, uropodal exopod.<br />
D
Atlantasellus cavernicolus 75<br />
belong to the superfamily Janiroidea, considered to be the most advanced. In<br />
place of a key to the four superfamilies, the chart shown in Figure 34 illus<br />
trates diagrammatically the arrangement of the pleopods in these groups.<br />
Superfamily <strong>As</strong>elloidea Rafinesque-Schmaltz, 1815<br />
DIAGNOSIS 6: Pleopod 1, peduncles separate, rami uniarticulate; pleopod 2<br />
having biarticulate exopod and copulatory endopod; pleopod 3 biramous,<br />
opercular. 9: Pleopod 1 absent; pleopod 2 absent, or of single article;<br />
pleopod 3 biramous, opercular.<br />
Family Atlantasellidae Sket, 1979<br />
DIAGNOSIS Body resembling that of a sphaeromatid isopod. Pleon consist<br />
ing of two free pleonites plus pleotelson. 6: Pleopod 1 of two articles; pleopod<br />
2 with sympod, small exopod, and uniarticulate copulatory endopod. 9:<br />
Pleopods 1 and 2 absent; pleopod 3 operculate but rami not fused.<br />
Atlantasellus Sket, 1979<br />
DIAGNOSIS Eyes lacking. Antennule flattened, short and broad. Antenna<br />
elongate, peduncle of four (?five) articles, flagellum of four articles. Man<br />
dibular palp of three articles; with incisor and lacinia, molar replaced by<br />
brushlike process. Maxillipedal palp of five articles of similar width and<br />
broad endite. Pereopod 1 subchelate; pereopods 2-7 ambulatory, slender,<br />
dactyli biunguiculate. Uropods uniarticulate, vestigial.<br />
Atlantasellus cavernicolus Sket, 1979<br />
Figure 35<br />
DIAGNOSIS 6 1.1 mm. Cephalon with strong triangular rostrum, equal to<br />
pereonites 1-3 combined in middorsal length. Pleotelson basally broad,<br />
tapering to trilobed posterior margin, uropods inserted in incisions between<br />
median and lateral lobes.<br />
RECORDS Walsingham Cave, Bermuda.
PLEOPOD 1<br />
PLEOPOD 2<br />
PLEOPOD 3<br />
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Figure 34. Comparison of pleopods 1-3 in the four superfamilies of the suborder <strong>As</strong>ellota.<br />
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STENETRIOIDEA<br />
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Superfamily Gnathostenetroidoidea Kussakin, 1967<br />
Gnathostenetroid.es pugio 11<br />
DIAGNOSIS 6: Pleopod 1 peduncles fused, uniarticulate rami separate, op<br />
ercular; pleopod 2 small, with biarticulate exopod and copulatory endopod;<br />
pleopod 3, broad endopod biarticulate, exopod slender, uniarticulate. 9:<br />
Pleopod 1 absent; pleopod 2 rami fused to form operculum; pleopod 3 as in<br />
6.<br />
Family Gnathostenetroididae Kussakin, 1967<br />
DIAGNOSIS Uropods with short sympod, rami relatively well developed. 6:<br />
Pleopod 1, protopodites short, fused, rami separate, together forming oper<br />
culum covering remaining pleopods. Pleopod 2, separate, much smaller than<br />
pleopod 1. Pleopod 3, exopod elongate, slender; endopod uniarticulate,<br />
broad. 9: Pleopod 2, rami fused to form operculum covering remaining<br />
pleopods.<br />
Key to genera of Gnathostenetroididae<br />
1. Eyes absent. Rostrum anteriorly rounded/truncate Neostenetroides<br />
Eyes present. Rostrum anteriorly bilobed Gnathostenetroides<br />
Gnathostenetroides Amar, 1957<br />
DIAGNOSIS Eyes present. Rostrum well developed, anteriorly bilobed. Per-<br />
eonite 1, especially in
78 ASELLOTA • GNATHOSTENETROIDIDAE<br />
Figure 35. Atlantasellus cavernicolus: A, 9; B,<br />
pereopod 1. (From Sket, 1979).<br />
bearing row of nine more slender spines, posterior margin setose. 9: Oper<br />
culum with distal incision acute.<br />
RECORDS Florida Middlegrounds, Gulf of Mexico, 55 m.<br />
REMARKS The type of the genus, and only other species known, G. laodicense<br />
Amar, 1957, was recorded from the Mediterranean Sea. This species is in<br />
cluded here, as several of the species first recorded from the Florida Mid<br />
dlegrounds (Hooker, 1985) have since been recorded from the Caribbean.<br />
Neostenetroides Carpenter and Magniez, 1982<br />
DIAGNOSIS Pleonites 1 and 2 very short. Operculum of 9 subcircular.<br />
Pleopod 2 in 6 with elongate protopodite, copulatory organ prolonged by<br />
hyaline tonguelike structure. Pleopod 4, exopod ovate, wider than endopod.<br />
Neostenetroides stocki Carpenter and Magniez, 1982<br />
Figure 36D,E<br />
DIAGNOSIS 6 1.46 mm, 9 1.87 mm. Eyes absent. Rostrum well developed,<br />
anteriorly rounded/truncate. Pleotelson wider than long, lateral margins en<br />
tire, posterior margin regularly convex. Pereopod 1, propodus elongate, wid<br />
ening distally, palm poorly demarked, with few spines; dactylus stout, over<br />
lapping palm. Uropods unknown.<br />
RECORDS San Salvador, Bahamas, from Dixon Hill Lighthouse Cave.
ASELLOTA • JANIROIDEA 79<br />
Figure 36. Gnathostenetroides pugio: A, 8; B, mandible; C, pereopod 1. Neostenetroides<br />
stocki (from Carpenter and Magniez, 1982): Dy 9; E, pereopod 1.<br />
Superfamily Janiroidea Sars, 1899<br />
DIAGNOSIS 6: Pleopod 1 with elongate peduncle, occasionally fused;<br />
pleopod 2 with short exopod and copulatory usually elongate endopod,<br />
pleopods 1 and 2 together forming operculum; pleopod 3 endopod uniarticu-
80 ASELLOTA • JANIROIDEA<br />
late, exopod biarticulate. 9: Pleopod 1 absent; pleopod 2, rami fused to form<br />
operculum; pleopod 3 as in
Figure 37. Mexicope kensleyi: A, 9; B,<br />
pereopod 1.<br />
ASELLOTA • JANIRIDAE 81<br />
REMARKS The inability to place Mexicope in a family reflects the fact that<br />
the arrangement of the families and genera of the Janiroidea is still unsettled.<br />
While having affinities with the Pleurocopidae and the Janiridae, placement<br />
in either of these would require redefinition of both families, making the<br />
Janiridae even more of a hodgepodge of phylogenetically unrelated genera.<br />
Mexicope kensleyi Hooker, 1985<br />
Figure 37<br />
DIAGNOSIS 6 1.7 mm, 9 2.9 mm. Antennae slightly longer than body. Per-<br />
eonites and pleon laterally finely serrate. Pleotelsonic lobe between uropodal<br />
bases narrowly rounded, barely produced.<br />
RECORDS Turks and Caicos Islands, 1 m; Florida Middlegrounds, Gulf of<br />
Mexico, 30 m.<br />
Family Janiridae Sars, 1899<br />
DIAGNOSIS Antennae longer than antennules. Mand<br />
M 3 at least as wide as endite,<br />
markedly broader than articles 4 and 5. Pereopod 1 prehensile, subchelate,<br />
sexually dimorphic, larger in 6 than in 9. Pereopods 2 bulatory<br />
tyli biunguiculate. Coxae visible at least on three posterior pereonites. Pleon
82 ASELLOTA • JANIRIDAE<br />
consisting of single free pleonite (often very narrow, short and in-<br />
conspicuous) plus pleotelson. Uropods with well-developed sympod; usually<br />
biramous.<br />
Carpias Richardson, 1902<br />
DIAGNOSIS Cephalon with dorsolateral eyes, lacking rostrum. Coxae visible<br />
dorsally on pereonites 1-7. Pleon with one short pleonite lacking free lateral<br />
margins, plus broad pleotelson. Antennules and antennae well developed,<br />
latter with scale. Articles 2 and 3 of maxillipedal palp expanded. Pereopod 1<br />
sexually dimorphic, often relatively enormous and/or elongate in male, car-<br />
pochelate, carpus expanded, propodus variously armed or expanded, dac-<br />
Key to species of Carpias (based on pereopod 1 of mature 8)<br />
1. Propodus distally acute 2<br />
Propodus distally broad 4<br />
2. Propodus apically acute, with proximal tooth minutus<br />
Propodus lacking teeth 3<br />
3. Carpus with two strong distal teeth; propodus (when chela closed)<br />
reaching to merus algicola<br />
Carpus with two strong and one small teeth; propodus (when chela<br />
closed) reaching proximal half of carpus serricaudus<br />
4. Propodus with distinct teeth 5<br />
Propodus lacking distinct teeth 6<br />
5. Carpus distally with broadly rounded area; merus elongate-slender,<br />
length about four times greater than width bermudensis<br />
Carpus lacking broadly rounded area; merus short, broader than long<br />
punctatus<br />
6. Carpus with two distal teeth; dactylus minute triton<br />
Carpus with three distal teeth; dactylus small, but not obsolete 7<br />
7. Propodus distally truncate; carpus with middle tooth of palm distally<br />
faintly bilobed brachydactylus<br />
Propodus distally faintly bilobed; carpus with middle tooth of palm<br />
distally narrowly rounded harrietae
Carpias bermudensis 83<br />
tylus reduced or rudimentary, bearing two claws (biunguiculate). Pereopods<br />
2—7 similar, ambulatory, dactylus with three claws (triunguiculate).<br />
Uropods often longer than pleotelson, with relatively elongate sympod and<br />
rami.<br />
REMARKS This genus has been, and continues to be, a source of taxonomic<br />
problems. Several authors (e.g., Pires, 1980) have separated the species into<br />
the genera Carpias and Bagatus; others (e.g., Bowman and Morris, 1979) have<br />
synonymized them. In part, this uncertainty reflects the general uncertainty<br />
of the state of taxonomy in the family Janiridae. In this work, the genus<br />
Carpias is used to contain all the species described under the names Carpias<br />
and Bagatus.<br />
While these tiny asellotes are frequently extremely abundant in certain<br />
habitats (e.g., in reef-crest algal turfs; Kensley, 1983), difficulty is experi<br />
enced in identifying specimens other than mature males. The first pereopod<br />
of the mature male is the feature best used for species separation, but varia<br />
tion with maturity and geographic locality have not been investigated. With<br />
more detailed work, some species will undoubtedly be synonymized.<br />
Carpias aIgicola (Miller, 1941)<br />
Figure 38A,B<br />
DIAGNOSIS 8 2.9 mm, ovigerous 9 2.0 mm. Frontal margin straight. Per<br />
eopod 1 in
84 ASELLOTA • JANIR1DAE<br />
RECORDS Bermuda; eastern and southern coasts of Florida, 1.5-15 m.<br />
Carpias brachydactylus Pires, 1982<br />
Figure 38E<br />
DIAGNOSIS 6 1.6 mm. Pereopod 1 6, carpus distally between two and<br />
three times wider than proximal width, with strong triangular outer tooth<br />
defining palm, middle tooth apically faintly bifid, inner tooth rounded; pro<br />
podus widening to distal truncate margin, overreaching carpal palm by short<br />
distance, with very low tubercle at about midlength of flexor margin.<br />
Pleopod 1 <br />
inner distal lobe rounded, outer lobe narrowly acute. Operculum of 9 with<br />
mediodistal margin gently concave. Uropod about twice length of pleotelson.<br />
RECORDS Biscayne Bay, Florida, intertidal to 2 m.<br />
Carpias minutus (Richardson, 1902)<br />
Figure 39A<br />
DIAGNOSIS 8 1.9 mm, ovigerous 9 1.8 mm. Pereopod 1
Carpias punctatus 85<br />
Figure 38. Carpias algicola: A, 9; B, pereopod 1,
86 ASELLOTA • JANIRIDAE<br />
Figure 39. Carpias minutus: A, pereopod 1, 6. Carpias punctatus: B, 6; C, pereopod<br />
1, 6. Carpias serricaudus: D, pereopod 1, d. Carpias triton: E, F, pereopod 1, 6.<br />
Pereopod 1 6, distal two-thirds parallel sided, with strong acute tooth defin<br />
ing palm, and second rounded tooth; propodus with three lobe-teeth on<br />
flexor surface, overreaching carpus by a third of its length. Pleopod 1
Carpias serricaudus Menzies and Glynn, 1968<br />
Figure 39D<br />
Joeropsis 87<br />
DIAGNOSIS 6 1.6 mm, 9 1.5 mm. Pereopod 1 d, palm of carpus with two<br />
strong outer teeth and one short inner tooth; propodus reaching back to<br />
proximal half of carpus, tapering distally; dactylus obsolete. Pleopod 1 d,<br />
outer distal lobe acute, reaching well beyond inner lobe. Pleotelsonic margins<br />
very faintly serrate. Uropod about 0.7 times length of pleotelson.<br />
RECORDS Turks and Caicos Islands, 1 m; Puerto Rico, intertidal to 1.5 m.<br />
Carpias triton Pires, 1982<br />
Figure 39E,F<br />
DIAGNOSIS 6 2.3 mm. Very similar to C. algicola. Pereopod 1 d, carpus<br />
with two strong basally broad distal teeth; propodus extending back to merus<br />
in adult 8y widening to broadly rounded distal margin; dactylus minute.<br />
Pleopod 1 d, outer distal lobe narrowly acute, reaching well beyond inner<br />
rounded lobe. Uropod about 1.5 times pleotelson length.<br />
RECORDS Carrie Bow Cay, Belize, intertidal reef crest.<br />
Family Joeropsidae Nordenstam, 1933<br />
DIAGNOSIS Cephalon free, with distinct rostrum. Molar process of mand<br />
ible reduced. Maxillipedal palp articles all of similar width. Antenna short,<br />
peduncle dilated, flagellum reduced. Pereonites similar, wider than long.<br />
Pereopods similar, biunguiculate. Uropods having short squat sympod and<br />
very reduced rami; inserted into submedian posterior notches of pleotelson.<br />
Joeropsis Koehler, 1885<br />
DIAGNOSIS Dorsolateral eyes present. Antennule, basal article widest and<br />
longest, often with transparent distal dentition. Antenna, peduncular articles<br />
3-5 somewhat dilated, article 2 often with fringe of transparent scales;<br />
flagellum of about six articles, together shorter than peduncle article 5. Per<br />
eonites similar, generally subequal in length and width. Pleotelson of single<br />
shield-shaped segment. Uropodal sympod usually with mesiodistal angle<br />
acute; rami reduced.
88 ASELLOTA • JOEROPSIDAE<br />
Key to species of Joeropsis<br />
1. Lateral margins of cephalon serrate; rostrum triangular personatus<br />
Lateral margins of cephalon entire; rostrum not triangular 2<br />
2. Body glabrous; strong band of pigment on cephalon and pereonite 4<br />
bifasciatus<br />
Body setose; pigment in reticulation over entire body 3<br />
3. Rostrum evenly convex; outer uropodal ramus longer than inner<br />
rathbunae<br />
Rostrum anteriorly shallowly notched; outer uropodal ramus shorter<br />
than inner coralicola<br />
Joeropsis bifasciatus Kensley, 1984<br />
Figure 40A-F<br />
DIAGNOSIS 6 2.5 mm, 9 2.4 mm. Body glabrous. Lateral margins of<br />
cephalon entire. Rostrum semicircular, with marginal flange of transparent<br />
teeth. Antennal flagellum of eight articles. Lateral margins of pleotelson ser<br />
rate. Apex of 9 operculum blunt. Broad band of pigment on cephalon be<br />
tween eyes and almost reaching posterior margin; broad band of pigment on<br />
pereonite 4.<br />
RECORDS Carrie Bow Gay, Belize, 1-6 m, often on Agaricia sp. and Porites<br />
sp. corals, and Halimeda sp. alga; Anguilla.<br />
Joeropsis coralicola Schultz and McCloskey, 1967<br />
Figure 40G<br />
DIAGNOSIS 6 2.0 mm, ovigerous 9 1.9 mm. Body setose. Lateral margins<br />
of cephalon entire. Rostrum anteriorly notched. Antennal flagellum of five<br />
articles. Lateral margins of pleotelson serrate. Apex of 9 operculum acute.<br />
Outer uropodal ramus shorter than inner. Pigment spread as reticulation<br />
over entire body.<br />
RECORDS Off North Carolina, on coral Oculina arbuscula; Florida Mid-<br />
dlegrounds, Gulf of Mexico, from sponge Agelas sp. and coral Madracis sp.,<br />
25-33 m.
Figure 40. Joeropsis bifasciatus: A, 8; B, pleopod 1, 6; C, uropod; D, operculum, 9;<br />
E> antennule; F, antenna. Joeropsis coralicola: G, d. Joeropsis pe?sonatas: Hy 6.<br />
Joeropsis rathbunae: I, 6.
90 ASELLOTA • MICROPARASELLIDAE<br />
Joeropsis personatus Kensley, 1984<br />
Figure 40H<br />
DIAGNOSIS 6 2.2 mm, 9 2.0 mm. Body glabrous. Lateral margin of<br />
cephalon serrate. Rostrum triangular. Lateral margins of pleotelson serrate.<br />
Antcnnal flagellum of five articles. Apex of 9 operculum acute. Outer uropo-<br />
dal ramus shorter than inner. Strong band of pigment on cephalon; rest of<br />
body with paler reticulation of pigment.<br />
RECORDS Carrie Bow Cay, Belize, on Pontes sp. and Madracis sp. corals,<br />
and on Halimeda sp. alga, 1—20 m.<br />
Joeropsis rathbunae Richardson, 1902<br />
Figure 401<br />
DIAGNOSIS 6 1.9 mm, ovigerous 9 1.6 mm. Body setose overall. Lateral<br />
margins of cephalon entire. Rostrum evenly convex with flange of trans<br />
parent teeth. Antennal flagellum of three articles. Lateral margins of<br />
pleotelson serrate. Apex of 9 operculum acute. Outer uropodal ramus longer<br />
than inner. Pigment in reticulation over entire body.<br />
RECORDS Bermuda; Florida Keys; Turks and Caicos Islands; Puerto Rico;<br />
Gulf of Mexico; intertidal to 36 m.<br />
Family Microparasellidae Karaman, 1933a<br />
DIAGNOSIS Eyes lacking. Antennule much shorter than antenna. Antenna<br />
with scale. Pereopods all similar, with biunguiculate dactyli. Pleon of one<br />
free pleonite plus pleotelson. Uropods with well-developed sympod and<br />
rami.<br />
Key to genera of Microparasellidae<br />
1. d, plcopod 1 narrow, not overlapping external part of pleopod 2;<br />
maxillipedal palp of five articles Microcharon<br />
6, pleopod 1 broad, almost completely covering pleopod 2;<br />
maxillipedal palp of four articles, terminal article ending in pointed<br />
process Angliera
Microcharon sabulum 91<br />
REMARKS All the members of this family are tiny (usually less than 2 mm),<br />
and most are interstitial in habit, being found in marine, brackish, and fresh<br />
water environments.<br />
Angliera Chappuis and Delamare Deboutteville, 1955<br />
DIAGNOSIS Mandibular palp with two proximal articles inflated, terminal<br />
article, slender hooklike, articles lacking setae and spines. Maxillipedal palp<br />
of four articles, articles 1 and 3 elongate, article 2 short, article 4 with termi<br />
nal acute process. Pleopod 1 in 6 forming broad lamella.<br />
Angliera psamathus Kensley, 1984<br />
Figure 41A-D<br />
DIAGNOSIS 6 1.0 mm, 9 1.0 mm. Maxillipedal endite with seven setae on<br />
distal margin. Posterior four pairs of pereopods with claw on dactylus dorsal<br />
to unguis. Uropodal endopod subequal in length to sympod.<br />
RECORDS Carrie Bow Cay, Belize, interstitial in intertidal sand bank.<br />
REMARKS Two other species of Angliera have been recorded from the Carib<br />
bean area: A. dubitans Stock, 1977, from Bonaire, and A. racovitzai Coineau<br />
and Botosaneanu, 1973, from Cuba. The reader should refer to the original<br />
descriptions to distinguish the species, as differences are extremely subtle.<br />
Microcharon Karaman, 1934<br />
DIAGNOSIS Mandibular palp of three articles, two distal articles bearing<br />
spines and/or setae. Maxillipedal palp of five articles, articles 1, 2 and 3<br />
expanded. c£, pleopod 1 narrow, elongate, not obscuring pleopod 2.<br />
REMARKS More than 20 species of Microcharon have been described world<br />
wide. The genus is unusual in that the species have been found in true ma<br />
rine environments, in brackish habitats such as wells, and inland in<br />
freshwater.<br />
Microcharon sabulum Kensley, 1984<br />
Figure 41E-H<br />
DIAGNOSIS 6 1.4 mm, 9 1.5 mm. Antennule of five articles. Inner ramus of<br />
maxilla 2 with pectinate spine. Pereopodal dactyli short, biunguiculate. En-
Figure 41. Angliera psamathus: A, 6; B, right, and part of left, mandible;<br />
maxilliped; Dy pleopod 1, 6. Microcharon sabulum: Ey 6\ Fy mandible; G,<br />
maxilliped; Hy pleopod 1, d.
Munna 93<br />
dopod of pleopod 3 with three distal plumose setae. Uropodal sympod stout,<br />
longer than rami.<br />
RECORDS Carrie Bow Gay, Belize, interstitial in intertidal sand bank.<br />
REMARKS Two other species of Microcharon have been described from the<br />
Caribbean area: M. phreaticus Coineau and Botosaneanu, 1973, from intersti<br />
tial freshwater on Cuba, and M. herrerai Stock, 1977, from freshwater wells on<br />
Bonaire. The reader should refer to the original descriptions to distinguish<br />
these species.<br />
Family Munnidae Sars, 1899<br />
DIAGNOSIS Body ovate. Cephalon and all pereonites free; pleon narrower<br />
than rest of body, longer than broad. Eyes on lateral processes of cephalon.<br />
Mandible with molar and incisor present; palp present or absent. Maxillipe-<br />
dal palp articles 2 and 3 broader than remaining articles. Pereopod 1 prehen<br />
sile; pereopods 2—7 ambulatory. Uropods tiny, without sympod. Anus ex<br />
posed, not covered by pleopods.<br />
REMARKS Poore (1984) has provided the most useful and recent survey of<br />
the genera, and especially of Munna.<br />
Key to genera of Munnidae<br />
1. Mandibular palp present; pereopod 1 enormous in
94 ASELLOTA • MUNNIDAE<br />
Munna petronastes Kensley, 1984<br />
Figure 42A-D<br />
DIAGNOSIS
Uromunna reynoldsi 95<br />
Figure 42. Munna petronastes: A, 8\ B, uropod; C, pereopod 1, 8] D, pereopod 1 9<br />
(C and D same scale). Uromunna caribea: E, 6; F, larger uropodal ramus. Uromunna<br />
reynoldsi: G, pereopod 1, 8; Hy larger uropodal ramus; /, 8.<br />
Uromunna reynoldsi Frankenberg and Menzies, 1966<br />
Figure 42G-I<br />
DIAGNOSIS 6 1.5 mm, 9 1.6 mm. Propodus of pereopod 1 two or three<br />
times longer than wide. Operculum of 9 distally rounded. Shorter uropodal<br />
ramus obscured by pleotelsonic margin, with single seta. Pigmentation a
96 ASELLOTA • PARAMUNNIDAE<br />
broad patch between eyes on cephalon, lateral bands on pereon, and anterior<br />
and lateral patches on pleon.<br />
RECORDS Sapelo Island, Georgia, in tidal saltmarsh creek; Lake<br />
Ponchartrain, Louisiana; Atlantic and Pacific locks of Panama Canal.<br />
Family Paramunnidae Vanhoffen, 1914<br />
DIAGNOSIS Body broad, ovate, often with laterally produced tergal or epi-<br />
meral plates. Cephalon recessed into pereonite 1. Eyes, if present, on lateral<br />
projections of cephalon. Antennule short, usually of six articles, with single<br />
terminal aesthetasc. Antenna never longer than body. Mandibular palp pres<br />
ent or absent. Pereopod 1 prehensile; pereopods 2-7 ambulatory. Pleopod 1<br />
6 distally sagittate. Uropods with sympod minute or absent; rami tiny. Anus<br />
covered by pleopods.<br />
Munnogonium George and Stromberg, 1968<br />
DIAGNOSIS Eyes present on short lateral processes of cephalon. Antennal<br />
peduncular scale present. Coxal plates visible on pereonites 2—7.<br />
Munnogonium wilsoni Hooker, 1985<br />
Figure 43A,B<br />
DIAGNOSIS 6 0.86 mm, 9 0.98 mm. Frontal margin of cephalon broadly<br />
rounded. Mandibular palp absent. Uropodal endopod twice length of ex-<br />
opod. Lateral margins of pleotelson to uropodal insertion serrate, posterior<br />
margin between uropodal insertions tapering to rounded apex.<br />
RECORDS Florida Middlegrounds, Gulf of Mexico, 55 m.<br />
Family Pleurocopidae Fresi and Schiecke, 1972<br />
DIAGNOSIS Cephalon broader than long. Eyes (or at least ocular peduncles)<br />
present. Mandible with or without palp; molar truncate. Maxillipedal palp<br />
articles narrow, less than half width of endite. At least coxae of pereonites 5-<br />
7 dorsally visible. Pereopods 2-7 uni- or biunguiculate. Pleopod 1 in 6 not<br />
sagittate. Uropod pedunculate, inserted laterally or slightly dorso- or ven<br />
trolateral^ on pleotelson; biramous, or with one ramus fused with sympod.
Pleurocope 97<br />
Figure 43. Munnogonium wilsoni: A, 9; B, pereopod 1,
98 ASELLOTA • SANTIIDAE<br />
Figure 44. Pleurocope jloridensis: adult in dorsal<br />
view.<br />
Pleurocope Jloridensis Hooker, 1985<br />
Figure 43C 44<br />
DIAGNOSIS 1.15 m m , $ 0 . 9 6 m m . B o d y o v a t e , t a p e r i n g p o s t e r i o r l y . I n <br />
t e g u m e n t v e r y finely t u b e r c u l a t e . M e s i o d i s t a l l o b e o n a n t e n n a l p e d u n c l e a i<br />
t i d e 3 b e a r i n g five d i s t a l s e t a e . P e r e o n l a c k i n g l o n g d o r s a l s e t a e . P e r e o p o d<br />
s u b c h e l a t e , b u t a l m o s t c a r p o c h e l a t e . P l e o n c o n s i s t i n g o f s i n g l e s e g m e n t , p o s <br />
t e r i o r l y n a r r o w l y t a p e r e d a n d p r o d u c e d . U r o p o d a l r a m i a s l o n g a s s y m p o d .<br />
RECORDS T u r k s a n d C a i c o s I s l a n d s , 1 m ; C a r r i e B o w C a y , B e l i z e , 3 - 1 0 m ;<br />
F l o r i d a M i d d l e g r o u n d s , G u l f o f M e x i c o , 5 5 m .<br />
F a m i l y S a n t i i d a e W i l s o n , 1 9 8 7<br />
D I A G N O S I S A n t e n n f l a g e l l u m w i t h a t m o s t , t h r e e a r t i<br />
s c a l e s o m e t i m e s p r e s e n t . P e r e o p o d 1 p r e h e n s i l e . P e r e o p o d s<br />
a n t e n n u l a r<br />
d a c t y l i<br />
b i u n g u i c u l a t e . C o x a e v i s i b l e a t l e a s t o n p e r e o n i 7. P l e o n c o n s i s t i n g o f<br />
s i n g l e s h o r t p l e o n i t e p l u s p l e o t e l s o n . U r o p o d s p e d u n c u l a t e , b i r a m o u s , i n s e r <br />
t e d d o r s a l l y o r l a t e r a l l y . ( O n e s p e c i e s o f Santia p o s s e s s e s a u n i r a m o u s<br />
u r o p o d . )<br />
S a n t i a S i v e r t s e n a n d H o l t h u i s , 1 9 8 0<br />
D I A G N O S I S C e p h a l o n a b o u t t w i c e w i d e r t h a n l o n g . E y e s p r e s e n t . A n t e n n u <br />
l a r p e d u n c l e o f t h r e e a r t i c l e s . P e r e o n i t e s l a t e r a l l y r o u n d e d , s o m e t i m e s b e a r <br />
i n g s h o r t l a t e r a l s p i n e s .
Santia milleri (Menzies and Glynn, 1968)<br />
Figure 43F-H<br />
ASELLOTA • STENETRIIDAE 99<br />
DIAGNOSIS 6 and 9 1.0 mm. Eye on short lateral process of cephalon.<br />
Mandibular palp of three articles. Maxillipedal palp articles all of similar<br />
width. Pereopod 1 barely subchelate. Uropod with sympod well developed,<br />
rami prominent, well developed.<br />
RECORDS Carrie Bow Cay, Belize, intertidal to 30 m; Puerto Rico, 1.5 m;<br />
San Salvador, Bahamas, 6 m; Turks and Caicos Islands, 1 m; Anguilla; Ja<br />
maica; Cozumel, Mexico; Gulf of Mexico.<br />
Brazil, 1-6 m.<br />
Superfamily Stenetrioidea Hansen, 1905a<br />
DIAGNOSIS 6: Pleopod 1 small, peduncles fused, rami separate, uniarticu-<br />
late; pleopod 2 small, copulatory; pleopod 3 biramous, opercular. 9 : Pleopod<br />
1 absent; pleopod 2 rami and peduncle fused to form operculum; pleopod 3<br />
as in 6.<br />
Family Stenetriidae Hansen, 1905a<br />
DIAGNOSIS 6: Pereopod 1 frequently much bigger than in 9, with distinc<br />
tive lobes and teeth. Pleopods 1 and 2 reduced; pleopod 1 protopodite short,<br />
fused, rami separate. Pleopod 2, endopod elongate, flexed, exopod short.<br />
Pleopod 3 exopod basally broad, distally narrowed; endopod broad, Particu<br />
late. 9: Pleopod 2, rami fused, short, covering base of pleopod 3. Uropod<br />
with short sympod, rami relatively well developed, styliform, of single article.<br />
Key to genera of Stenetriidae<br />
1. Rostrum narrowly triangular, spikelike; two free, very short pleonites<br />
anterior to pleotelson Stenobermuda<br />
Rostrum short, basally broad, anteriorly truncate or broadly rounded;<br />
two or three very short free pleonites anterior to pleotelson<br />
Stenetrium
100 ASELLOTA • STENETRI1DAE<br />
Stenetrium Haswell, 1881<br />
DIAGNOSIS Eyes present. Cephalon broader than long. Rostrum short,<br />
basally broad, anteriorly truncate or rounded. Pereonites 1—4 with ante<br />
rolateral projections; pereonites 5-7 projecting posteriorly. Pleotelson with<br />
sharp tooth anterior to small lateral notch.<br />
REMARKS If fresh material is not available, and color pattern is lost in pres<br />
ervation, mature male material is needed as identification is based on the<br />
structure of male pereopod 1.<br />
Key to species of Stenetrium<br />
1. Eyes of few (not more than 10) ommatidia, not reniform 2<br />
Eyes of many ommatidia, reniform 3<br />
2. Eyes of four ommatidia; 6 pereopod 1 small, propodus not unusually<br />
broad minocule<br />
Eyes of more than four ommatidia; 6 pereopod 1, propodus broad,<br />
with wide palm patulipalma<br />
3. Pleotelsonic margins serrate 4<br />
Pleotelsonic margins entire 5<br />
4. Rostrum convex, with fine marginal teeth; pereopod 1 c?, propodal<br />
palm with three straight teeth bowmani<br />
Rostrum truncate; pereopod 1 cJ, propodal palm with two teeth, outer<br />
tooth elongate, curved serratum<br />
5. Pereopod 1 cJ, carpus produced, apically acute stebbingi<br />
Pereopod 1 cJ, carpus produced, apically rounded spathulicarpus<br />
Stenetrium bowmani Kensley, 1984<br />
Figure 45<br />
DIAGNOSIS 8 5.0 mm, 9 5.2 mm. Rostrum convex, with tiny marginal<br />
teeth. Lateral lobes of cephalon acute, margins serrate. Color pattern in<br />
small scattered red-brown chromatophores; irregular unpigmented patches<br />
on cephalon. pereonite 4, and pleon; chalky-white bands on antennae and<br />
uropods. 8: Pereopod 1 propodus broad, palm with three teeth, outermost
Stenetrium bowmani 101<br />
Figure 45. Stenetrium bowmani: A, 6; B, pereopod 1, 6; C, pereopod 1, 9; D,<br />
pleopod \,6;E, pleopod 2, 6; F, pleopod 3; G, pleopod 4; H, operculum, 9.
102 ASELLOTA • STENETRIIDAE<br />
longest, slender. 9: Pereopod 1 propodus with strong denticulate spine de-<br />
marking palm, latter straight, with row of about seven slender spines.<br />
RECORDS Cozumel, Mexico; Carrie Bow Cay, Belize, 0.5-15.2 m, on algae<br />
and corals in reefcrest, and spur and groove zone of reef.<br />
Stenetrium minocule Menzies and Glynn, 1968<br />
Figure 46A-C<br />
DIAGNOSIS 8 2.8 mm, 9 3.7 mm. Eye of four ommatidia. Anterolateral<br />
lobes of cephalon blunt, barely produced. Rostrum poorly defined, truncate.<br />
8: Pereopod 1, carpus produced posterodistally into broadly rounded lobe;<br />
propodus broad, palm demarked by strong spine, with six low rounded teeth.<br />
9: Pereopod 1 propodus little broadened, palm demarked by strong denticu<br />
late spine, bearing several more spines.<br />
RECORDS Puerto Rico, intertidal to 3 m; Carrie Bow Cay, Belize, intertidal<br />
to 36 m, from rubble, algal turfs, and seagrass.<br />
Stenetrium patulipalma Kensley, 1984<br />
Figure 46D,E<br />
DIAGNOSIS 8 2.0 mm, 9 2.7 mm. Eyes of about 10 ommatidia in cluster.<br />
Rostrum poorly defined, truncate. Two basal articles of maxillipedal palp<br />
not enlarged. 8: Pereopod 1 unknown. 9: Pereopod 1 broadening to palm,<br />
and bearing row of about 12 small fringed spines. Color pattern: entire body<br />
with red-brown reticulation, dark transverse bars anteriorly on cephalon,<br />
pereonites 2 and 3, posteriorly on pereonites 4-7.<br />
RECORDS Carrie Bow Cay, Belize, 9.1—27.4 m; Barbados; Jamaica.<br />
Stenetrium serratum Hansen, 1904<br />
Figure 46F-H<br />
DIAGNOSIS 8 4.0 mm, ovigerous 9 4.9 mm. Rostrum truncate. Pereonites<br />
1-5 with acute anterolateral angles. Pleotelsonic lateral margins with five<br />
teeth. 8: Pereopod 1, propodus broad, palm with three teeth, outermost<br />
elongate, curved; dactylus reaching well beyond outermost palmar tooth. 9:<br />
Pereopod 1, propodus much smaller than in 8, palm bearing series of about<br />
nine fringed spines. Color: tiny red-brown chromatophores arranged in re-
Figure 46. tart. —«* *. * A P«-P od ' * C " "TT" ' '' ^TTH<br />
pLpalma: D,2;E, pereopod 1 «. ta*te —to.' F, 3; C, pereopod 1 9, H,<br />
pereopod 1 6.
104 ASELLOTA • STENETRUDAE<br />
ticulatc bands; distinctive open patches on cephalon and pereonite 1; pleon<br />
with two broad transverse bands.<br />
RECORDS Looe Key, Florida, 0.5-6 m; Turks and Caicos Islands, 1 m;<br />
Jamaica; Puerto Rico, intertidal to 3 m; St. Thomas, U.S. Virgin Islands;<br />
Carrie Bow Cay, Belize, intertidal to 15 m.<br />
Stenetrium spathulicarpus Kensley, 1984<br />
Figure 47A-C<br />
DIAGNOSIS 6 4.1 mm, 9 4.1 mm. Rostrum truncate. 6: Pereopod 1, merus<br />
and ischium each with setose fingerlike anterodistal projection; carpus with<br />
large spatulate and setose lobe almost reaching level of palm; propodal palm<br />
with large outer tooth and four or five low rounded teeth, broad band of setae<br />
near anterior margin; dactylus reaching slightly beyond palm, with band of<br />
setae along anterior margin. 9: Propodal palm straight, with row of slender<br />
spines; band of setae in similar position as in 6. Color: pigment in ill-defined<br />
and scattered reticulation; strong band on cephalon between eyes.<br />
RECORDS Carrie Bow Cay, Belize, intertidal to 36 m; Puerto Rico,<br />
intertidal.<br />
Stenetrium stebbingi Richardson, 1902<br />
Figure 47 D-H<br />
DIAGNOSIS 6 4.8 mm, ovigerous 9 4.1 mm. Rostrum truncate. Ommatidia<br />
of eye more bunched than in S. spathulicarpus. 6: Pereopod 1 variable accord<br />
ing to maturity; carpus produced posterodistally into narrowly triangular,<br />
apically acute lobe; propodal palm poorly defined, with group of two to four<br />
teeth near dactylar articulation. 9: Pereopod 1, palm straight, defined by<br />
strong outer tooth and bearing row of six or seven low rounded tubercles.<br />
Color: irregular reticulation of red-brown pigment, no strong band between<br />
eyes.<br />
RECORDS Bermuda, 0.5-4 m; Florida Keys, 18.3 m; Bahamas, 5 m; Turks<br />
and Caicos Islands, 1 m; Cuba; Jamaica; U.S. Virgin Islands, 50 m; Carrie<br />
Bow Cay, Belize, 0.5-36 m; Gulf of Mexico.
Figure 47. Sterutrium sPathulicarPus: A,6;B, pereopod 1, 6, many setae removed;<br />
C pereopod 1,9. Stenetnum stebbingi: D, 8; E, pereopod 1, 9; F, G, H, vanauon<br />
in pereopod 1,
106 ASELLOTA • STENETRIIDAE<br />
Figure 48. Stenobermuda acutirostrata: A, 9; By pereopod 1,9.<br />
Stenobermuda Schultz, 1979<br />
DIAGNOSIS Eyes of few ommatidia; rostrum narrow-based, elongate and<br />
spikelike. Pleon consisting of two free pleonites plus pleotelson, with<br />
posterolateral notch marked by tooth.<br />
Stenobermuda acutirostrata Schultz, 1979<br />
Figure 48<br />
DIAGNOSIS 6 4.8 mm. Eyes having five ommatidia. Spikelike rostrum<br />
reaching well beyond anterolateral angles of cephalon. Pereopod 1, propodus<br />
longer than wide, palm straight, bearing eight fringed spines, posterior mar<br />
gin with six spines and several setae.<br />
RECORDS Off Bermuda, 90 m; Turks and Caicos, 1 m.
Suborder Epicaridea Latreille, 1831<br />
EP1CARIDEA 107<br />
DIAGNOSIS Predominantly ectoparasites of marine crustaceans, feeding on<br />
blood. Eyes sessile, usually present in
Figure 49. A, epicaridium larva, lateral view; B> epicaridium larva, ventral view;<br />
C, cryptoniscium larva, ventral view (from Bonnier, 1900).
EP1GAR1DEA 109<br />
Figure 50. A, caridean shrimp with bopyrid parasite in branchial chamber.<br />
Probopyrus pandalicola: B, 9 and 6 in dorsal view, same scale; C, 6 enlarged; D, 9,<br />
ventral view, eggs removed from marsupium.<br />
cephalothorax of the host, attached dorsally to the carapace, ventrally and<br />
laterally in the gill chambers and on the pereopods, or in the brood<br />
chambers.<br />
The Entoniscidae are internal parasites of decapod crustaceans, being<br />
found in the visceral cavity, with the parasite's head in the position of the<br />
host's gonads or hepatopancreas. Veillet (1945) demonstrated that a pore to<br />
the host's branchial chamber connecting the parasite to the exterior is pres<br />
ent only in hosts with mature parasites, to facilitate the release of epi-<br />
caridium larvae.<br />
The Cryptoniscidae are protandrous hermaphrodites. The female is
110<br />
TABLE 2. CARIBBEAN EPICARIDEAN ISOPODS, THEIR HOSTS AND LOCALITIES<br />
Achelion occidentalis Hartnoll, 1966<br />
Microphrys bicornutus (Latreille)<br />
Stenorhynchus seticornis (Herbst)<br />
Jamaica<br />
Synalpheus pectiniger Coutiere<br />
North Carolina to Florida; Bahamas;<br />
Hispaniola; Jamaica; Bonaire;<br />
Curasao; Belize; Gulf of Mexico<br />
Aporobopyrina anomala Markham, 1973 Balanopleon tortuganus Markham, 1973<br />
Munida valida Smith<br />
Florida Keys; off Colombia; Gulf<br />
of Mexico<br />
Aporobopyrus curtatus (Richardson,<br />
1904)<br />
Petrochirus diogenes (Linnaeus)<br />
Petrolisthes armatus (Gibbes)<br />
Petrolisthes galathinus (Bosc)<br />
Petrolisthes marginatus Stimpson<br />
Porcellana sayana (Leach)<br />
Florida Keys; U.S. Virgin Islands;<br />
North Carolina<br />
Argeia atlantica Markham, 1977<br />
Sclerocrangon jacqueti (A. Milne<br />
Edwards)<br />
Bahamas; Newfoundland<br />
<strong>As</strong>talione cruciaria Markham, 1975b<br />
Clastotoechus vanderhorsti (Schmitt)<br />
U.S. Virgin Islands<br />
<strong>As</strong>ymmetrione clibanarii Markham,<br />
1975d<br />
Clibanarius tricolor (Gibbes)<br />
Florida; Bahamas; <strong>As</strong>cension<br />
Island<br />
<strong>As</strong>ymmetrione desultor Markham, 1975d<br />
Pagurus bonairensis Schmitt<br />
Pagurus longicarpus Say<br />
Pagurus provenzanoi Forest and de<br />
Saint Laurent<br />
Pylopagurus sp.<br />
North Carolina; Florida Keys;<br />
Curasao; Bonaire<br />
Azygopleon schmitti (Pearse, 1932)<br />
Synalpheus brooksi Coutiere<br />
Synalpheus hemphilli Coutiere<br />
Synalpheus longicarpus Coutiere<br />
Synalpheus mcclendoni Coutiere<br />
Munida simplex Benedict<br />
Tortuga Island<br />
Bopyrella harmopleon Bowman, 1956<br />
Synalpheus brevicarpus (Herrick)<br />
Synalpheus fritzmuelleri Coutiere<br />
Synalpheus hemphilli Coutiere<br />
Synalpheus minus Say<br />
Venezuela; Brazil<br />
Bopyrina abbreviata Richardson, 1904<br />
Hippolyte curacaoensis Schmitt<br />
Hippolyte pleuracanthus (Stimpson)<br />
Hippolyte zostericola (Smith)<br />
North Carolina to Florida; Belize;<br />
West Indies; Gulf of Mexico<br />
Bopyrinella thorii (Richardson, 1904)<br />
Thor Jloridanus Kingsley<br />
Florida; Curasao; Yucatan<br />
Peninsula, Mexico<br />
Bopyrione synalphei Bourdon and<br />
Markham, 1980<br />
Synalpheus goodei Coutiere<br />
Synalpheus bousfieldi Chace<br />
Synalpheus brevicarpus (Herrick)<br />
Synalpheus pectiniger Coutiere<br />
Florida; Haiti; Curasao; Gulf of<br />
Mexico<br />
Bopyrissa wolffi Markham, 1978<br />
Clibanarius tricolor (Gibbes)<br />
Clibanarius vittatus (Bosc)<br />
Bermuda; North Carolina to Florida;<br />
Bahamas; Puerto Rico; Gulf of<br />
Mexico<br />
Cabirops sp.<br />
Synsynella deformans Hay<br />
Bermuda<br />
Cancricepon choprae (Nierstrasz and<br />
Brender a Brandis, 1925)
Domecia acanthophora (Desbonne<br />
and Schramm)<br />
Domecia hispida Eydoux and<br />
Souleyet<br />
Eriphia gonagra (Fabricius)<br />
Hexapanopeus angustifrons (Benedict<br />
and Rathbun)<br />
Micropanope barbadensis Rathbun<br />
Neopanope packardii (Kingsley)<br />
Neopanope texana sayi (Smith)<br />
Panopeus herbstii H. Milne Edwards<br />
Panoplax depressa Stimpson<br />
Paraliomera dispar (Stimpson)<br />
Rithropanopeus harrisii (Gould)<br />
Carolinas to Florida; Bermuda;<br />
Curagao; Gulf of Mexico<br />
Cancrion carolinas Pearse and Walker,<br />
1939<br />
Panopeus herbstii H. Milne Edwards<br />
North Carolina; Bahamas<br />
Dactylokepon caribaeus Markham,<br />
1975c<br />
Iliacantha liodactyla Rathbun<br />
Iliacantha subglobosa Stimpson<br />
Dominican Republic; Costa Rica-<br />
Panama<br />
Dicropleon periclimenis Markham, 1972a<br />
Periclimenes americanus Kingsley<br />
St. Lucia Island<br />
Diplophryxus sp. (see Markham, 1985)<br />
Alpheus formosus Gibbes<br />
Georgia; Florida; Yucatan, Mexico<br />
Eophrixus subcaudalis (Hay, 1917)<br />
Synalpheus brooksi Coutiere<br />
Synalpheus goodei Coutiere<br />
Synalpheus hemphilli Coutiere<br />
Synalpheus longicarpus (Herrick)<br />
Synalpheus mcclendoni Coutiere<br />
Synalpheus pandionis Coutiere<br />
Synalpheus pectiniger Coutiere<br />
North Carolina to Florida;<br />
Yucatan Pensinsula, Mexico;<br />
Belize; Hispaniola; Curasao<br />
Gigantione mortenseni Adkison, 1984b<br />
Dromidia antillensis Stimpson<br />
Hypoconcha sabulosa (Herbst)<br />
Hypoconcha spinosissima Rathbun<br />
Florida; Haiti; Yucatan, Mexico;<br />
U.S. Virgin Islands; Gulf of<br />
Mexico<br />
Hemiarthrus synalphei (Pearse, 1950)<br />
Synalpheus fritzmuelleri Coutiere<br />
Synalpheus hemphilli Coutiere<br />
Synalpheus longicarpus (Herrick)<br />
North Carolina to Florida; Haiti;<br />
Gulf of Mexico<br />
Leidya bimini Pearse, 1951<br />
Cyclograpsus interger (H. Milne<br />
Edwards)<br />
Pachygrapsus transversus (Gibbes)<br />
Sesarma ricordi H. Milne Edwards<br />
Bermuda; Florida Keys; Bahamas;<br />
U.S. Virgin Islands; Jamaica;<br />
Panama<br />
Leidya distorta (Leidy, 1855)<br />
Uca pugilator (Bosc)<br />
Uca spp.<br />
New Jersey to Florida; Guadeloupe;<br />
Trinidad<br />
Loki circumsaltanus Markham, 1972a<br />
Thor Jloridanus Kingsley<br />
Thor manningi Chace<br />
Southern Florida; U.S. Virgin<br />
Islands; Belize<br />
Metaphrixus carolii Nierstrasz and<br />
Brender a Brandis, 1931<br />
Hippolyte pleuracanthus Stimpson<br />
Southern Florida; U.S. Virgin<br />
Islands<br />
Munidion cubense Bourdon, 1972<br />
Munidajlinti Benedict<br />
Munida stimpsoni A. Milne Edwards<br />
Cuba; Venezuela<br />
Munidion irritans Boone, 1927<br />
Munida irrasa A. Milne Edwards<br />
Florida Keys; Belize<br />
111<br />
{continued)
12<br />
TABLE 2. {Continued)<br />
Munidion longipedis Markham, 1975a<br />
Munida longipes A. Milne Edwards<br />
Munida schroederi Chace<br />
East coast of Florida; Florida<br />
Keys; Cuba; Gulf of Mexico<br />
Parabopyrella lata (Nierstrasz and<br />
Brender a Brandis, 1929)<br />
Alpheus normanni Kingsley;<br />
Upogebia affinis (Say)<br />
Florida; U.S. Virgin Islands;<br />
Brazil<br />
Parabopyrella mortenseni (Nierstrasz<br />
and Brender a Brandis, 1929)<br />
Lysmata rathbunae Chace<br />
Lysmata wurdemanni (Gibbes)<br />
Florida; U.S. Virgin Islands;<br />
Venezuela<br />
Parabopyrella richardsonae (Nierstrasz<br />
and Brender a Brandis, 1929)<br />
Alpheus formosus Gibbes<br />
Alpheus heterochaelis (Say)<br />
U.S. Virgin Islands; Gulf of<br />
Mexico<br />
Parabopyrella thomasi (Nierstrasz and<br />
Brender a Brandis, 1929)<br />
Tozeuma carolinense Kingsley<br />
U.S. Virgin Islands<br />
Parathelges piriformis Markham, 1972b<br />
Paguristes oxyophthalmus Holthuis<br />
Pagurus brevidactylus (Stimpson)<br />
Pagurus provenzanoi Forest and de<br />
Saint Laurent<br />
Bermuda; Bahamas; Colombia<br />
Parathelges tumidipes Markham, 1972b<br />
Allodardanus bredini Haig and<br />
Provenzano<br />
Dardanus fucosus Biffar and<br />
Provenzano<br />
Bermuda; Jamaica<br />
Pleurocrypta floridana Markham, 1974<br />
Galathea rostrata A. Milne Edwards<br />
Alligator Reef, Florida<br />
Pleurocryptella fimbriata Markham,<br />
1973<br />
Munida constricta A. Milne Edwards<br />
Munida miles A. Milne Edwards<br />
Western Caribbean; Cuba<br />
Probopyria alphei (Richardson, 1900a)<br />
Alpheus armillatus H. Milne Edwards<br />
Alpheus heterochaelis Say<br />
Alpheus normanni Kingsley<br />
Alpheus viridari (Armstrong)<br />
North Carolina to Florida; Antilles;<br />
Brazil; Gulf of Mexico<br />
Parapagurion imbricata Markham, 1978 Probopyrinella latreuticola (Gissler, 1882)<br />
Paguristes tortugae Schmitt<br />
Parapagurus sp.<br />
Cuba; Colombia<br />
Parathelges foliatus Markham, 1972b<br />
Clibanarius vittatus (Bosc)<br />
Pagurus brevidactylus (Stimpson)<br />
Barbados; Curasao; Trinidad<br />
Parathelges occidentalis Markham,<br />
1972b<br />
Clibanarius tricolor (Gibbes)<br />
Iridopagurus sp.<br />
Pylopagurus corallinus (Benedict)<br />
North Carolina; Florida Keys;<br />
Bahamas; Venezuela<br />
Latreutes fucorum (Fabricius)<br />
Bermuda; Sargasso Sea to Azores;<br />
North Carolina to Florida;<br />
Bahamas; Antilles; Gulf of<br />
Mexico<br />
Probopyrus pandalicola (Packard, 1879)<br />
Macrobrachium acanthurus (Wiegmann<br />
Macrobrachium amazonicum (Heller)<br />
Macrobrachium bonelli (Nobili)<br />
Macrobrachium carcinus (Linnaeus)<br />
Macrobrachium faustinum (de Saussure<br />
Macrobrachium ohione (Smith)<br />
Macrobrachium olfersii (Wiegmann)<br />
Macrobrachium surinamicum Holthuis
Palaemon northropi (Rankin)<br />
Palaemon pandaliformis (Stimpson)<br />
Palaemonetes exilipes Stimpson<br />
Palaemonetes intermedins Holthuis<br />
Palaemonetes kadiakensis Rathbun<br />
Palaemonetes paludosus (Gibbes)<br />
Palaemonetes pugio Holthuis<br />
Palaemonetes vulgaris (Say)<br />
Periclimenes americanus (Kingsley)<br />
New Hampshire to Florida;<br />
Caribbean to Brazil; Pacific<br />
Panama<br />
Pseudasymmetrione sp. (see Adkison<br />
and Heard, 1978)<br />
Iridopagurus iris (A. Milne<br />
Edwards)<br />
Venezuela<br />
Pseudione affinis (Sars, 1882)<br />
Pandalus annulicornis Leach<br />
Pandalus bonnieri Caullery<br />
Pandalus leptorhynchus Kinahan<br />
Pandalus montagui Leach<br />
Plesionika antiguai Zariquiey<br />
Plesionika edwardsi (Brandt)<br />
Plesionika ensis (A. Milne Edwards)<br />
Plesionika heterocarpus (Costa)<br />
Plesionika martia (A. Milne<br />
Edwards)<br />
Bermuda; Northeastern Atlantic;<br />
South Africa; Java<br />
Schizobopyrina urocaridis (Richardson,<br />
1904)<br />
Periclimenes longicaudatus (Stimpson)<br />
Pontonia margarita Smith<br />
North Carolina to Florida; Belize;<br />
Gulf of Mexico<br />
Stegias clibanarii Richardson, 1904<br />
Clibanarius tricolor (Gibbes)<br />
Bermuda<br />
Stegophryxus hyptius Thompson, 1902<br />
Iridopagurus sp.<br />
Pagurus annulipes (Stimpson)<br />
Pagurus bonairensis Schmitt<br />
Pagurus brevidactylus (Stimpson)<br />
Pagurus longicarpus Say<br />
Pagurus provenzanoi Forest and de<br />
Saint Laurent<br />
Massachusetts to Florida; Curasao<br />
Synalpheion giardi Coutiere, 1908<br />
Synalpheus longicarpus Herrick<br />
Yucatan, Mexico<br />
Synsynella choprae (Pearse, 1932)<br />
Synalpheus brooksi Coutiere<br />
Synalpheus longicarpus (Herrick)<br />
Synalpheus minus (Say)<br />
Synalpheus pandionis Coutiere<br />
113<br />
Bermuda; North Carolina to Florida;<br />
Bahamas; Haiti; U.S. Virgin<br />
Islands; Gulf of Mexico<br />
Synsynella deformans Hay, 1917<br />
Synalpheus brooksi Coutiere<br />
Synalpheus longicarpus (Herrick)<br />
Synalpheus pectiniger Coutiere<br />
Bermuda; Carolinas to West Indies;<br />
Gulf of Mexico<br />
Urobopyrus processae Richardson,<br />
1904<br />
Ambidexter symmetricus Manning and<br />
Chace<br />
Processa acutirostris Nouvel and<br />
Holthuis<br />
Processa canaliculata Leach<br />
Processa edulis (Risso)<br />
Processa Jimbriata Manning and Chace<br />
Processa tenuipes Manning and Chace<br />
Caribbean; Gulf of Mexico; Brazil;<br />
Mediterranean; Eastern Atlantic
1 14 FLABELLIFERA<br />
reduced to a simple or lobed sac, generally without appendages. The brood-<br />
pouch is formed by invagination of the ventral body wall. The eggs are re<br />
leased by the bursting of the sac. Cryptoniscids have been recorded as<br />
parasites of ostracods, cirripedes, mysidaceans, amphipods, isopods, and<br />
cumaceans. The majority feed on blood, but the females of some forms have<br />
been reported to be egg predators.<br />
Given the highly variable morphology of the epicarideans, and the neces<br />
sity of examining large series of specimens, keys are not provided and species<br />
are not treated individually here. <strong>As</strong> there is a degree of genus- and species-<br />
specificity for the hosts, Table 2 is provided to give a clue to the possible<br />
identity of a specimen. The student is then advised to consult one of the<br />
detailed works on the group. The most useful single work on the speciose<br />
Bopyridae for the area covered here is Markham (1985).<br />
Suborder Flabellifera Sars, 1882<br />
DIAGNOSIS Eyes usually well developed, reduced or absent in cave forms.<br />
Antennules and antennae uniramous; antennal peduncle of five or six arti<br />
cles. Mandible usually strong, adapted for cutting and grinding, occasionally<br />
for piercing; lacinia mobilis, spine-row, and molar usually present, although<br />
latter sometimes reduced; usually with triarticulate palp. Maxilla 1<br />
biramous, sometimes adapted for piercing; maxilla 2 biramous, outer ramus<br />
Key to families of Flabellifera<br />
1. Pleon consisting of four or five free pleonites plus pleotelson 3<br />
Pleon consisting of not more than three free pleonites plus<br />
pleotelson 2<br />
2. Pleon consisting of one or two free pleonites plus pleotelson; body<br />
usually dorsally strongly convex; pleopods subequal<br />
<strong>Sphaeromatidae</strong><br />
Pleon consisting of three free pleonites plus pleotelson; body strongly<br />
depressed; pleopods 1—3 small, natatory, pleopods 4 and 5 large and<br />
broadly ovate Serolidae<br />
3. Uropodal rami flattened, generally not reduced 4<br />
Uropodal rami reduced, exopod often hooklike Limnoriidae
FLABELLIFERA • AEGIDAE 115<br />
4. Pereopods 4—7 prehensile, with dactyli longer than propodi; antennae<br />
reduced, with no clear distinction between peduncle and flagellum<br />
Cymothoidae<br />
Pereopods 4—7 ambulatory, with dactyli shorter than propodi; antennae<br />
normal, peduncle and flagellum clearly distinguished 5<br />
5. Maxilliped bearing distal recurved hooks; pereopods 1-3 strongly<br />
prehensile Aegidae<br />
Maxilliped lacking distal recurved hooks; pereopods 1-3 ambulatory or<br />
at most weakly prehensile 6<br />
6. Maxilliped lacking, or with very reduced endite; maxilla 1 a strongly<br />
falcate hook Corallanidae<br />
Maxilliped with strong endite; maxilla 1 not strongly falcate 7<br />
7. Mandibular incisor distally narrowed, lacinia lacking; maxilla 1 slender<br />
and elongate, with 3-5 distal hooked spines Tridentellidae **<br />
Mandibular incisor distally broad, cusped; maxilla 1 relatively broad,<br />
with several distal spines and setae Cirolanidae {<br />
usually consisting of two lobes. Pereopods generally ambulatory, sometimes<br />
prehensile; pereopods 1 and 2 subchelate only in Serolidae, ancinine<br />
<strong>Sphaeromatidae</strong>, and some Cirolanidae; posterior pereopods sometimes sec<br />
ondarily natatory in some cirolanids. Pleon consisting of as many as five free<br />
pleonites plus pleotelson, but pleonites variously fused in several families.<br />
Five pairs of pleopods usually present, Uropods lateral, usually forming tail-<br />
fan with pleotelson.<br />
REMARKS This suborder contains a large group of diverse families, largely<br />
held together by primitive features such as the tailfan structure. Future work<br />
will undoubtedly show the Flabellifera to be an artificial polyphyletic group.<br />
Family Aegidae Leach, 1815<br />
DIAGNOSIS Dorsal integument usually unornamented. Coxae distinct on<br />
pereonites 2-7. Eyes usually present, large, often almost, or complete con<br />
tiguous. Mandible lacking lacinia mobilis, spine-row, and molar. Maxilla 1<br />
slender, with apical spines. Maxilla 2 with two terminal unequal lobes bear<br />
ing apical spines. Maxillipedal palp of two, three, or five articles. Pereopods<br />
1—3 prehensile, with dactyli strongly curved; pereopods 4—7 ambulatory.
116 FLABELLIFERA • AEGIDAE<br />
Pleopods biramous, bearing plumose marginal setae. Uropods forming tail-<br />
fan with plcotclson. Pleon of four or five free pleonitcs plus pleotelson.<br />
REMARKS Although these large isopods (up to 60 mm) are often referred to<br />
as fish parasites, Brusca (1983) prefers the term "carnivorous scavengers and<br />
micropredators," as they attach to fish hosts infrequently and only long<br />
enough to feed. When feeding, they engorge themselves on the host's blood.<br />
Aegids show almost no host- (or rather prey-) specificity, being opportunistic<br />
feeders, and are most frequently captured by bottom trawls on the ocean bed.<br />
In ovigerous females, the maxillipedal articles become expanded and, along<br />
with the anterior oostegites, cover the buccal field, thereby making feeding<br />
impossible.<br />
Key to genera and subgenera of Aegidae<br />
1. Maxillipedal palp of two or three articles; frontal lamina small, narrow<br />
Rocinela<br />
Maxillipedal palp of five articles; frontal lamina large, broad 2<br />
2. Antennular peduncle articles 1 and 2 expanded; cephalon lacking true<br />
rostrum, not completely separating antennular bases .... Aega (Aega)<br />
Antennular peduncle articles 1 and 2 not expanded; cephalon with true<br />
rostrum completely separating antennular bases . . . Aega (Rhamphion)<br />
Aega Leach, 1815<br />
DIAGNOSIS Eyes large, contiguous or separate. Cephalon with or without<br />
true rostrum. Frontal lamina broad, separating bases of antennae. Mandibu<br />
lar palp article 2 elongate. Maxilla 1 bearing strong apical and subapical<br />
spines. Maxilla 2 of two usually unequal lobes bearing stout spines. Max<br />
illipedal palp of four or five articles, terminal article often small, with setae or<br />
recurved spines; article 4 with stout recurved spines; endite small, seldom<br />
reaching beyond palp article 2. Pleon not much narrower than pereon.<br />
REMARKS Brusca (1983) published a useful account of the genus Aega in the<br />
Eastern Pacific.
1. Eyes contiguous<br />
Eyes separate .<br />
Key to species of Aega (Aega)<br />
Key to species of Aega (Rhamphion)<br />
Aega (A eg a) ecarinata 117<br />
1. Posterior margin of pleotelson distinctly dentate dentata<br />
Posterior margin of pleotelson at most faintly crenulate tenuipes<br />
Aega (Aega) deshaysiana (PL Milne Edwards, 1840)<br />
Figure 51A<br />
DIAGNOSIS 9 18.0 mm. Eyes large, contiguous. Cephalon with frontal mar<br />
gin acute to subacute. Frontal lamina large, shield shaped. Antennular ped<br />
uncle articles 1 and 2 not expanded; flagellum of more than 15 articles.<br />
Uropodal endopod with deep notch in lateral (outer) margin. Pleotelson with<br />
basal width subequal to middorsal length, triangular, lateral margins faintly<br />
to markedly convex, tapering to narrowly rounded to subacute apex.<br />
RECORDS Cuba; Yucatan Peninsula, Mexico; Gulf of Mexico.<br />
Azores; Cape Verde Islands; Tristan da Cunha; Mediterranean; St. Paul and<br />
Amsterdam Islands; Seychelles; east coast of South Africa; Philippines; Ja<br />
pan; Hawaii; northeast Australia; Tasmania; Cocos Islands; Costa Rica.<br />
REMARKS This species is more familiarly known in the Caribbean region as<br />
Aega antillensis Schioedte and Meinert, 1879.<br />
Aega (Aega) ecarinata Richardson, 1898<br />
Figure 51B<br />
deshaysiana<br />
ecarinata<br />
DIAGNOSIS 9 21.0 mm. Eyes well separated. Articles 1 and 2 of antennular<br />
peduncle expanded. Propodus of pereopod 3 with posterodistal lobe. Uropo
118 FLABELLIFERA • AEGIDAE<br />
Figure 51. A, Aega (Aega) deshayesiana; B, Aega (Aega) ecarinata; Cy Aega (Rhamphion)<br />
dentata; Dy Aega (Rhamphion) tenuipes.<br />
dal exopod narrower than endopod; latter distally truncate, lacking marginal<br />
notch. Pleotelson dorsally smooth, with posterior margin broadly trilobed.<br />
RECORDS Bahamas, 776 m; Puerto Rico; Gulf of Mexico, 176 m.
Aega (Rhamphion) dentata Schioedte and Meinert, 1879<br />
Figure 51C<br />
Rocinela 119<br />
DIAGNOSIS 9 7.5 mm. Eyes large, just contiguous in midline. Frontal lam<br />
ina distally acute. Uropodal exopod shorter than and half width of endopod;<br />
latter with lateral margin entire. Pleotelson with two obscure dorsal depres<br />
sions anteriorly; posterior margin crenulate with seven teeth.<br />
RECORDS Cuba.<br />
Aega (Rhamphion) tenuipes Schioedte and Meinert, 1879<br />
Figure 51D<br />
DIAGNOSIS 9 11.5 mm. Eyes large, contiguous. Frontal lamina distally<br />
broadly rounded. Uropodal exopod shorter and narrower than endopod; lat<br />
ter with entire lateral margin. Pleotelson dorsally smooth; posterior margin<br />
evenly and broadly convex, obscurely crenulate.<br />
RECORDS Cuba.<br />
Rocinela Leach, 1818<br />
DIAGNOSIS Cephalon with short rostrum sometimes covering antennular<br />
bases. Eyes well developed. Frontal lamina small, often indistinct. Mandibu<br />
lar palp of three articles, article 1 elongate. Maxillipedal palp of two or three<br />
articles. Pereopods 1—3 usually with spine-bearing expanded lobe on pos<br />
terior margin of propodi.<br />
Key to species of Rocinela<br />
1. Eyes contiguous oculata<br />
Eyes not contiguous 2<br />
2. Cephalon produced anteriorly into broadly rounded rostrum . . cubensis<br />
Cephalon lacking obvious broadly rounded rostrum 3<br />
3. Eyes well separated, cephalon anteriorly broadly triangular .... signata<br />
Eyes barely separate, cephalon anteriorly narrowly triangular insularis
120 FLABELLIFERA • AEG1DAE<br />
Rocinela cubensis Richardson, 1898<br />
Figure 52A<br />
DIAGNOSIS 6 16 mm. Cephalon with two small tubercles between well-<br />
separated eyes; rostrum broadly rounded, extending anteriorly, very ob<br />
vious. Flagellum of antenna with about 15 articles. Propodi of pereopods 1-3<br />
with two spines. Pleotelson basally wider than middorsal length, lateral mar<br />
gins convex, apex rounded.<br />
RECORDS Off Cuba, 290 m.<br />
Rocinela insularis Schioedte and Meinert, 1879<br />
Figure 52B<br />
DIAGNOSIS Ovigerous 9 24.5 mm. Eyes medially barely separated but not<br />
contiguous. Flagellum of antenna of more than 12 articles. Propodus of per<br />
eopods 1-3 with two to four spines on posterior lobe. Uropodal endopod<br />
barely reaching beyond pleotelsonic apex. Pleotelson basally slightly wider<br />
than middorsal length, lateral margins convex, apex rounded.<br />
RECORDS Florida Keys; West Indies; between Mississippi delta and west<br />
coast of Florida, Gulf of Mexico; 550 m.<br />
Rocinela oculata Harger, 1883<br />
Figure 52C<br />
DIAGNOSIS 9 21.0 mm. Eyes contiguous. Cephalon with rostrum truncate<br />
in dorsal view. Antennal flagellum with 12 articles. Propodi of pereopods 1-3<br />
with six to eight spines on lobed posterior margin. Pleotelson with basal<br />
width subequal to middorsal length; posterior margin broadly rounded.<br />
RECORDS Off Georgia; Gulf Stream off Florida, 360-400 m; Puerto Rico,<br />
84 m; Gulf of Mexico, 380-750 m.<br />
New South Wales, Queensland, Australia, 450-630 m.<br />
Rocinela signata Schioedte and Meinert, 1879<br />
Figure 52D<br />
DIAGNOSIS Ovigerous 9 13.0-15.0 mm. Cephalon anteriorly broadly tri<br />
angular, produced over bases of antennules. Eyes widely separate. Flagellum<br />
of antenna with 10 or 11 articles. Pereopods 1—3, propodi unarmed or with
c<br />
L<br />
r<br />
/ /<br />
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- • *<br />
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• '<br />
1 ' * 1<br />
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^fc ^bP^I<br />
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Figure 52. ^4, Rocinela cubensis; B, Rocinela insularis; C, Rocinela oculata; D, Rocinela<br />
signata.<br />
D<br />
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fc?l<br />
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122 FLABELLIFERA • CIROLANIDAE<br />
single spine on posterior margin. Plcotelson with posterior margin evenly<br />
and broadly rounded; usually with inverted W-shaped band of pigment.<br />
RECORDS Florida Keys, shallow infratidal-4 m; Tortugas, from gills ofjew-<br />
fish Epinephelus itajara, mutton snapper Lutjanus analis; U.S. Virgin Islands,<br />
on mutton snapper Lutjanus analis, on yellowfin grouper Mycteroperca venenosa;<br />
Bahamas, on sheepshead Archosargus probatocephalus, on mutton snapper Lut<br />
janus analis, on blackfin snapper Lutjanus buccanella, on queen triggerfish Bat<br />
istes vetula, on saucereye porgy Calamus calamus; Jamaica, on French grunt<br />
Haemulon Jlavolineatum, hogfish Lachnolaimus maximus, on parrotfish Sparisoma<br />
viride; Haiti; Yucatan Peninsula, Mexico, 60—93 m, on gills of tiger shark<br />
Galeocerdo cuvieri; Puerto Rico, in gill slits of southern stingray Dasyatis amer-<br />
icana, in gill slits of nurse shark Ginglymostoma cirratum; Jamaica; Carrie Bow<br />
Cay and Blue Ground Range, Belize, 0.5—2 m, on jolthead porgy Calamus<br />
bajonado and sheepshead porgy Calamus penna, on peacock flounder Bothus lu-<br />
natus, on queen triggerfish Balistes vetula, on Caranx sp., on barracuda<br />
Sphyraena barracuda, on hogfish Lachnolaimus maximus, on mutton snapper Lut<br />
janus analis; Venezuela, on Orthopristis ruber, on Haemulon steindachneri; Sur<br />
inam, on gills of sheepshead porgy Calamus penna; Gulf of Mexico off Florida,<br />
shallow infratidal-55 m, on red grouper Epinephelus morio, on Lutjanus black-<br />
fordi, on black grouper Mycteroperca bonaci, on clearnose skate Raja eglanteria.<br />
Pacific records: Southern California and Gulf of California; Socorro Is<br />
land; Panama; Costa Rica.<br />
REMARKS While often taken from fish hosts (sometimes in the gill cham<br />
ber), this species is equally frequently found freeliving in shallow water over<br />
sand and coral rubble. The species will also attach itself to humans, inflicting<br />
a sharp bite as it tries to feed.<br />
Family Cirolanidae Dana, 1852<br />
DIAGNOSIS Eyes when present, relatively small, lateral. Frontal lamina<br />
present. Mandible with tridentate incisor, lacinia mobilis, blade- or sawlike<br />
molar, and palp. Maxillipedal palp of five articles, endite present. Coxal<br />
plates present on pereonites 2-7, distinctly separated by suture from tergite.<br />
Pereopods generally ambulatory, although anterior three pairs prehensile in<br />
some genera, and posterior four pairs natatory in some genera. Picon of five<br />
free pleonites plus pleotelson in most genera; pleonite 5 with free lateral mar<br />
gins or overlapped by pleonite 4. Pleopods membranous, lacking ridges or<br />
folds. Uropods situated at anterolateral angles of pleotelson, freely articulat<br />
ing, both rami well developed, mobile.
FLABELLIFERA • CIROLANIDAE 123<br />
REMARKS Of the many recent publications on the cirolanids, the most com<br />
prehensive is that of Bruce (1986) on the cirolanids of Australia. Botosa-<br />
neanu, Bruce, and Notenboom (1986) tabulate all the known troglobitic<br />
cirolanids of the world.<br />
Key to subfamilies of Cirolanidae<br />
1. Clypeus projecting; pleonite 5 with free lateral margins (except in<br />
Xylolana) Eurydicinae<br />
Clypeus flattened, not projecting; pleonite 5 lacking free lateral margin,<br />
overlapped by pleonite 4 2<br />
2. Pereopods 1-3 with ischium and merus not anterodistally produced;<br />
antennal peduncular articles 4 and 5 subequal; secondary unguis<br />
present on pereopodal dactyli Cirolaninae<br />
Pereopods 1—3 with ischium and merus anterodistally produced;<br />
antennal peducular articles 3 and 4 subequal; no secondary unguis<br />
on pereopodal dactyli Conilerinae<br />
Subfamily Cirolaninae Dana, 1852<br />
DIAGNOSIS Frontal lamina short, flat. Clypeus flattened, not projecting.<br />
Antennal peduncular articles 4 and 5 subequal, longer than articles 1-3.<br />
Pereopods with secondary unguis on dactyli. Penes reduced or absent.<br />
Pleonite 5 always overlapped by pleonite 4. Pleopod 2 in
124 FLABELLIFERA • CIROLANIDAE<br />
Key to genera of Cirolaninae (Continued)<br />
4. Merus of pereopod 1 posterodistally produced; merus of pereopods 2<br />
and 3 anterodistally produced Bahalana<br />
Meri of pereopods 1—3 not markedly produced 5<br />
5. Animal able to conglobate Creaseriella<br />
Animal unable to conglobate 6<br />
6. Mandibular palp directed anteriorly Anopsilana<br />
Mandibular palp directed posteriorly Haptolana<br />
7. Pleopod 1, exopod longer and broader than endopod Oncilorpheus<br />
Pleopod 1, endopod longer and broader than exopod Calyptolana<br />
Anopsilana Paulian and Delamare Deboutteville, 1956<br />
DIAGNOSIS Body unable to conglobate. Eyes present or absent. Frontal<br />
lamina well developed, as long as broad, or longer than broad, anteriorly<br />
Key to species of Anopsilana<br />
1. Estuarine-brackish water species; integument pigmented when alive 2<br />
Cave species; lacking integumental pigment 3<br />
2. Frontal lamina distally rounded, projecting browni<br />
Frontal lamina distally acute, not projecting jonesi<br />
3. Posterior margin of pleotelson with 10 or more spines 4<br />
Posterior margin of pleotelson with less than 10 spines 5<br />
4. Posterior margin of pleotelson with 10 spines; found in cave on Cuba<br />
Posterior margin of pleotelson with 10—12 spines; found in cave on<br />
Haiti<br />
5. Posterior margin of pleotelson with eight spines; found in cave on<br />
cubensis<br />
acanthura<br />
Grand Cayman crenata<br />
Posterior margin of pleotelson with four spines; found in cave on Haiti<br />
radicicola
Anopsilana crenata 125<br />
somewhat expanded. Antennular peduncle of two articles. Maxillipedal en-<br />
dite with two coupling hooks. Pereopod 1 prehensile, pereopods 2-3 weakly<br />
prehensile, pereopods 4-7 ambulatory. Pleopod 2
126 FLABELLIFERA • GIROLANIDAE<br />
Figure 53. Anopsilana acanthura: A, 9; B, anterior cephalon. Anopsilana browni: C, 9;<br />
D, anterior cephalon. Anopsilana crenata: E, 9; F, anterior cephalon. Anopsilana<br />
cubensis: G, 9; H, anterior cephalon.<br />
Anopsilana cubensis (Hay, 1903)<br />
Figure 53G,H<br />
DIAGNOSIS 9 7.0 mm. Lacking eyes and integumental pigmentation.<br />
Frontal lamina longer than wide, anteriorly expanded, rounded. Posterior<br />
margin of pleotelson with 10 spines.
Bahalana 127<br />
RECORDS Caves in provinces of Pinar del Rio, La Habana, Matanzas, and<br />
on Isla de Pinos, Cuba.<br />
Anopsilana jonesi Kensley, 1987<br />
Figure 54A-D<br />
DIAGNOSIS 6 7.2 mm, ovigerous 9 5.9 mm. Eyes well developed and pig<br />
mented. Dorsal integument strongly pigmented with almost solid central<br />
area on pereonites 1-7. 6 cephalon with three low tubercles near posterior<br />
margin; pereonite 1 with four to six low tubercles. 9 lacking tubercles on<br />
cephalon or pereonite 1. Pereonites 2—7 with low submedian longitudinal<br />
ridges near posterior margin. Frontal lamina narrow, pentagonal, anteriorly<br />
acute, not projecting. Posterior margin of pleotelson with 9 or 10 spines.<br />
RECORDS Salt Creek, and Sittee River, Stann Creek District, Belize, in es-<br />
tuarine mangroves.<br />
Anopsilana radicicola (Notenboom, 1981)<br />
Figure 54E,F<br />
DIAGNOSIS 6 6.3 mm, 9 6.5 mm. Lacking eyes and integumental pigmen<br />
tation. Frontal lamina longer than wide, anteriorly expanded and rounded.<br />
Posterior margin of pleotelson with four spines.<br />
RECORDS Source Debarasse, a spring near Jeremie, Haiti.<br />
Bahalana Carpenter, 1981<br />
DIAGNOSIS Eyes lacking. Frontal lamina basally triangular, anteriorly nar<br />
rowed into poorly developed carina. Pereopods 1—3 prehensile, dactyli and<br />
propodi relatively elongate; pereopod 1 carpus small, almost concealed;<br />
merus with strong posterodistal extension almost reaching dactylar base and<br />
armed with spines. Pereopods 2 and 3, meri with elongate anterodistal exten<br />
sion, meri and carpi with shorter posterodistal extensions. Pereopods 4-7<br />
slender, ambulatory. Pleopod 2 in 6 with copulatory stylet articulating<br />
basally on endopod. Pleopods 3-5, exopods biarticulate, endopods with few<br />
distal marginal setae, or lacking setae. Pleonite 5 with free lateral margin,<br />
hardly overlapped by pleonite 4.
128 FLABELLIFERA • CIROLANIDAE<br />
Key to species of Bahalana<br />
1. Pleopods 3—4, endopods lacking setae; maxillipedal endite with one<br />
coupling hook cardiopus<br />
Pleopods 3—4, endopods with few setae; maxillipedal endite with one or<br />
two coupling hooks 2<br />
2. Antennular peduncle, article 3 longest; maxillipedal endite with two<br />
coupling hooks geracei<br />
Antennular peduncle, article 2 longest; maxillipedal endite with one<br />
coupling hook mayana<br />
Bahalana cardiopus Notenboom, 1981<br />
Figure 55A,B<br />
DIAGNOSIS 9 10.0 mm. Maxillipedal endite with single coupling hook. Per-<br />
eopod 1, meral projection bearing five distal spines. Pleopods 3—5, endopods<br />
lacking marginal setae. Uropodal exopod half width of endopod, four spines<br />
on outer margin.<br />
RECORDS Mount Misery Cave, Mayaguana Island, Bahamas.<br />
Bahalana geracei Carpenter, 1981<br />
Figure 55C-G<br />
DIAGNOSIS 9 15.0 mm, 6 8 mm. Maxillipedal endite with two coupling<br />
hooks. Pereopod I, meral projection bearing seven distal spines. Pleopods 3-<br />
5, endopods with 9—13 distal marginal setae; pleopod 5 endopod with four<br />
distal setae. Uropodal endopod bearing few spines at outer distal margin,<br />
margin not serrate, lacking distinct apex; exopod half width of endopod, with<br />
four spines on outer margin.<br />
RECORDS Lighthouse Cave, San Salvador Island, Bahamas.<br />
Bahalana mayana Bowman, 1987<br />
Figure 55H<br />
DIAGNOSIS 9 8.4 mm, 6 10.0 mm. Clypeus acutely pointed. Antennular<br />
peduncle, article 2 longest. Maxillipedal endite with one coupling hook. Per-
B<br />
Bathynomus 129<br />
Figure 54. Anopsilana jonesi: A> 8\ By anterior cephalon; C, pleopod 2, 8; D,<br />
pleopod 3. Anopsilana radicicola: E> 8\ F> anterior cephalon.<br />
eopod 1, meral projection rudimentary. Pleopods 3—4, endopods with few<br />
marginal setae, pleopod 5 lacking marginal setae. Uropodal exopod narrow,<br />
V3 width of endopod; endopod with distal margin slightly concave.<br />
RECORDS Anchialine caves on Cozumel Island and at Tulum, Yucatan<br />
Peninsula, Mexico.<br />
Bathynomus A. Milne Edwards, 1879<br />
DIAGNOSIS Frontal lamina triangular; clypeus projecting anteriorly. Anten-<br />
nal peduncle with articles 3 and 4 subequal, article 5 longest. Maxillipedal<br />
endite with four to seven coupling hooks. Pereopods 1—3 with anterodistal<br />
margins of ischia and meri produced. Pleopods with all rami bearing margi<br />
nal setae; endopods bearing accessory gills at bases. Posterior margin of<br />
pleotelson dentate.
Figure 55. Bahalana cardiopus: A, 9\ B, pleopod 3. Bahalana geracei: C, 9; D,<br />
pleopod 3; E, pereopod 1; F, pereopod 2; G, anterior cephalon. Bahalana mayana:<br />
H, 9.
Figure 56. Bathynomus giganteus<br />
Bathynomus giganteus A. Milne Edwards, 1879<br />
Figure 56<br />
Calyptolana 131<br />
D I A G N O S I S U p t o 2 8 0 m m . L a r g e p i g m e n t e d e y e s p r e s e n t , n o t v i s i b l e d o r -<br />
s a l l y . A n t e n n u l e w i t h s m a l l e x o p o d d i s t a l l y o n p e d u n c u l a r a r t i c l e 3 . P l e o p o d s<br />
w i t h m a r g i n a l s e t a e o n a l l r a m i ; p l e o n i t e s 3 a n d 4 w i t h e p i m e r a p r o d u c e d<br />
p o s t e r i o r l y . P o s t e r i o r m a r g i n o f p l e o t e l s o n w i t h m e d i a n t o o t h a n d f i v e o r s i x<br />
t e e t h o n e a c h s i d e .<br />
R E C O R D S G u l f o f M e x i c o ; C a r i b b e a n ; B a h a m a s ; F l o r i d a K e y s ; 3 6 0 - 2 3 0 0<br />
m .<br />
R E M A R K S G u t - c o n t e n t a n a l y s i s o f t h e s e d e e p - w a t e r g i a n t s h a s s h o w n t h e m<br />
t o b e s c a v e n g e r s , c o m m o n l y f e e d i n g o n d e a d f i s h , c e p h a l o p o d s , c r a b s , a n d<br />
p o l y c h a e t e w o r m s .<br />
Calyptolana Bruce, 1985<br />
D I A G N O S I S A l l p e r e o p o d s a m b u l a t o r y ; e a c h d a c t y l u s w i t h s e c o n d a r y u n <br />
g u i s . P l e o p o d 1 o p e r c u l a t e , l o n g e r t h a n f o l l o w i n g p l e o p o d s . A l l p l e o p o d a l<br />
r a m i e x c e p t e n d o p o d o f p l e o p o d 5 w i t h m a r g i n a l s e t a e .
132 FLABELLIFERA • C1ROLANIDAE<br />
Calyptolana hancocki Bruce, 1985<br />
Figure 57<br />
DIAGNOSIS 9 3.0 mm. Body dorsally strongly convex. Cephalon with small<br />
rostrum curving ventrally to meet frontal lamina; latter pentagonal. Eyes<br />
small, well pigmented. Coxae of pereonites barely produced. Pleonite 5 lack<br />
ing free lateral margins. Pleotelson with broadly rounded posterior margin.<br />
Uropodal sympod produced along mesial margin of endopod; exopod<br />
slightly less than half length of endopod; latter distally rounded.<br />
RECORDS Dominican Republic; Aruba Island, Netherlands Antilles, 43.2<br />
m.<br />
Cirolana Leach, 1818<br />
DIAGNOSIS Frontal lamina usually twice as long as wide, not projecting;<br />
clypeus flat. Mandible with strong incisor, dentate molar, palp of three arti<br />
cles. Pereopods all ambulatory. Pleon consisting of five free pleonites plus<br />
pleotelson, pleonite 5 overlapped laterally by pleonite 4. All pleopodal rami<br />
Key to species of Cirolana<br />
1. Pleotelson posteriorly very broad to subtruncate 2<br />
Pleotelson posteriorly narrowed 4<br />
2. Uropodal endopod broad, distally rounded, lacking marginal spines<br />
Uropodal endopod broad, distal margin having apical tooth or<br />
obtruncata<br />
angle 3<br />
3. Posterior margin of pleotelson bearing spines; uropodal endopod not<br />
dentate minuta<br />
Posterior margin of pleotelson faintly crenulate, lacking spines;<br />
uropodal endopod strongly dentate crenulitelson<br />
4. Uropodal endopod evenly tapering to acute apex; uropodal exopod<br />
length about four times greatest width albidoida<br />
Uropodal endopod, outer margin convex; uropodal exopod length less<br />
than four times greatest width parva
Figure 57. Calyptolana hancocki: A} 9; B, ventral pleon.<br />
Cirolana crenulitelson 133<br />
bearing marginal setae except endopod of pleopod 5. Copulatory stylet on<br />
endopod of 6 pleopod 2 inserted proximally.<br />
Cirolana albidoida Kensley and Schotte, 1987<br />
Figure 58A-C<br />
DIAGNOSIS 6 7.8 mm. Integument sparsely pitted. Antenna reaching pos<br />
teriorly to pereonite 3. Uropodal endopod triangular, evenly tapering, mar<br />
gins serrate; exopod length about four times greatest width, apically acute.<br />
Pleopod 1 6, endopod markedly narrowed in distal half. Pleopod 2 6, copu<br />
latory stylet reaching beyond rami by about half its length. Pleotelson with<br />
sides gently convex, tapering to rounded posterior margin bearing eight<br />
spines and three or four small serrations anterior to first spine.<br />
RECORDS OffLucaya, Grand Bahama, 180-220 m.<br />
Cirolana crenulitelson Kensley and Schotte, 1987<br />
Figure 58D-F<br />
DIAGNOSIS 6 7.0 mm, 9 7.0 mm. Antenna reaching posteriorly to anterior<br />
of pereonite 3. Pleopod 2 8, copulatory stylet reaching by Vs its length be-
134 FLABELLIFERA • CIROLANIDAE<br />
Figure 58. Cirolana albidoida: A> uropod; C, pleotelsonic apex. Cirolana<br />
crenulitelson: D, 9; Ey pleotelsonic apex; Fy uropod. Cirolana minuta: G, 6; H,<br />
anterior cephalon; /, uropod.<br />
yond rami. Uropodal endopod with mesial margin broadly convex, serrate,<br />
apicaily acute; exopod about 2.5 times longer than wide, mesial margin<br />
weakly convex, apicaily acute. Posterior margin of pleotelson truncate,<br />
faintly crenulate, lacking spines.<br />
RECORDS Carrie Bow Cay, Belize, 36 m.
Cirolana minuta Hansen, 1890<br />
Figure 58G-I<br />
Cirolana parva 135<br />
DIAGNOSIS 8 8.9 mm, 9 5.0 mm. Antenna reaching posteriorly to per-<br />
eonite 3. Pleopod 2 copulatory stylet just reaching to distal mar<br />
gin of rami. Uropodal endopod distally broadly rounded, margin with<br />
rounded teeth; exopod 2.5 times longer than wide, margin with rounded<br />
teeth, apically obscurely subacute. Posterior margin of pleotelson subtrun<br />
cate, with about eight spines.<br />
RECORDS Jamaica; Puerto Rico; Cozumel, Mexico; Gulf of Mexico.<br />
Cirolana parva Hansen, 1890<br />
Figures 59C-E, 60<br />
DIAGNOSIS 6 6.9 mm, 9 7.9 mm. Cephalon with furrow between eyes hav<br />
ing middorsal posterior deflection. Antenna reaching posteriorly to pereonite<br />
4. Pleopod 1 (5, endopod narrowed in distal third. Pleopod 2 6, copulatory<br />
stylet reaching by 'A its length beyond rami. Uropodal endopod with mesial<br />
margin convex, apically acute; exopod 2.5 times longer than wide, apically<br />
acute. Pleotelson evenly tapering to angled posterior margin, with seven or<br />
eight spines.<br />
RECORDS North and South Carolina; Turks and Caicos Islands; St.<br />
Thomas and St. Croix, U.S. Virgin Islands; Andros Island, Bahamas; Pu<br />
erto Rico; Jamaica; Florida Keys; Dry Tortugas; Barbados; Carrie Bow Cay,<br />
Belize; Cozumel, Mexico; Panama; Gulf of Mexico; intertidal to 55 m.
Figure 59. Cirolana obtruncata: A, $; B, uropod. Cirolana parva: C, 9; D, uropod; E,<br />
pleotelson and uropods.
Figure 60. Cirolana parva,<br />
anteroventral cephalon.<br />
C r e a s e r i e l l a R i o j a , 1 9 5 3<br />
Haptolana 137<br />
D I A G N O S I S A n t e n n u l a r p e d u n c l e o f t w o a r t i c l e s ( a r t i c l e s a n d f u s e d ) .<br />
A n t e n n a l p e d u n c l e o f f i v e a r t i c l e s . P e r e o p o d s a l l a m b u l a t o r y . P e n i a l r a m i<br />
f u s e d t o f o r m s h o r t s t o u t p r o c e s s . P l e o p o d 2 i n cT w i t h c o p u l a t o r y s t y l e t i n s e r -<br />
t e d a t b a s e o f e n d o p o d . P l e o p o d s<br />
P l e o n i t e 5 l a c k i n g f r e e l a t e r a l m a r g i n s .<br />
Creaseriella anops (Creaser, 1936)<br />
Figure 61A,B<br />
, e n d o p o d s l a c k i n g m a r g i n a l s e t a e .<br />
DIAGNOSIS 1 5 . 5 m m . A n i m a l a b l e t o c o n g l o b a t e . E y e s l a c k i n g . F r o n t a l<br />
l a m i n a p e n t a g o n a l , l o n g e r t h a n w i d e , w i t h t r a n s v e r s e r i d g e a t w i d e s t p o i n t .<br />
M a x i l l i p e d a l e n d i t e w i t h f o u r o r five c o u p l i n g h o o k s . P l e o t e l s o n w i d e r t h a n<br />
l o n g , p o s t e r i o r m a r g i n v e r y b r o a d l y r o u n d e d a n d f i n e l y c r e n u l a t e . B o t h<br />
u r o p o d a l r a m i d i s t a l l y r o u n d e d , m a r g i n s b e a r i n g s p i n e s a n d s e t a e , s y m p o d<br />
p r o d u c e d a l o n g m e s i a l m a r g i n o f e n d o p o d .<br />
RECORDS S e v e r a l c a v e s a n d c e n o t e s o n t h e Y u c a t a n P e n i n s u l a , M e x i c o .<br />
H a p t o l a n a B o w m a n , 1 9 6 6<br />
D I A G N O S I S M a n d i b u l a r p a l p d i r e c t e d p o s t e r i o r l y . A n t e n n u l a r p e d u n c l e o f<br />
t w o a r t i c l e s , b a s a l a r t i c l e e x p a n d e d . P e r e o p o d s a l l p r e h e n s i l e , w i t h d a c t y l i<br />
c l o s i n g i n p r o p o d a l g r o o v e . P l e o p o d s 3 - 5 , e x o p o d s w i t h p a r t i a l s u t u r e a n d<br />
m a r g i n a l s e t a e ; e n d o p o d s u n d i v i d e d , l a c k i n g m a r g i n a l s e t a e . P l e o n i t e 4 o v e r <br />
l a p p i n g p l e o n i t e 5 l a t e r a l l y .
138 FLABELLIFERA • CIROLANIDAE<br />
Figure 61. Creaseriella anops: A, adult; B, frontal lamina. Haptolana trichostoma: C,<br />
adult; D, frontal lamina. Oncilorpheus stebbingi: £, juvenile (from Paul and Menzies,<br />
1971).<br />
Haptolana trichostoma Bowman, 1966<br />
Figure 61C,D<br />
DIAGNOSIS 6 13.8 mm. Eyes lacking. Frontal lamina broad, pentagonal,<br />
with anterior angle very broad. Pleotelson wider than long, roughly rec<br />
tangular, posterior margin faintly crenulate, with spine between and setae on<br />
crenulations. Uropodal endopod distally broad, exopod distally narrowly<br />
rounded, 2 fo width of endopod, both rami with marginal setae and spines.<br />
RECORDS Cave in Camaguey Province, Cuba.<br />
REMARKS A second species of Haptolana, H. somata Messana and Chelazzi,<br />
has been described from northern Somalia in Africa.
Oncilorpheus Paul and Menzies, 1971<br />
Natatolana 139<br />
DIAGNOSIS Frontal lamina projecting ventrally. Pleopod 1, exopod indu<br />
rate, opercular; endopod membranous, less than half width of exopod.<br />
Uropodal sympod longer than rami, slightly produced along mesial margin<br />
of endopod; rami inserted subapically.<br />
Oncilorpheus stebbingi Paul and Menzies, 1971<br />
Figure 61E<br />
DIAGNOSIS 9 11.0 mm. Body narrow, about five times longer than wide.<br />
Pleotelson triangular, bearing faint middorsal longitudinal ridge, apex nar<br />
rowly rounded.<br />
RECORDS Off Venezuela, 73 m.<br />
Subfamily Conilerinae, new name<br />
DIAGNOSIS Clypeus flattened. Frontal lamina flat, narrow. Antennal ped<br />
uncular articles 3 and 4 subequal. Pereopods 1-3, ischium and merus pro<br />
duced anterodistally. Pereopods lacking secondary unguis on dactyli. Nata<br />
tory setae present on pereopods 4—7.<br />
REMARKS In a discussion of the Cirolanidae, Botosaneanu, Bruce, and<br />
Notenboom (1986:412) refer to the subfamilies Eurydicinae and Cirolaninae<br />
but place the Conilera group under the heading "Unnamed Subfamily." For<br />
consistency, the Conilera group is here recognized as a subfamily.<br />
Key to genera of Conilerinae<br />
1. Uropodal endopod with distal notch; pereopods 4-7 natatory, ischium,<br />
merus, and carpus flattened Politolana<br />
Uropodal endopod lacking distal notch; pereopods 4-7 with basis<br />
flattened and expanded, bearing natatory setae Natatolana<br />
Natatolana Bruce, 1981<br />
DIAGNOSIS Frontal lamina narrow; clypeus flat. Pereopods 1-3 bearing<br />
long setae. Pleopod 2
140 FLABELLIFERA • CIROLANIDAE<br />
Natatolana gracilis (Hansen, 1890)<br />
Figure 62<br />
DIAGNOSIS 8 8.0 mm. Antennule short, not reaching distal end of antcnnal<br />
peduncle. Antenna reaching posteriorly to pereonite 4. Pleopod 2, copulatory<br />
stylet of endopod cylindrical, bowed, distally rounded. Pleotelson with ob<br />
tuse apex, with slight transverse indentation near anterior margin.<br />
RECORDS Probably St. Thomas, U.S. Virgin Islands; off Sombrero Light,<br />
Florida, 100-120 m.<br />
Northern Brazil, 7-85 m.<br />
REMARKS Hansen (1890) indicated some uncertainty about the exact type<br />
locality, but thought it likely to have been St. Thomas. Koening (1972) re<br />
corded this species from several localities off northern Brazil, but did not<br />
illustrate her material.<br />
Politolana Bruce, 1981<br />
DIAGNOSIS Frontal lamina slender, flattened; clypeus flattened. Antennal<br />
peduncular articles 3—5 subequal. Pereopods 1—3 with ischium and merus<br />
anterodistally produced. Pereopods 4—7 with ischium, merus, and carpus<br />
flattened. 6 pleopod 2 with copulatory stylet inserted basally on endopod.<br />
Endopod of pleopod 5 lacking marginal setae. Pleonite 5 overlapped laterally<br />
by pleonite 4. Uropodal endopod with distal emargination; exopod slender,<br />
elongate; sympod produced along mesial margin of endopod.<br />
Key to species of Politolana<br />
1. Uropodal endopod broad distal to emargination, margin obliquely<br />
truncate; coxae of pereonites 2—6 with impressed line impressa<br />
Uropodal endopod distal to emargination somewhat narrowed, margin<br />
evenly convex; coxae of pereonites lacking impressed line polita<br />
Politolana impressa (Harger, 1883)<br />
Figure 63A,B<br />
DIAGNOSIS 6 and 9 up to 27 mm. Frontal lamina slightly expanded ante<br />
riorly. Coxae of pereonites 2-6 with impressed oblique line. Uropodal endo-
Figure 62. Natatolana gracilis: A, 9; B, frontal lamina; C, pleotelson; D> uropod; E><br />
pereopod 7.
142 FLABELLIFERA • CIROLANIDAE<br />
Figure 63. Politolana impressa: A, 6, lateral view; B, uropod, (setae and spines<br />
omitted). Politolana polita: Cy 6; D, pereopod 1; Ey uropod, setae and spines<br />
omitted; F, pereopod 1.<br />
pod broad distal to emargination, margin subtruncate. Pleotelson posteriorly<br />
broadly rounded.<br />
RECORDS Massachusetts to Palm Beach, Florida, 32-650 m.
Figure 64. Politolana polita,<br />
anteroventral cephalon.<br />
Politolana polita (Stimpson, 1853)<br />
Figures 63C-F, 64<br />
Politolana polita 143<br />
DIAGNOSIS 2 7 . 0 m m , 9 2 9 . 0 m m . A n t e n n u l e b a r e l y r e a c h i n g d i s t a l e n d<br />
o f a n t e n n a l p e d u n c l e . F r o n t a l l a m i n a b a s a l l y s l e n d e r , a n t e r i o r l y e x p a n d e d ,<br />
j u s t v i s i b l e i n d o r s a l v i e w . C o x a e l a c k i n g i m p r e s s e d o b l i q u e l i n e . U r o p o d a l<br />
e n d o p o d d i s t a l t o e m a r g i n a t i o n n a r r o w e d , m a r g i n c o n v e x . P l e o t e l s o n p o s <br />
t e r i o r l y b r o a d l y r o u n d e d .<br />
RECORDS<br />
Mexico.<br />
Bay of Fundy, Canada, to Florida Keys, 2-600 m; Gulf of<br />
Subfamily Eurydicinae Stebbing, 1905<br />
D I A G N O S I S C l y p e u s p r o j e c t i n g . P l e o n i t e 5 w i t h f r e e l a t e r a l m a r g i n s ( e x c e p t<br />
i n X y l o l a n a ) . P e n e s p r o m i n e n t . P l e o p o d 2 o f 6 w i t h c o p u l a t o r y s t y l e t a r t i c <br />
u l a t i n g s u b b a s a l l y , m e d i a l l y , o r s u b a p i c a l l y .<br />
Key to genera of Eurydicinae<br />
Uropodal sympod not produced along mesial margin of endopod<br />
Eurydice<br />
Uropodal sympod produced along mesial margin of endopod<br />
Rostrum prominent, fused with frontal lamina, separating antennal<br />
bases<br />
Rostrum not prominent<br />
{continued)
144 FLABELLIFERA • CIROLANIDAE<br />
Key to genera of Eurydicinae {Continued)<br />
3. Picon of five free plconites plus pleotelson 4<br />
Picon of three free plconites plus pleotelson Colopisthus<br />
4. Endopods of plcopods 3-5 lacking marginal setae, or with no more<br />
than three marginal setae; copulatory stylet of endopod of pleopod 2<br />
in 6 articulating subterminally Arubolana<br />
Marginal setae lacking only on endopod of pleopod 5; copulatory stylet<br />
of pleopod 2 endopod in 8 articulating basally Metacirolana<br />
5. Clypeus conical; uropodal endopod lacking notch in outer distal margin<br />
Xylolana<br />
Clypeus flattened; uropodal endopod with notch in outer distal margin<br />
Arubolana Botosaneanu and Stock, 1979<br />
Excirolana<br />
DIAGNOSIS Animal not able to conglobate. Blind, or with very small eyes.<br />
Anterior margin of frontal lamina broad. Antennular peduncle of three arti<br />
cles. Maxillipedal palp of four articles (articles 2 and 3 fused). Maxilla 2<br />
reduced, endite unarmed, exopod with few marginal setae. Pereopods 1 and<br />
2 and sometimes pereopod 3 prehensile; pereopods 4—7 ambulatory.<br />
Pleopods 1 and 2, rami undivided. Pleopod 2 6 with copulatory stylet artic<br />
ulating subterminally on endopod. Pleopods 3-5, exopods biarticulate; endo<br />
pods lacking marginal setae, or with few setae on endopods of 3 and 4;<br />
pleopod 5 exopod with marginal setae interrupted.<br />
Key to species of Arubolana<br />
1. Eyes present, small parvioculata<br />
Eyes absent 2<br />
2. Pleotelson posteriorly rounded; rostrum not distinct in dorsal view<br />
Pleotelson posteriorly subtruncate; rostrum distinct in dorsal view<br />
aruboides<br />
imula
Arubolana aruboides (Bowman and IlifTe, 1983)<br />
Figure 65A-C<br />
Arubolana parvioculata 145<br />
DIAGNOSIS 6 3.9 mm, 9 4.1 mm. Body about three times longer than wide.<br />
Eyes absent. Antennular peduncle article 3 longer than articles 1 and 2 to<br />
gether. Antenna reaching posteriorly to pereonite 6 or 7. Frontal lamina vis<br />
ible in dorsal view between antennal bases, anteriorly rounded and only<br />
slightly wider than proximally, with distally flared ridge on ventral (exposed)<br />
surface. Pleotelson as long as basal width, evenly narrowing to broadly<br />
rounded posterior margin, latter with few small serrations and setae. Uropo-<br />
dal exopod four times longer than wide; endopod length about twice basal<br />
width, distally obliquely truncate, with elongate setae on inner margin.<br />
RECORDS Church Cave and Wonderland Cave, Bermuda.<br />
Arubolana imula Botosaneanu and Stock, 1979<br />
Figure 65D<br />
DIAGNOSIS 6 and 9 6.25 mm. Body 2.3 times longer than wide. Eyes ab<br />
sent. Antenna reaching posteriorly to pereonite 4 or 5. Rostrum distinct,<br />
anteriorly truncate, separating antennal bases, fused with rectangular frontal<br />
lamina ventrally. Pleotelson basally slightly wider than long, posterior mar<br />
gin broadly rounded to subtruncate, with irregular crenuiations or faint<br />
teeth. Uropodal exopod apically acute, reaching to about midlength of endo<br />
pod; latter distally broad, with slight tooth at distolateral angle.<br />
RECORDS Mangel Cora Tunnel, Aruba.<br />
Arubolana parvioculata Notenboom, 1984<br />
Figure 65E,F<br />
DIAGNOSIS 6 2.8 mm, 9 2.9 mm. Body 3.3 times longer than wide.<br />
Cephalon with tiny pigmented eyes. Antenna reaching posteriorly to per<br />
eonite 5. Frontal lamina projecting, dorsally visible. Pleotelson basally wider<br />
than long, tapering to broadly rounded/subtruncate posterior margin bear<br />
ing about six low teeth. Uropodal exopod distally acute, almost three times<br />
longer than basal width; endopod distally serrate, apically acute.<br />
RECORDS Interstitial water near Discovery Bay, Jamaica.
146 FLABELLIFERA • GIROLANIDAE<br />
Figure 65. Arubolana aruboides: A, 8; B, pereopod 1; C, pereopod 2. Arubolana imula<br />
D, 9. Arubolana parvioculata: Ey 6\ Fy pleopod 2, 8.<br />
Colopislhus Richardson, 1902<br />
DIAGNOSIS Cephalon broader than long, becoming triangular between an-<br />
tennal bases. Pleon consisting of three short free pleonites (often obscured<br />
beneath pereonite 7), plus triangular pleotelson.<br />
D
Colopisthus parvus Richardson, 1902<br />
Figure 66A<br />
Eurydice convex a 147<br />
DIAGNOSIS 9 3.6 mm. Eyes large, well pigmented. Frontal lamina prox-<br />
imally narrow, anteriorly widened to truncate distal margin. Antennules and<br />
antennae short, latter reaching posteriorly to pereonite 1. Pleotelson with<br />
strong middorsal ridge.<br />
RECORDS Bermuda; Puerto Rico, intertidal rocks and algae.<br />
Eurydice Leach, 1815<br />
DIAGNOSIS Antennular peduncle article 2 at right angle to article 1. Anten-<br />
nal peduncle of four articles. Frontal lamina usually slender; clypeus usually<br />
a ventrally directed triangular blade. Maxillipedal endite reduced, lacking<br />
coupling hooks. 6 pleopod 2 with copulatory stylet articulating at mid-<br />
length. Pleopod 5, endopod lacking marginal setae. Pleonite 5 with free lat<br />
eral margins, not overlapped by pleonite 4. Uropodal sympod not produced<br />
along medial margin of endopod.<br />
Key to species of Eurydice<br />
1. Frontal lamina distally truncate to faintly bilobed 2<br />
Frontal lamina lanceolate, distally acute personata<br />
2. Posterior margin of pleotelson between notches rounded, with four<br />
moderate spines convexa<br />
Posterior margin of pleotelson between notches almost straight, with<br />
four very short spines and several elongate setae piperata<br />
Eurydice convexa Richardson, 1900<br />
Figures 66B-E, 67A,B<br />
DIAGNOSIS 6 6.1 mm, 9 6.1 mm. Frontal lamina slender, anteriorly widen<br />
ing slightly and becoming truncate to faintly bilobed. Posterior margin of<br />
pleotelson between lateral notches convex, with four spines and few setae<br />
between serrations; spines between three and four times longer than wide. 6:<br />
Plicate process on antennal flagellar articles about l /s length of article.<br />
Pleopod 2 with copulatory stylet distally blunt, reaching well beyond rami.
148 FLABELLIFERA • CIROLANIDAE<br />
Figure 66. Colopisthus parvus: A> 9. Eurydice convexa: B, 6; C, uropod; Dy antennule;<br />
E, pleotelsonic apex. Eurydice per sonata: F> 8, lateral view; Gy pleotelsonic apex.<br />
Eurydice piperata: Hy pleotelsonic apex.<br />
RECORDS South Carolina to Florida Keys; Bahamas; Gulf of Mexico and<br />
Caribbean.<br />
REMARKS Whether E. convexa and E. littoralis are conspecific needs further<br />
investigation. Differences can be detected in the male plicate organs of the<br />
antennae, and in the mandibular palp spination, but range of variation in
Excirolana 149<br />
these features is still unknown. It seems unlikely that the species recorded as<br />
E. littoralis by Moreira (1972) from Brazil is the same species.<br />
Eury dice per sonata Kensley, 1987a<br />
Figures 66F,G; 67C<br />
DIAGNOSIS 6 6.0 mm, ovigerous 9 5.1—6.4 mm. Frontal lamina slender,<br />
lanceolate, anteriorly acute. Posterior margin between notches faintly con<br />
vex, with four relatively elongate spines (inner pair five or six times longer<br />
than wide) and few setae between dentitions. 6: Plicate organ on antennal<br />
flagellar articles half length of article. Pleopod 2, copulatory stylet on endo-<br />
pod clavate, barely reaching beyond ramus.<br />
RECORDS Bermuda; off Georgia, 18-27 m; off South Carolina, 34 m; off<br />
Miami, Florida; Puerto Rico, 13—17 m; Bahamas, 1—2 m and surface<br />
plankton tow; Haiti; Cuba; Venezuela.<br />
REMARKS This recently discovered species has masqueraded under the<br />
names of E. convexa and E. littoralis for some time, which may explain some of<br />
the inconsistencies in the literature, especially in variation in the pleotelsonic<br />
apex.<br />
Eury dice piper at a Menzies and Frankenberg, 1966<br />
Figure 66H<br />
DIAGNOSIS 6 4.0 mm, 9 4.5 mm. Frontal lamina slender, widening ante<br />
riorly to become slightly bilobed. Posterior margin of pleotelson between<br />
notches straight to faintly convex, with four spines barely twice longer than<br />
wide, and several much longer setae between dentition. 6: Plicate organ on<br />
antennal flagellar articles about l /e length of article but situated subdistally.<br />
Pleopod 2, copulatory stylet clavate, reaching well beyond ramus.<br />
RECORDS Georgia to Florida, Gulf of Mexico, 37-150 m.<br />
Excirolana Richardson, 1912a<br />
DIAGNOSIS Cephalon with prominent rostrum separating antennular bases;<br />
fused with flattened frontal lamina. Clypeus with short, broadly triangular<br />
blade projecting anteroventrally. Antennal peduncle with four or five arti<br />
cles. Maxillipedal endite with single coupling hook. Pleopods 3—5, endopods
150 FLABELLIFERA • GIROLANIDAE<br />
F i g u r e 6 7 . E u r y d i c e c o n v e x a : A , a n t e r o v e n t r a l c e p h a l o n ; B , f r o n t a l l a m i n a . E u r y d i c e<br />
per sonata: C, frontal lamina.<br />
l a c k i n g m a r g i n a l s e t a e . P l e o n i t e 5 w i t h f r e e l a t e r a l m a r g i n s , n o t o v e r l a p p e d<br />
b y p l e o n i t e 4 . U r o p o d a l s y m p o d p r o d u c e d a l o n g m e s i a l m a r g i n o f e n d o p o d .<br />
Key to species of Excirolana<br />
1 . P l e o t e l s o n w i t h t w o a n t e r i o r h o l l o w s c l e a r l y j o i n e d b y i m p r e s s e d l i n e ;<br />
u r o p o d a l e n d o p o d a b o u t h a l f l e n g t h o f e x o p o d braziliensis<br />
P l e o t e l s o n w i t h t w o a n t e r i o r h o l l o w s n o t c o n n e c t e d b y i m p r e s s e d l i n e ;<br />
u r o p o d a l e n d o p o d a b o u t t w o - t h i r d s l e n g t h o f e x o p o d mayana<br />
Excirolana braziliensis Richardson, 1912a<br />
F i g u r e s 6 8 A - C , 6 9 A - C<br />
D I A G N O S I S 6 6 . 0 m m , 9 7 . 5 m m . F r o n t a l l a m i n a v e r y s l e n d e r b e t w e e n<br />
a n t e n n a l b a s e s , w i d e n i n g a n t e r i o r l y i n t o r o u n d e d s t r u c t u r e b e t w e e n a n t e n -
Figure 68. Excirolana braiiliensis: Ay 6; B, pleopod 2 8; C, uropod, (setae and<br />
spines omitted). Excirolana mayana: D, 9; Ey pleopod 2 6\ F, uropod, setae and<br />
spines omitted.
152 FLABELLIFERA • GIROLANIDAE<br />
400HM M'Hrl<br />
F i g u r e 6 9 . Excirolana b r a z i l i e n s i s : A , a n t e r o v e n t r a l c e p h a l o n ; B , p l e o t e l s o n ;<br />
u r o p o d a l e n d o p o d . Excirolana mayana: D , a n t e r o d o r s a l c e p h a l o n ; E , a n t e r c<br />
c e p h a l o n .<br />
n u l a r b a s e s , j o i n e d t o r o s t r u m b y v e r y s l i m i s t h m u s . C l y p e u s d i s t a l l y s u b <br />
a c u t e . U r o p o d a l e n d o p o d a b o u t h a l f l e n g t h o f e x o p o d . P l e o t e l s o n w i t h t w o<br />
l a t e r a l h o l l o w s d e f i n e d a n d c o n n e c t e d b y c l e a r i m p r e s s e d l i n e ; p o s t e r i o r m a r <br />
g i n e v e n l y c o n v e x , b e a r i n g n u m e r o u s p l u m o s e s e t a e .
Metacirolana 153<br />
RECORDS Caribbean to Brazil; common in the intertidal of sandy beaches;<br />
Gulf of Mexico.<br />
Gulf of California to Chile.<br />
REMARKS Glynn et al. (1975) produced a thorough study of the taxonomy,<br />
zonation, and distribution of this Pan-American species.<br />
Excirolana may ana (Ives, 1891)<br />
Figures 68D-F, 69D,E<br />
DIAGNOSIS 6 8.2 mm, 9 10.0 mm. Frontal lamina between antennal bases<br />
about half anterior width. Clypeus anteriorly rounded. Uropodal endopod<br />
about 2 /3 length of exopod. Pleotelson with two faint lateral hollows in ante<br />
rior half, not connected by impressed line.<br />
RECORDS Florida to Venezuela, intertidal.<br />
Metacirolana Nierstrasz, 1931<br />
DIAGNOSIS Frontal lamina anteriorly dilated, free, projecting; clypeus tri<br />
angular, projecting ventrally. Maxillipedal endite with one coupling hook.<br />
Pleon with five free segments, pleonite 5 not overlapped laterally by pleonite<br />
4. Eyes often larger, and antennular flagellum of more articles in 6 than in<br />
2.<br />
Key to species of Metacirolana<br />
1. Telson posteriorly truncate halia<br />
Telson posteriorly rounded or angulate 2<br />
2. Posterior margin of telson an obtuse angle agaricicola<br />
Telson posteriorly rounded 3<br />
3. Posterior margin of telson narrowly rounded; uropodal rami, margins<br />
strongly dentate menziesi<br />
Posterior margin of telson broadly rounded; uropodal rami, margins<br />
obscurely dentate sphaeromiformis
154 FLABELLIFERA • CIROLANIDAE<br />
Metacirolana agaricicola Kensley, 1984<br />
Figure 70A-C<br />
DIAGNOSIS 6 2,6 mm, ovigerous 9 2.1 mm. Antennular flagellum of six or<br />
seven articles. Antennal flagellum of 10 articles. Posterior margin of telson<br />
finely dentate, with broadly obtuse median point. Uropodal exopod about<br />
half width of endopod, margins dentate, apically acute; endopod, margins<br />
dentate, distally angled, apically acute.<br />
RECORDS Carrie Bow Cay, Belize, 1-20 m, in coral on reef slope, and spur<br />
and groove zone.<br />
Metacirolana halia Kensley, 1984<br />
Figure 70D-F<br />
DIAGNOSIS 6 2.9 mm, ovigerous 9 2.7 mm. Antennular flagellum of 10<br />
articles in 9, 14 in 6. Antennal flagellum of 10 articles in 9, 11 in 6. Pos<br />
terior margin of telson truncate, bearing about eight sensory spines. Uropo<br />
dal exopod distally broadly rounded, more than half distal width of endopod,<br />
outer margin dentate, with about 11 sensory spines; endopod distally broad,<br />
margin straight, bearing about 12 sensory spines.<br />
RECORDS Carrie Bow Cay, Glover's Reef, Belize; intertidal to 23 m; Turks<br />
and Caicos Islands, 1 m; Bahamas; Jamaica; Cozumel, Mexico.<br />
Metacirolana menziesi Kensley, 1984<br />
Figure 71A,B<br />
DIAGNOSIS 8 2.3 mm, ovigerous 9 2.4 mm. Antennular flagellum of six<br />
articles in 9, eight in 6. Antennal flagellum of nine articles in 9, 10 in
Figure 70. Metacirolana agaricit<br />
halia: D, uropod; E, 8; F, 9.<br />
rou<br />
pair<br />
Metacirolana sphaeromijormis 155<br />
nded, obscurely dentate. Telson with low rounded middorsal ridge and<br />
of lateral ridges. Uropodal exopod more than half width ofendopod,<br />
margin dentate; exopod distally broadened, margin dentate, with few sen<br />
sory spines
156 FLABELLIFERA • CIROLANIDAE<br />
Figure 71. Metacirolana menziesi: A, 9; B, uropod. Metacirolana sphaeromiformis: C,<br />
D, uropod.<br />
RECORDS Looe Key, Florida, intertidal reef crest; Turks and Caicos Is<br />
lands, 1 m; St. Thomas, U.S. Virgin Islands.<br />
Xylolana Kensley, 1987a<br />
DIAGNOSIS Frontal lamina and rostrum fused, broad, separating antennu-<br />
lar bases. Clypeus conical, projecting. Maxillipedai endite reduced, lacking
A Icirona 15 7<br />
coupling hooks; palp of five articles. Gopulatory stylet in 6 articulating in<br />
distal half of mesial margin of pleopod 2 endopod. Pleopods 3-5, exopods<br />
biarticulate; endopods lacking marginal setae. Pleonite 5 lacking free lateral<br />
margins, overlapped by pleonite 4. Uropodal sympod produced along mesial<br />
margin of endopod.<br />
Xylolana radicicola Kensley, 1987a<br />
Figure 72<br />
DIAGNOSIS 8 2.6 mm, 9 3.3 mm. Body about four times longer than great<br />
est width. Uropodal exopod about 2 fo width of endopod, bearing single short<br />
subapical spine; both uropodal rami distally rounded. Pleotelson with lateral<br />
margins subparallel, with poorly defined middorsal longitudinal ridge.<br />
RECORDS Twin Gays, Belize, in dead red mangrove roots, 1 m.<br />
Family Corallanidae Hansen, 1890<br />
DIAGNOSIS Outer ramus of maxilla 1 apex with single strong falcate spine,<br />
with single strong spine with one or more smaller hooked spines at base, or<br />
with two large recurved spines, occasionally with one to three smaller spines<br />
between them. Maxillipedal endite reduced or lacking.<br />
Key to genera of Corallanidae<br />
1. Maxilla 1 with single strong falcate spine 2<br />
Maxilla 1 with two large falcate spines Alcirona<br />
2. Maxilla 2, distal article slender-elongate; article 2 of maxillipedal palp<br />
longest Nalicora<br />
Maxilla 2 distally bluntly bilobed; article 3 of maxillipedal palp longest<br />
Alcirona Hansen, 1890<br />
Excorallana<br />
DIAGNOSIS Antennular peduncle of two articles. Mandible lacking molar.<br />
Maxilla 1 having two large recurved spines, with one or more smaller spines<br />
between these. Maxilla 2 a simple rounded lobe. Maxilliped lacking endite.<br />
Posterior half of body hirsute.
158 FLABELLIFERA • CORALLANIDAE<br />
Key to species of Alcirona<br />
1. Golden-brown setae starting dorsally on pereonite 3; pereopod 1,<br />
dactylus having several elongate spines insularis<br />
Golden-brown setae starting dorsally on pereonites 5 or 6; pereopod 1,<br />
dactylus having accessory spine only krebsi<br />
Alcirona insularis Hansen, 1890<br />
Figure 73A<br />
DIAGNOSIS 8 and 9, 5.0 mm. Posterior half of body, especially in (J, bear<br />
ing stiff golden-brown setae, these beginning as posterior row on pereonite 3,<br />
and becoming dense on posterior pereonites, pleonites and pleotelson. Per-<br />
eopods 1— 3, dactylus strongly falcate, having distal unguis equal in length to<br />
rest of article, and with three or four strong teeth on posterior margin. Apex<br />
of pleotelson rounded, bearing six short marginal spines in addition to setae.<br />
RECORDS Looe Key, Florida, 0.5-6 m; St. Thomas, U.S. Virgin Islands,<br />
40—46 m; Puerto Rico, from intertidal coral rubble and from gills of nurse<br />
shark Ginglymostoma sirratum; St. Lucia, from coral rubble.<br />
Alcirona krebsii Hansen, 1890<br />
Figure 73B-D<br />
DIAGNOSIS 8 10 mm, ovigerous 9 15.5 mm. Posterior half of body, espe<br />
cially in (J bearing stiff golden-brown setae, these beginning in posterior row<br />
on pereonites 5 or 6, becoming dense on posterior pereonites, pleonites,<br />
uropods, and pleotelson. Pereopod 1, dactylus strongly falcate, with unguis<br />
equal in length to rest of article, and with one strong tooth and several low<br />
tubercles on posterior margin. Apex of pleotelson rounded, bearing six short<br />
spines in addition to setae.<br />
RECORDS Bermuda, in sponges; Florida Keys; Quintana Roo, Yucatan<br />
Peninsula; St. Thomas, U.S. Virgin Islands; Venezuela.<br />
REMARKS A single 8-mm 8 specimen of Alcirona from Panama Bay (Pa<br />
cific) has the characteristic pereopod 1 of A. krebsii, but has the rows of stiff<br />
setae beginning on about pereonite 3. The possibility that A. krebsii is another<br />
amphi-Panamic species needs to be investigated.<br />
*
Excorallana 159<br />
Figure 72. Xylolana radicicola: A, By 6; C, maxilliped; D, pleopod 2 6; E, pleopod 3.<br />
Excorallana Stebbing,1904<br />
DIAGNOSIS Eyes well developed and pigmented, sometimes contiguous or<br />
nearly so. Maxilla 1, outer ramus a single falcate spine. Maxillipedal palp of<br />
five articles; endite reduced or absent. Pereopods 1—3 subprehensile or pre-
160 FLABELLIFERA • CORALLANIDAE<br />
Figure 73. Alcirona insularis: A, pereopod 1. Alcirona krebsi: B3 d\ C, maxilla 1; D,<br />
pereopod 1.<br />
hensile, pereopods 4—7 ambulatory. All rami of pleopods bearing marginal<br />
setae. Pleotelson often with characteristic spination and tubcrculation; lat<br />
eral margins often with incision.<br />
REMARKS Excorallana subtilis (Hansen, 1890) was described from St.<br />
Thomas, U.S. Virgin Islands, based on a specimen undergoing ecdysis; the<br />
true identity of this species remains uncertain.<br />
In the key, two species have been included which are not illustrated. These<br />
are E. mexicana Richardson, 1905, from the Gulf of Mexico, and E. delaneyi
Excorallana 161<br />
Stone and Heard, 1989, from the northeastern Gulf of Mexico. The latter<br />
species, particularly, could conceivably be encountered in the Florida Keys.<br />
Delaney (1984) provides a useful review of the genus Excorallana, and of the<br />
distribution of the species.<br />
Key to species of Excorallana<br />
1. Eyes contiguous 2<br />
Eyes well separated 4<br />
2. Apex of pleotelson with deep slit Jissicauda<br />
Apex of pleotelson entire 3<br />
3. Pleotelson with lateral incision oculata<br />
Pleotelson lacking lateral incision warmingii<br />
4. Pleotelson with lateral incision 6<br />
Pleotelson lacking lateral incision 5<br />
5. Frontal lamina linguiform, anteriorly rounded berbicensis<br />
Frontal lamina posteriorly with faintly concave margins, anteriorly<br />
subacute delaneyi<br />
6. Frontal lamina strongly grooved for entire length 7<br />
Frontal lamina with ventral surface flat 8<br />
7. Pleotelson with medial row of small tubercles flanked by row of larger<br />
tubercles in posterior half mexicana<br />
Pleotelson lacking rows of tubercles in posterior half antillensis<br />
8. Frontal lamina distinctly bell shaped; 8 cephalon with two pairs of<br />
tubercles and antennular bases not tuberculate quadricornis<br />
Frontal lamina, and 6 cephalon and antennules not as above 9<br />
9. Frontal lamina anteriorly broadly rounded, length 1.5 or less times<br />
basal width; 6 cephalon with two pairs of tubercles and basal<br />
antennular article each with tubercle sexticornis<br />
Frontal lamina anteriorly narrowly rounded, length about twice basal<br />
width; 8 cephalon with three tubercles, tubercles lacking on<br />
antennular bases tricornis tricornis
162 FLABELL1FERA • G0RALLAN1DAE<br />
Excorallana antillensis (Hansen, 1890) - t. ar- '
Excorallana oculata 163<br />
Figure 74. Excorallana antillensis: A, 6; B, frontal lamina; C, maxilla 2; D,<br />
maxilliped. Excorallana berbicensis: E, 6\ F, frontal lamina.<br />
Excorallana oculata (Hansen, 1890)<br />
Figure 75A,B<br />
DIAGNOSIS 6 6.9 mm, 9 8.5 mm. Eyes contiguous. Cephalon unorna-<br />
mented. Frontal lamina slender, linguiform, widest posteriorly. Pleonites 3-5<br />
each with slightly hollowed middorsal area containing strong flattened tuber<br />
cle; pleonite 5 with two strong flanking tubercles; pleotelson basally with low<br />
median ridge and two strong submedian tubercles; short strong spines in two<br />
roughly triangular submedian patches; lateral incisions present; apex nar<br />
rowly rounded.<br />
RECORDS Bahamas; Cuba, Puerto Rico, 40 m; Barbados.<br />
Brazil.
J<br />
Figure 75. Excorallana oculata: A, d; B> frontal lamina. Excorallana quadricomis: C,<br />
d; Dy frontal lamina. Excorallana sexticomis: Ey d; F, frontal lamina. Excorallana<br />
tricomis tricomis: G, d; H, frontal lamina. Excorallana warmingi: I, d; J, frontal<br />
lamina.<br />
H
Excorallana quadricornis (Hansen, 1890)<br />
Figures 75C,D; 76A-C<br />
Excorallana tricornis tricornis 165<br />
DIAGNOSIS 6 13.2 mm, 9 12.1 mm. Eyes well separated. Cephalon in 6<br />
with two pairs of tubercles, anterior pair connected by low rounded ridge,<br />
posterior pair situated mesial to eyes. Frontal lamina bell shaped, broadest<br />
posteriorly. Pereonite 1 with submedian pair of low tubercles. Posterior mar<br />
gin of pereonite 7 and pleonites faintly tuberculate. Pleotelson with two sub-<br />
median raised areas bearing short spines; lateral margins with incision; few<br />
low tubercles basally.<br />
RECORDS Bermuda; St. Thomas, U.S. Virgin Islands; Jamaica, intertidal<br />
in grassflats and between mangrove roots; Martinique; Belize; Venezuela.<br />
Excorallana sexticornis (Richardson, 1901)<br />
Figures 75E,F; 76D-F<br />
DIAGNOSIS 6 7.9 mm, ovigerous 9 6.9—8.3 mm. Eyes well separated. Basal<br />
antennular peduncular article in 6 with short anterodorsally directed tuber<br />
cle. Cephalon with two pairs of prominent tubercles, anterior pair shorter<br />
than posterior pair, latter situated mesial to eyes. Frontal lamina, length less<br />
than twice width, sides parallel to faintly converging anteriorly, apically<br />
broadly rounded. Posterior margins of pleonites 2—5 with low rounded tuber<br />
cles, those near middorsal line largest. Pleotelson with two basal submedian<br />
tubercles, numerous scattered dorsal spines, lateral margins with incision,<br />
apex narrowly rounded.<br />
RECORDS Key West, Florida; Cuba; Puerto Rico; Twin Cays, Belize, shal<br />
low infratidal from dead mangrove wood.<br />
Excorallana tricornis tricornis (Hansen, 1890)<br />
Figures 75G,H; 77<br />
DIAGNOSIS 6 8.2 mm, $ 9.9 mm. Basal antennular article narrow, not<br />
dilated. Cephalon in 6 with one median and two dorsal "horns." Eyes large,<br />
well separated. Frontal lamina between two and three times longer than<br />
basal width, sides subparallel, anteriorly rounded to subacute. Pereon<br />
smooth. Margins of lateral incision of pleotelson separated by gap; anterior<br />
margin of incision lined with short spines; scattered short spines on dorsum<br />
of telson, but especially concentrated in two submedian patches. Uropodal<br />
exopod, length 2.3-2.5 times width; apical notch nearly symmetrical.
166 FLABELLIFERA • CORALLANIDAE<br />
Figure 76. Excorallana quadricornis: A, cephalon and pereonite 1 6; B, ventral<br />
cephalon; C, frontal lamina enlarged. Excorallana sexticornis: D, cephalon and<br />
o<br />
pereonite 1 8; E, cephalon 9; F, ventral cephalon.<br />
RECORDS Turks and Caicos Islands, 1 m; St. Thomas and St. Croix, U.S.<br />
Virgin Islands, 48—55 m; Cuba; Puerto Rico, on gills of rays Aetobatus narinari<br />
and Dasyatis americana, and on squirrel fish; Belize, intertidal to 15.2 m, in<br />
tertid bble sediments in Syringodium and Thalassia<br />
g r a s s b e d s , o n b r o w n a l g a T u r b i n a r i a , o n M a d r a c i s s p . s p o n g e , o n A g a r i c i a s p<br />
c o r a l ; G u l f o f M e x i c o .
Excorallana warmingii 167<br />
F i g u r e 7 7 . Excorallana tricornis tricornis: A , c e p h a l o n a n d p e r e o n i t e 1 6 ; B , v e n t r a l<br />
c e p h a l o n ; C, p l e o t e l s o n a n d u r o p o d s .<br />
R E M A R K S T h e s u b s p e c i e s E x c o r a l l a n a t r i c o r n i s o c c i d e n t a l i s R i c h a r d s o n , 1 9 0 5 a ,<br />
f r o m s o u t h e r n C a l i f o r n i a , d i f f e r s f r o m t h e G u l f a n d C a r i b b e a n s u b s p e c i e s i n<br />
l a c k i n g a g a p b e t w e e n t h e m a r g i n s o f t h e p l e o t e l s o n i c i n c i s i o n , a n d i n h a v i n g<br />
a r e l a t i v e l y w i d e r u r o p o d a l e x o p o d w h i c h s h o w s a d i s t i n c t l y a s y m m e t r i c a l<br />
a p i c a l n o t c h .<br />
E x c o r a l l a n a w a r m i n g i i ( H a n s e n , 1 8 9 0 )<br />
F i g u r e 7 5 1 , J<br />
D I A G N O S I S ( J 9 . 7 m m , 9 1 2 . 0 m m . C e p h a l o n u n o r n a m e n t e d . E y e s c o n <br />
t i g u o u s , o c c u p y i n g m o s t o f d o r s a l s u r f a c e o f h e a d . P o s t e r i o r m a r g i n s o f<br />
p l e o n i t e s v e r y f a i n t l y t u b e r c u l a t e . F r o n t a l l a m i n a , l e n g t h s l i g h t l y m o r e t h a n<br />
t w i c e b a s a l w i d t h , t a p e r i n g a n t e r i o r l y t o r o u n d e d a p e x . P l e o t e l s o n u n o r n a <br />
m e n t e d e x c e p t f o r t w o f a i n t s u b m e d i a n t u b e r c l e s b a s a l l y ; l a t e r a l i n c i s i o n s<br />
l a c k i n g ; a p e x b r o a d l y r o u n d e d , w i t h f i v e l o w b u t d i s t i n c t m a r g i n a l t e e t h .<br />
*
168 FLABELLIFERA • CORALLAN1DAE<br />
Figure 78. Nalicora rapax: A, 9; B> maxilla 1; C> maxilla 2; D, maxilliped.<br />
RECORDS Bahamas; between Cuba and the Yucatan Pensinsula; Puerto<br />
Rico.<br />
Off Brazil near Rio de Janeiro.<br />
Nalicora Moore, 1901<br />
DIAGNOSIS Maxilla 1 exopod a single strongly falcate distal spine with<br />
knoblike mesial process, and basal caplike convex papilla-bearing structure.<br />
Maxilla 2 of four articles, distal article slender. Maxillipedal palp of five<br />
articles; endite lacking.
Nalicora rapax Moore, 1901<br />
Figure 78<br />
FLABELL1FERA • CYMOTHOIDAE 169<br />
DIAGNOSIS 6 6.9 mm, ovigerous 9 10.0 mm. Eyes well developed. Frontal<br />
lamina basally slender, widening anteriorly, apex subacute. Posterior half of<br />
body bearing numerous scattered stiff setae. Pereonites 4-7 with row of low<br />
rounded tubercles near posterior margin. Posterior margins of pleonites 3—5<br />
faintly tuberculate, more noticeable in 6. Pleotelson wider than long; lateral<br />
margins faintly sinuous; apex rounded.<br />
RECORDS Florida Keys, 55 m; Puerto Rico, 50-150 m; Gulf of Mexico off<br />
Florida, 37-73 m.<br />
Family Cymothoidae Leach, 1818<br />
DIAGNOSIS Antennules and antennae reduced, no clear distinction between<br />
peduncles and flagella. Mandibular palp of three articles. Maxilla 1 with four<br />
terminal spines. Maxilla 2 apically bilobed, armed with several spines. Max-<br />
illipedal palp of two articles, terminal article bearing hooks. All seven pairs of<br />
pereopods prehensile, ending in strongly hooked dactyli. Pleopods lacking<br />
marginal setae in adults.<br />
REMARKS The cymothoids are exclusively ectoparasites on marine, fresh<br />
water, and brackish-water fishes. Most cymothoids occur in shallow water,<br />
mainly in tropical and subtropical areas. The position of attachment on the<br />
host (externally, in the buccal cavity, or in the gill chamber) is usually genus-<br />
or species-specific. The body of gill parasites is often asymmetrical, being<br />
slightly twisted, perhaps an effect of the position on the host. The mouthparts<br />
are highly adapted for the parasitic mode of life, while all seven pairs of<br />
pereopods are strongly prehensile. The posterior pereopods of some genera<br />
have the basal article expanded and carinate, allowing for increased mus<br />
culature. The secretion of anticoagulants in the juvenile stages further aids<br />
the blood-feeding habit. The surface area of the pleopods is often increased<br />
by the development of lobes on the bases or the lamellae, providing an in<br />
creased respiratory ability.<br />
The post-mancal juvenile stages (sometimes referred to as the aegathoid<br />
stage) have large eyes, and highly setose pleopods for active swimming. The<br />
juveniles will attach themselves indiscriminantly to any convenient fish host,<br />
but eventually attach to the preferred host-species. The juvenile then de<br />
velops into a functional male, losing the swimming setae of the pleopods.<br />
Both juveniles and males feed actively, drawing blood from the host fish. The
170 FLABELLIFERA • GYMOTHOIDAE<br />
Key to genera of Cymothoidae<br />
1. Antennule broader and usually longer than antenna; cephalon very<br />
weakly sunk into pcreonite 1 2<br />
Antennule not broader or longer than antenna; cephalon distinctly<br />
immersed in, or not at all immersed in pereonite 1 4<br />
2. Bases of antennules widely separated 3<br />
Bases of antennules contiguous Glossobius<br />
3. Body curved to one side; pleonitc 1 extended laterally more on one side<br />
than on other Mothocya<br />
Body rarely curved to one side; pleonitc 1 extended equally on each<br />
side Renocila<br />
4. Pereonites and coxal plates 4-7 strongly expanded on one side only<br />
Agarna<br />
No pereonites or coxal plates strongly expanded 5<br />
5. Cephalon not immersed in pereonite 1; posterior margin of cephalon<br />
trisinuate 6<br />
Cephalon to some degree immersed in pereonite 1; posterior margin of<br />
cephalon not trisinuate 7<br />
6. Posterolateral angles of pereonites 2-6 not produced; coxal plates short,<br />
rarely reaching posterior margin of their pereonites Anilocra<br />
Posterolateral angles of pereonites 2-6 posteriorly increasingly<br />
produced; coxal plates usually reaching to posterior margin of their<br />
pereonites Nerocila<br />
7. Basal antennular articles expanded and contiguous Ceratothoa<br />
Basal antennular articles expanded but not contiguous, or basal<br />
antennular articles neither expanded nor contiguous 8<br />
8. Basal antennular articles expanded but not contiguous Kuna<br />
Basal antennular articles neither expanded nor contiguous 9<br />
9. Plconal margins continuous with pereonal margins, pleon not abruptly<br />
narrowed, only weakly immersed in pereonite 7 Lironeca<br />
Pleon to some degree narrower than pereon; pleon usually deeply<br />
immersed in pereonite 7 Cymothoa
FLABELLIFERA • CYMOTIIOIDAE 171<br />
male eventually becomes a female (all cymothoids are protandrous) should a<br />
female not already be present. In some species, the female is nonfeeding. In<br />
those species which settle either in the mouth cavity or gill chamber of the<br />
host, integumental pigment is frequently lost, and the eyes become reduced.<br />
Given the highly variable morphology of the cymothoids, in part imposed<br />
by the parasitic mode of life, and the existence of polymorphism and possible<br />
sibling species, the taxonomy of this family demands the examination of large<br />
numbers of specimens. <strong>As</strong> a further aid to identification, Table 3 is provided,<br />
giving host species, parasite, and site of attachment.<br />
TABLE 3. GYMOTHOID PARASITES FROM THE CARIBBEAN AREA, LISTED BY FISH<br />
HOST SPECIES<br />
Fish host Cymothoid parasite Site of attachment<br />
Abudefduf saxatilis<br />
Acanthurus bahianus<br />
Acanthurus chirurgus<br />
Alutera schoepji<br />
Anchoa lamprotaenia<br />
Apogon lachneri<br />
Apogon maculatus<br />
Apogon townsendi<br />
Ariusfelis<br />
<strong>As</strong>trapogon stellatus<br />
Batrachoides surinamensis<br />
Caranx hippos<br />
Caranx latus<br />
Caranx ruber<br />
Caranx sp.<br />
Chaetodipterus faber<br />
Chaetodon capistratus<br />
Chaetodon ocellatus<br />
Chaetodon sedentarius<br />
Chaetodon striatus<br />
Anilocra abudefduji<br />
Kuna insularis<br />
Anilocra acanthuri<br />
Anilocra acanthuri<br />
Nerocila acuminata<br />
Lironeca tenuistylis<br />
Mothocya bohlkeorum<br />
Renocila colini<br />
Renocila colini<br />
Nerocila acuminata<br />
Mothocya bohlkeorum<br />
Nerocila acuminata<br />
Cymothoa oestrum<br />
Cymothoa oestrum<br />
Cymothoa oestrum<br />
Cymothoa oestrum<br />
Nerocila acuminata<br />
Anilocra chaetodontis<br />
Anilocra chaetodontis<br />
Anilocra chaetodontis<br />
Anilocra chaetodontis<br />
beneath eye<br />
gill chamber<br />
9 at base of pectoral fin;<br />
immature on or near<br />
pectoral or pelvic fin<br />
9 at base of pectoral fin;<br />
immature on or near<br />
pectoral or pelvic fin<br />
on or at base of fin<br />
posterior to pectoral fin<br />
in gill chamber<br />
next to dorsal fin<br />
next to dorsal fin<br />
on or at base of fin<br />
in gill chamber<br />
on or at base of fin<br />
inside mouth<br />
inside mouth<br />
inside mouth<br />
inside mouth<br />
on or at base of fin<br />
beneath eye<br />
beneath eye<br />
beneath eye<br />
beneath eye<br />
(continued)
172<br />
TABLE 3. {Continued)<br />
Fish host Cymothoid parasite Site of attachment<br />
Chilomycterus schoepfi<br />
Chromis cyaneus<br />
Chromis multilineatus<br />
Cynoscion nebulosus<br />
Cynoscion sp.<br />
Epinephelus cruentatus<br />
Epinephelus Julvus<br />
Epinephelus guttatus<br />
Epinephelus itajara<br />
Epinephelus sp.<br />
Exocoetus spp.<br />
Genes rhombeus<br />
Haemulon aurolineatum<br />
Haemulon carbonarium<br />
Haemulon chrysargyreum<br />
Haemulon Jlavolineatum<br />
Haemulon macrostomum<br />
Haemulon plumieri<br />
Haemulon sciurus<br />
Nerocila acuminata<br />
Anilocra chromis<br />
Anilocra chromis<br />
Cymothoa excisa<br />
Cymothoa oestrum<br />
Anilocra haemuli<br />
Anilocra haemuli<br />
Anilocra haemuli<br />
Nerocila acuminata<br />
Cymothoa oestrum<br />
Glossobius impressus<br />
Lironeca redmanni<br />
Anilocra haemuli<br />
Anilocra haemuli<br />
Anilocra haemuli<br />
Anilocra haemuli<br />
Anilocra haemuli<br />
Anilocra haemuli<br />
Anilocra haemuli<br />
Hemirhamphus brasiliensis Glossobius hemirhamphi<br />
Hirundichthys speculifer<br />
Holacanthus tricolor<br />
Holocentrus ascensionis<br />
Glossobius impressus<br />
Anilocra holacanthi<br />
Anilocra holocentri<br />
Hyporhamphus unifasciatus Mothocya nana<br />
Leiostomus xanthurus<br />
Lepiosteus spatula<br />
Lutjanus analis<br />
Lutjanus mahogoni<br />
Lutjanus synagris<br />
Megalops atlanticus<br />
Monacanthus ciliatus<br />
Mugil cephalus<br />
Myripristis jacobus<br />
Ocyurus chrysurus<br />
Orthopristis chrysoptera<br />
Nerocila acuminata<br />
Cymothoa excisa<br />
Lironeca redmanni<br />
Nerocila acuminata<br />
Cymothoa excisa<br />
Cymothoa excisa<br />
Cymothoa excisa<br />
Cymothoa oestrum<br />
Nerocila acuminata<br />
Nerocila acuminata<br />
Anilocra myripristi<br />
Cymothoa excisa<br />
Cymothoa excisa<br />
on or at base of fin<br />
beneath eye<br />
beneath eye<br />
inside mouth<br />
inside mouth<br />
beneath eye<br />
beneath eye<br />
beneath eye<br />
on or at base of fin<br />
on or at base of fin<br />
inside mouth<br />
in gill chamber<br />
beneath eye<br />
beneath eye<br />
beneath eye<br />
beneath eye<br />
beneath eye<br />
beneath eye<br />
beneath eye<br />
inside mouth<br />
inside mouth<br />
beneath eye<br />
? between eyes, 8 and<br />
immature beneath<br />
eye<br />
in gill chamber<br />
on or at base of fin<br />
inside mouth<br />
in gill chamber<br />
on at base of fin<br />
inside mouth<br />
inside mouth<br />
inside mouth<br />
inside mouth<br />
on or at base of fin<br />
on or at base of fin<br />
9 between eyes, imma<br />
ture beneath eye<br />
inside mouth<br />
inside mouth
Agarna cumulus 173<br />
Fish host Cymothoid parasite Site of attachment<br />
Orthopristis ruber<br />
Paranthias Jurcifer<br />
Phaeoptyx conklini<br />
Phaeoptyx pigmentaria<br />
Pogonias cromis<br />
Pomacentrus partitas<br />
Priacanthus arenatus<br />
Scomberomorus cavalla<br />
Scomberomorus maculatus<br />
Scomberomorus regalis<br />
Selar crumenophthalmus<br />
Serranus tigrinus<br />
Sphoeroides maculatus<br />
Synodus foetens<br />
Anilocra haemuli<br />
Anilocra haemuli<br />
Mothocya bohlkeorum<br />
Mothocya bohlkeorum<br />
Nerocila acuminata<br />
Anilocra partiti<br />
Cymothoa oestrum<br />
Lironeca redmanni<br />
Lironeca redmanni<br />
Lironeca redmanni<br />
Cymothoa oestrum<br />
Renocila bowmani<br />
Renocila waldneri<br />
Nerocila acuminata<br />
Cymothoa excisa<br />
Agarna Schioedte and Meinert, 1883<br />
beneath eye<br />
beneath eye<br />
in gill chamber<br />
in gill chamber<br />
on or at base of fin<br />
beneath eye<br />
inside nouth<br />
in gill chamber<br />
in gill chamber<br />
in gill chamber<br />
inside mouth<br />
next to dorsal fin<br />
next to dorsal fin<br />
on or at base of fin<br />
inside mouth<br />
DIAGNOSIS Cephalon with posterior margin not trilobed; immersed in per-<br />
eonite 1. Antennular bases contiguous. Pereonites 4-7 on one side flattened<br />
and expanded; coxal plates of pereopods 4-7 also expanded and flattened<br />
but generally hidden by lateral expansion of pereonites. Bases of posterior<br />
three pereopods with well-formed carinae. Pleonites 1 and 2 immersed in<br />
pereonite 7; pleonites 2-5 with free fingerlike lateral margins.<br />
Agarna cumulus (Haller, 1880)<br />
Figure 79<br />
DIAGNOSIS 9 18 mm. Eyes present, indistinct. Pereon strongly "humped"<br />
dorsally. Uropod about { fo length of pleotelson; uropodal exopod slightly<br />
longer, and twice width of endopod. Pleotelson triangular, length 3 A basal<br />
width, apex rounded.<br />
RECORDS No host recorded: Key West, Florida.
174 FLABELLIFERA • GYMOTHOIDAE<br />
Figure 79. Agarna cumulus: A, 9, dorsal view; By 9, ventral view, coxal plates<br />
stippled.<br />
Anilocra Leach, 1818<br />
DIAGNOSIS Cephalon usually narrowed anteriorly to triangular apex folded<br />
ventrally between bases of antennules; posterior margin trilobed; not im<br />
mersed, or only weakly immersed in pereonite 1. Coxal plates small, com<br />
pact, not reaching level of posterior margin of their respective pereonites.<br />
Pereopods increasing in length posteriorly, pereopod 7 often markedly longer<br />
than 6. Pleon not immersed or only slightly immersed in pereonite 7.<br />
Pleopods 3—5 often formed into deep pockets or pleats. Uropods often ex<br />
tending beyond pleotelsonic apex.<br />
REMARKS Williams and Williams (1981) have provided a comprehensive<br />
treatment of this genus and nine of its species in the West Indies. Table 1 in<br />
this latter paper provides characters for separating these nine species. This<br />
table also indicates that for each species, the site of attachment of the adult to<br />
the host fish is specific, with six species attaching under the eye of the host.
Key to species of Anilocra<br />
Anilocra abudefdufi 175<br />
1. Pereopods 2-4 with swelling on outer margin of dactylus 2<br />
Pereopods 2-4 lacking swelling on outer margin of dactylus 5<br />
2. Body axis distorted by more than 10° holacanthi<br />
Body axis distorted by less than 5° 3<br />
3. Dactylus of pereopod 7 longer than propodus partiti<br />
Dactylus of pereopod 7 shorter than propodus 4<br />
4. Posteroventral angle of pereonite 7 overlapping pleonite 1 only<br />
Posteroventral angle of pereonite 7 overlapping pleonites 1 and 2<br />
abudefdufi<br />
chaetodontis<br />
5. Posteroventral angle of pereonite 7 produced 6<br />
Posteroventral angle of pereonite 7 not produced 7<br />
6. Uropod reaching posterior margin of pleotelson myripristis<br />
Uropod not reaching posterior margin of pleotelson haemuli<br />
7. Posteroventral angle of pereonite 7 overlapping pleonite 1 . . . . holocentri<br />
Posteroventral angle of pereonite 7 not overlapping pleonite 1 8<br />
8. Uropod reaching posterior margin of pleotelson acanthuri<br />
Uropod not reaching posterior margin of pleotelson chromis<br />
Anilocra abudefdufi Williams and Williams, 1981<br />
Figure 80A-C<br />
DIAGNOSIS Ovigerous 9 19.0-31.0 mm, 6 7.0-8.5 mm. Pereopods 2-4<br />
with swelling on outer margin of dactylus. Posteroventral angle of pereonite 6<br />
slightly produced, of pereonite 7 more produced, overlapping pleonite 1.<br />
Uropodal endopod variable, not reaching, to extending well beyond, apex of<br />
exopod. Color: upper lateral half to three-fourths of dorsal surface of 9 when<br />
attached to host is dark brown; rest of dorsal surface light brown to yellow.<br />
Attaching beneath eye of host.
176 FLABELLIFERA • GYMOTHOIDAE<br />
Figure 80. Anilocra abudefdufi: A, 9, lateral view; B, pereopod 3; C, pereopod 7.<br />
Anilocra acanthuri: D, pleotelson and uropods. Anilocra chaetodontis: E, 9, lateral<br />
view. Anilocra chromis: F, pleotelson and uropods.<br />
Anilocra acanthuri Williams and Williams, 1981<br />
Figure 80D<br />
DIAGNOSIS Ovigerous 9 29.0-40.0 mm, 6 4.0-8.0 mm. Pereopods 2-4<br />
without swelling on outer margin of dactylus. Posteroventral angles of per-<br />
eonites not produced. Uropod not reaching posterior margin of pleotelson.<br />
Endopod of uropod variable, not reaching, to extending well beyond, apex of<br />
exopod. Color: dorsal surface of 9 black to lead gray, ventral surface gray.<br />
Attaching under pectoral fin of host.<br />
RECORDS Doctorfish Acanthurus chirurgus: Florida Keys; Bahamas; Puerto<br />
Rico; U.S. Virgin Islands. Ocean surgeon Acanthurus bahianus: Florida Keys;<br />
Bahamas; Cuba; Jamaica; Dominican Republic; Puerto Rico; U.S. Virgin<br />
Islands.
Anilocra chaetodontis Williams and Williams, 1981<br />
Figure 80E<br />
Anilocra haemuli 177<br />
DIAGNOSIS Ovigerous 9 18-28 mm, 6 4-5 mm. Pereopods 2-4 with swell<br />
ing on outer margin of dactylus, Posteroventral angles of pereonites 4—7 be<br />
coming progressively produced, that of pereonite 7 overlapping pleonite 2.<br />
Uropod not reaching posterior margin of pleotelson; uropodal endopod ex<br />
tending beyond apex of exopod. Pleotelson as wide as long to slightly wider<br />
than long. Color: dorsal surface of 9 black to lead gray, ventral surface gray.<br />
Attaching beneath eye of host.<br />
RECORDS Foureye butterflyfish Chaetodon capistratus: Bahamas; Puerto Rico;<br />
British and U.S. Virgin Islands. Banded butterflyfish Chaetodon striatus:<br />
Bahamas; Puerto Rico; British Virgin Islands. Spotfin butterflyfish Chaetodon<br />
ocellatus: Bahamas; Puerto Rico; U.S. Virgin Islands. Reef butterflyfish<br />
Chaetodon sedentarius: Puerto Rico.<br />
Anilocra chromis Williams and Williams, 1981<br />
Figure 80F<br />
DIAGNOSIS Ovigerous 9 16-28 mm, 8 4-9 mm. Pereopods 2-4 lacking<br />
swelling on outer margin of dactylus. Posteroventral angles of pereonites not<br />
produced. Uropod extending beyond posterior margin of pleotelson; uropo<br />
dal endopod not reaching beyond exopod. Color: upper lateral one-fourth to<br />
two-thirds of dorsal surface of 9 when attached is dark gray, shading to off-<br />
white lower lateral area. Attaching beneath eye of host.<br />
RECORDS Brown chromis Chromis multilineatus: Puerto Rico; British and<br />
U.S. Virgin Islands. Blue chromis Chromis cyaneus: Bahamas; Dominican Re<br />
public. No host recorded: Anguilla.<br />
Anilocra haemuli Williams and Williams, 1981<br />
Figure 81A,B<br />
DIAGNOSIS Ovigerous 9 21-40 mm, 6 7 mm. Body axis distorted less than<br />
5°. Pereopods 2—4 lacking swelling on outer margin of dactylus. Postero<br />
ventral angle of pereonites 6 and 7 produced, latter overlapping pleonite 1.<br />
Uropod not reaching posterior margin of pleotelson; uropodal endopod<br />
reaching beyond apex of exopod. Color: dorsal surface of 9 yellow to light<br />
brown. Attaching beneath eye of host.
178 FLABELLIFERA • GYMOTHOIDAE<br />
Figure 81. Anilocra haemuli: A, 9, dorsal view; B, 9, lateral view. Anilocra<br />
holacanthi: C, 9. Anilocra holocentri: D, 9. Anilocra myripristis: E, pleotelson and<br />
uropods. Anilocra partiti: F, 9; G, pereopod 7.<br />
RECORDS French grunt Haemulon Jlavolineatum: Florida Keys; Puerto Rico;<br />
British and U.S. Virgin Islands. Tomtate Haemulon aurolineatum: Jamaica;<br />
Puerto Rico. Smallmouth grunt Haemulon chrysargyreum: Puerto Rico; U.S.<br />
Virgin Islands. Caesar grunt Haemulon carbonarium: Puerto Rico; U.S. Virgin
Anilocra holocentri 179<br />
Islands. Spanish grunt Haemulon macrostomum: Puerto Rico. White grunt<br />
Haemulon plumieri: Florida Keys; Yucatan Peninsula. Bluestriped grunt<br />
Haemulon sciurus: Florida Keys. Cora cora Orthopristis ruber: Margarita Island,<br />
Venezuela. Coney Epinephelus Julvus: Bahamas; Dominican Republic; Puerto<br />
Rico; U.S. Virgin Islands; Guadeloupe. Red hind Epinephelus guttatus: Puerto<br />
Rico; British and U.S. Virgin Islands. Graysby Epinephelus cruentatus:<br />
Bahamas; Dominican Republic; U.S. Virgin Islands. Creole-fish Paranthias<br />
jurcifer: Dominican Republic; Puerto Rico; Colombia. No host recorded:<br />
Cuba; Jamaica; Dominica; Barbados; Venezuela; Brazil.<br />
Anilocra holacanthi Williams and Williams, 1981<br />
Figure 81C<br />
DIAGNOSIS Ovigerous 9 21-33 mm, 6 4-7 mm. Body axis distorted by<br />
more than 10°. Pereopods 2-4 with swelling on outer margin of dactylus.<br />
Posteroventral angles of pereonites 5—7 progressively more produced, that of<br />
pereonite 7 overlapping pleonite 1. Uropod not reaching posterior margin of<br />
pleotelson; uropodal endopod reaching beyond apex of exopod. Color: dorsal<br />
surface of 9 black to lead gray. Attaching beneath eye of host.<br />
RECORDS Rock beauty Holacanthus tricolor: Bahamas; Jamaica; Dominican<br />
Republic; Puerto Rico; British and U.S. Virgin Islands.<br />
Anilocra holocentri Williams and Williams, 1981<br />
Figure 81D<br />
DIAGNOSIS Ovigerous 9 32-46 mm, 6 5-9 mm. Body axis distorted less<br />
than 5°. Pereopods 2—4 lacking swelling on outer margin of dactylus.<br />
Posteroventral angle of pereonite 7 produced, overlapping pleonite 1.<br />
Uropod not reaching posterior margin of pleotelson; uropodal endopod<br />
reaching beyond apex of exopod. Color: dorsal surface of 9 dark brown,<br />
ventral surface light brown. 9 attaching between eyes of host; 6 or transi<br />
tional stage beneath eye.<br />
RECORDS Squirrelfish Holocentrus ascensionis: Puerto Rico; U.S. Virgin<br />
Islands.<br />
No host recorded: Patagonia, Straits of Magellan.
180 FLABELLIFERA • CYMOTHOIDAE<br />
Anilocra myripristis Williams and Williams, 1981<br />
Figure 81E<br />
DIAGNOSIS Ovigerous 9 29-40 mm, 6 6-7 mm. Body axis distorted less<br />
than 5°. Pereopods 2-4 lacking swellings on outer margin of dactylus.<br />
Posteroventral angle of pereonites 6 and 7 produced, latter overlapping<br />
pleonite 1. Uropod reaching beyond posterior margin of pleotelson; uropodal<br />
endopod reaching beyond apex of exopod. Color: dorsal surface of 9 light<br />
reddish brown, ventral surface yellow. 9 attaching between eyes of host;<br />
immature or transitional forms sometimes beneath eve.<br />
4<br />
RECORDS Blackbar soldierfish Myripristis jacobus: Bahamas; Dominican Re<br />
public; Puerto Rico.<br />
Anilocra partiti Williams and Williams, 1981<br />
Figure 81F,G<br />
DIAGNOSIS Ovigerous 9 12-16 mm, transitional 7.6-9.0 mm. Body axis<br />
distorted less than 5°. Pereopods 2-4 with swelling on outer margin of dac<br />
tylus. Pereopod 7 with dactylus longer than propodus. Posteroventral angle<br />
of pereonite 7 produced, overlapping pleonite 1. Uropod not reaching pos<br />
terior margin of pleotelson; uropodal endopod not reaching apex of exopod.<br />
Color: dorsal surface black to slate gray. Attaching beneath eye of host.<br />
RECORDS Bicolor damselfish Pomacentrus partitas: Jamaica.<br />
Ceratothoa Dana, 1852<br />
DIAGNOSIS Cephalon more or less immersed in pereonite 1, posterior mar<br />
gin not trisinuate. Bases of antennules expanded, contiguous. Coxal plates<br />
compact; anterior plates not extending beyond posterior margins of their<br />
respective pereonites; posterior coxal plates may or may not be produced<br />
beyond the posterior margins of the pereonites. Anterior pleonites narrowed,<br />
immersed in pereonite 7, Copulatory stylet lacking on pleopod 2 of 6 of some<br />
species.<br />
Ceratothoa deplanata Bovallius, 1885<br />
Figure 82A<br />
DIAGNOSIS 9 18 mm. Cephalon subtriangular, anterior margin rounded.<br />
Pereopods 4—7 with strongly carinate bases. Uropod reaching or extending
Ceratothoa deplanata 181<br />
Figure 82. A, Ceratothoa deplanata (from Bovallius, 1885); B, Cymothoa caraibica; C,<br />
Cymothoa excisa; D, Cymothoa oestrum.<br />
slightly beyond posterior margin of pleotelson; rami subequal in length and<br />
width. Pleotelson basally wider than long, posterior margin broadly<br />
rounded. Color: bright yellow.<br />
RECORDS Haiti, host not recorded.
182 FLABELUFERA • GYMOTHOIDAE<br />
Cymothoa Fabricius, 1793<br />
DIAGNOSIS Body usually not distorted. Cephalon with posterior margin not<br />
trilobed; more or less immersed in pereonite 1; latter with anterolateral cor<br />
ners produced to embrace cephalon. Bases of antennules not expanded, well<br />
separated. Anterior coxal plates not reaching posterior borders of their re<br />
spective pereonites, posterior coxal plates nearly reaching or extending be<br />
yond posterior borders of pereonites. Pleon narrower than, and immersed in<br />
pereonite 7. Pleonites increasing in length and width posteriorly.<br />
Key to species of Cymothoa<br />
1. Anterolateral angles of pereonite 1 reaching to half length of cephalon<br />
or less; eyes or traces of eyes present 2<br />
Anterolateral angles of pereonite 1 broad, reaching to anterior margin<br />
of cephalon; eyes absent oestrum<br />
2. Anterolateral angles of pereonite 1 narrow, subacute excisa<br />
Anterolateral angles of pereonite 1 broad, rounded caraibica<br />
Cymothoa caraibica Bovallius, 1885<br />
Figure 82B<br />
DIAGNOSIS 9 17 mm, 6 12-16 mm. Anterior margin of cephalon broadly<br />
rounded. Eyes large, distinct. Broadly rounded anterolateral angles of per<br />
eonite 1 reaching to about midlength of cephalon. Bases of pereopods 4—7<br />
with strong, rounded carina. Uropodal rami subequal in length, equal to<br />
peduncle in length. Pleotelson width about twice length, posterolateral mar<br />
gin broadly rounded.<br />
RECORDS Puerto Rico; Gulf of Mexico.<br />
Cymothoa excisa Perty, 1833<br />
Figure 82C<br />
DIAGNOSIS Ovigerous 9 20-24 mm. Anterior margin of cephalon in dorsal<br />
view truncate to slightly excavate; eyes small, indistinct. Anterolateral angles<br />
of pereonite 1 narrowly rounded to subacute, reaching anteriorly to about<br />
midlength of cephalon. Pereopods 4—7 with high rounded carina on basis.
Glossobius 183<br />
Uropods hardly reaching halfway along lateral margin of pleotelson; exopod<br />
slightly longer than endopod. Pleotelson about twice wider than long;<br />
broadly rounded and somewhat bilobed.<br />
RECORDS Yellowtail snapper Ocyurus chrysurus: Yucatan Peninsula, Mexico;<br />
Carrie Bow Cay, Belize; Margarita Island, Venezuela; Panama. Mutton<br />
snapper Lutjanus analis: Yucatan Peninsula, Mexico; Panama. Lane snapper<br />
Lutjanus synagris: Panama. Mahogany snapper Lutjanus mahogoni: Panama.<br />
Pigfish Orthopristis chrysoptera: Florida, Gulf of Mexico. Spot Leiostomus<br />
xanthurus: Texas, Gulf of Mexico. Spotted seatrout Cynoscion nebulosus: Texas,<br />
Gulf of Mexico. Inshore lizardfish Synodusfoetens: Texas, Gulf of Mexico. No<br />
host recorded: Massachusetts; South Carolina; Georgia; Florida Keys;<br />
Bahamas; Cuba; Trinidad; Brazil.<br />
Cymothoa oestrum (Linnaeus, 1793)<br />
Figure 82D<br />
DIAGNOSIS Ovigerous 9 38 mm. Cephalon in dorsal view with ante<br />
rolateral angles rounded, anterior margin slightly excavate; eyes absent. An<br />
terolateral angles of pereonite 1 expanded, broadly rounded, reaching to<br />
level of anterior margin of cephalon. Pereonites 4—7 with high rounded<br />
carina on basis. Uropod reaching posteriorly beyond midlength of<br />
pleotelson; exopod slightly longer than endopod. Pleotelson length slightly<br />
more than half basal width.<br />
RECORDS Bigeye scad Selar crumenophthalmus: Bermuda; U.S. Virgin Is<br />
lands. Bigeye Priacanthus arenatus: Bermuda. Bar jack Caranx ruber: Florida<br />
Keys; Carrie Bow Cay, Belize. Horse-eye jack Caranx latus: Bahamas; Bar<br />
bados. Crevalle jack Caranx hippos: Venezuela. Jack Caranx sp.: Jamaica; Cu<br />
rasao. Hind Epinephelus sp.: Grenada. Parrotfish: Jamaica. Seatrout Cynoscion<br />
sp.: Panama. Tarpon Megalops atlantica: Texas, Gulf of Mexico. No host re<br />
corded: Honduras; Haiti.<br />
Glossobius Schioedte and Meinert, 1883<br />
DIAGNOSIS Cephalon not immersed in pereonite 1; excavate on either side<br />
in anterior half, forming broad and anteriorly rounded median area; anten<br />
nae fitting into excavate areas. Bases of antennules contiguous, expanded.<br />
Antennules broader and longer than antennae. Bases of pereopods 4—7 with<br />
posterior margin expanded and flattened. Pleonites 1-3 immersed in per<br />
eonite 7.
184 FLABELLIFERA • CYMOTHOIDAE<br />
Key to species of Glossobius<br />
1. Coxal plates of perconites 1 and 2 anteroventrally protruding impressus<br />
Coxal plates of pereonites 1 and 2 close to body, not protruding<br />
Glossobius hemiramphi Williams and Williams, 1985a<br />
Figure 83A<br />
hemirhamphi<br />
DIAGNOSIS Ovigerous 9 27 mm. Eyes small but distinct. Cephalon pointed<br />
anteriorly. Fused coxa of pereonite 1 and free coxa of pereonite 2 carinate but<br />
not protruding. Coxa of pereonite 7 semicircular in dorsal view. Pleotelson<br />
with middorsal length more than half basal width; lateral margins somewhat<br />
tapered; posterior margin variable, sinuate or excavate. Uropods reaching to<br />
or slightly beyond posterior pleotelsonic margin; rami subequal in length,<br />
exopod slightly broader than endopod.<br />
RECORDS Ballyhoo Hemiramphus brasiliensis; Puerto Rico.<br />
Glossobius impressus (Say, 1818)<br />
Figure 83B<br />
DIAGNOSIS Ovigerous 9 33 mm. Eyes small but distinct. Cephalon<br />
rounded anteriorly. Fused coxal plate of pereonite 1 and distinct coxal plate<br />
of pereonite 2 protruding strongly in oblique anteroventral direction. Uropod<br />
reaching to posterior half of pleotelson; exopod shorter and narrower than<br />
endopod. Pleotelson basal width twice length, posteriorly broadly bilobed.<br />
Attaching inside mouth of host.<br />
RECORDS Flyingfish Exocoetus spp.: Rio de Janeiro, Brazil; North Atlantic,<br />
especially in the Gulf Stream.<br />
Mirrorwing flyingfish Hirundichthys speculifer: North Atlantic. No host rec<br />
ord: Senegal, West Africa.<br />
Kuna Williams and Williams, 1986<br />
DIAGNOSIS Cephalon somewhat immersed in pereonite 1. Anterior margin<br />
of pereonite 1 not trisinuate. Number of articles in antennules and antennae
Kuna 185<br />
Figure 83. A, Glossobius hemiramphi; B, Glossobius impressus; C, Kuna insularis; D,<br />
Lironeca redmani; E, Lironeca tenuistylis.<br />
reduced. Antennule somewhat expanded; basal article expanded but not<br />
contiguous. Copulatory stylet present on pleopods 1-3 in 6. Pleonites dor-<br />
sally strongly convex, not immersed in pereonite 7.
186 FLABELLIFERA • GYMOTHOIDAE<br />
Kuna insularis (Williams and Williams, 1985b)<br />
Figure 83C<br />
DIAGNOSIS Ovigerous 9 11.1-17.2 mm, 6 4.2-8.7 mm, transitional 9.6-<br />
9.8 mm. Antennules and antennae consisting of four articles each. Uropods<br />
short, not reaching posterior margin of pleotelson. Clavate eopulatory stylet<br />
present on pleopods 1—3 in 6. Pleotelson basally broader than long, pos<br />
terior margin broadly rounded.<br />
RECORDS Sergeant major Abudefdufsaxatilis: Carrie Bow Cay, Belize; Cura<br />
sao; Panama.<br />
Lironeca Leach, 1818<br />
DIAGNOSIS Cephalon weakly to deeply immersed in pereonite 1; posterior<br />
border rarely trisinuate. Bases of antennules not expanded, well separated.<br />
Posterior pereopods with carinae on bases in 6, carinae present or absent in<br />
9. Pleonites subequal in width; pleonites 1 and 2 rarely narrowed and<br />
weakly to moderately immersed in pereonite 7. Pleopods highly folded, and<br />
with lamellar or digitiform accessory gills in some species.<br />
Key to species of Lironeca<br />
1. Uropodal endopod about twice longer than wide; pleon somewhat<br />
immersed in pereon redmanni<br />
Uropodal endopod about three times longer than wide; pleon barely<br />
immersed in pereon tenuistylis<br />
Lironeca redmanni Leach, 1818<br />
Figure 83D<br />
DIAGNOSIS Ovigerous 9 19.5-25.0 mm. Cephalon barely immersed in per<br />
eonite 1. Pleon somewhat immersed in pereon, but lateral margins of pleonite<br />
1 free. Pleotelson basally wider than long. Uropodal rami reaching well be<br />
yond posterior margin of pleotelson; exopod longer than endopod, both rami<br />
somewhat broad, endopod about twice longer than wide. Attaching to gills of<br />
host.
Mothocya 187<br />
RECORDS New Jersey to Florida; gills of kingfish, Jamaica; Cuba; St.<br />
Christopher; Spanish mackerel Scomberomorus maculatus and cero Scomberomorus<br />
regalisy Puerto Rico; king mackerel Scomberomorus cavalla, Colombia; Genes rho-<br />
mbeus, Panama; spot Leiostomus xanthurus, Gulf of Mexico.<br />
Brazil.<br />
Lironeca tenuistylis (Richardson, 1912b)<br />
Figure 83E<br />
DIAGNOSIS 9 13 mm. Cephalon barely immersed in pereonite 1. Uropodal<br />
rami reaching beyond rounded posterior margin of pleotelson; exopod longer<br />
than endopod; endopod slender, about three times longer than wide. Pleonite<br />
1 barely immersed in pereonite 7. Pleotelson basally wider than long. Attach<br />
ing to host between pectoral and anal fin.<br />
RECORDS Longnose anchovy Anchoa lamprotaenia: Panama.<br />
Mothocya Costa, 1851<br />
DIAGNOSIS Cephalon more or less immersed in pereonite 1. Bases of anten-<br />
nules widely separated; antennules longer and more robust than antennae.<br />
Coxae nearly reaching or extending beyond posterior margin of respective<br />
pereonites. Pleon somewhat immersed in pereonite 7. Uropodal exopod<br />
longer than endopod.<br />
REMARKS Bruce (1986b) revised the genus Mothocya, The species of Moth<br />
ocya are almost entirely gill parasites on the fish families Hemiramphidae,<br />
Apogonidae, Belonidae, and Atherinidae.<br />
Key to species of Mothocya<br />
1. Cephalon anteriorly narrowed, slightly immersed in pereonite 1;<br />
pleotelson subrectangular bohlkeorum<br />
Cephalon anteriorly broad, deeply immersed in pereonite 1; pleotelson<br />
subtriangular nana
188 FLABELLIFERA • GYMOTHOIDAE<br />
Mothocya bohlkeorum Williams and Williams, 1982<br />
Figure 84B<br />
DIAGNOSIS Ovigerous 9 7.6-8.5 mm, 6 3.7 mm. Cephalon anteriorly nar<br />
rowed in dorsal view, ventrally flexed, broadly rounded; slightly immersed in<br />
pereonite 1. Pleotelson subrectangular. Uropods extending slightly beyond<br />
posterior margin of pleotelson; exopod only slightly longer than endopod. 9<br />
lateral lobes of pleopodal peduncles not developed. Endopods of pleopods 3-<br />
5 with small proximomedial lobe.<br />
RECORDS Whitestar cardinalfish Apogon lachneri: Puerto Rico. Dusky car-<br />
dinalfish Phaeoptyx pigmentaria: Bahamas. Freckled cardinalfish Phaeoptyx con-<br />
klini: Florida Keys; Bahamas. Conchfish <strong>As</strong>trapogon stellatus: Leeward<br />
Islands.<br />
Mothocya nana (Schioedte and Meinert, 1884)<br />
Figure 84A<br />
DIAGNOSIS Ovigerous 9 11.0-17.0 mm, 6 7.9-8.3 mm. Cephalon deeply<br />
immersed in pereonite 1; rostrum anteroventrally narrowly rounded. Uropo-<br />
dal exopod markedly longer than endopod. Pleotelson broad, with posterior<br />
margin rounded sufficiently to give appearance of being subtriangular.<br />
RECORDS Halfbeak Hyporhamphus unifasciatus: Chesapeake Bay, Maryland;<br />
Georgia; Florida; Colon, Panama. Halfbeak Hemiramphus bermudensis:<br />
Bermuda.<br />
Nerocila Leach, 1818<br />
DIAGNOSIS Body generally more depressed than in most cymothoid genera,<br />
rarely curved. Cephalon with anterior margin convex, narrowly rounded, or<br />
concave; not, or only slightly, immersed in pereonite 1. Pereonite 1 anterior<br />
margin trisinuate. Posterolateral angles of pereonites weakly to strongly pro<br />
duced, increasing in length posteriorly. Coxal plates prominent, usually al<br />
most reaching or extending to posterior margin of their respective pereonites.<br />
Juveniles and 6 usually with spines on posterior pereopods; 9 lacking these<br />
spines. Pleon not immersed in pereonite 7. Pleonites subequal in length;<br />
pleonites 1 and 2 usually produced posterolateral^. Pleopods typically with<br />
small lamellar accessory gills; pleopods 3-5 folded into deep pockets or<br />
pleats. Uropods usually extending beyond pleotelsonic apex.
Figure 84. A, Mothocya nana; By Mothocy<br />
acuminata; D, Nerocila acuminata f. aster.<br />
B
190 FLABELLIFERA • CYMOTHOIDAE<br />
Nerocila acuminata Schioedte and Meinert, 1881<br />
DIAGNOSIS Ovigerous 9 16.2-19.0 mm. Cephalon with anterior margin<br />
convex. Posterolateral angles of all, or of posterior pereonites only, produced<br />
into acute or subacute angles.<br />
RECORDS Striped burrfish Chilomycterus schoepfi: Texas, Gulf of Mexico.<br />
Northern puffer Sphoeroides maculatus: New York. Striped mullet Mugil<br />
cephalus: Texas, Gulf of Mexico. Jewfish Epinephelus itajara: Texas, Gulf of<br />
Mexico. Hogfish: Bermuda. Alligator gar Lepisosteus spatula: Louisiana, Gulf<br />
of Mexico. Hardhead catfish Ariusfelis: Texas, Gulf of Mexico. Sawfish: Flor<br />
ida (Atlantic). Black drum Pogonias cromis: Texas, Gulf of Mexico. Orange<br />
filefish Alutera schoepfi: Texas, Gulf of Mexico. Toadfish Batrachoides sur-<br />
inamensis: Colon, Panama. Spot Leiostomus xanthurus: Florida, Gulf of Mexico.<br />
Spadefish Chaetodipterus faber: Florida, Gulf of Mexico; Virginia. Fringed fil<br />
efish Monacanthus ciliatus: Florida, Gulf of Mexico. No host recorded: Mas<br />
sachusetts; Florida Keys; Florida, Gulf of Mexico. Louisiana, Gulf of Mex<br />
ico. Texas, Gulf of Mexico.<br />
REMARKS Brusca (1981) has shown that this highly variable species occurs<br />
on both sides of the Isthmus of Panama, in two relatively distinct forms.<br />
Intergrades between the two forms do occur but are uncommon. Brusca<br />
(1981:159) also lists all the host-records for this species in the eastern Pacific.<br />
Nerocila acuminata Schioedte and Meinert, 1881, forma acuminata<br />
Figure 84C<br />
DIAGNOSIS Cephalon width equal to or greater than length; frontal margin<br />
narrowly rounded. Posterolateral angles of anterior pereonites weakly pro<br />
duced, rounded to subacute; of posterior pereonites more strongly produced,<br />
subacute to acute. Coxal plates 3—7, 4—7, or 5—7 with acute posterolateral<br />
angles; coxae rarely reaching beyond posterior margins of their respective<br />
pereonites.<br />
Nerocila acuminata Schioedte and Meinert, 1881, forma aster<br />
Figure 84D<br />
DIAGNOSIS Cephalon always wider than long; anterior margin broadly<br />
rounded. Posterolateral angles of all pereonites strongly produced, acute, all<br />
reaching well beyond posterior margins of their respective pereonites. Coxal<br />
plates 2—7 strongly produced with acute posterior angles.
Renocila Miers, 1880<br />
Renocila colini 191<br />
DIAGNOSIS Body rarely curved. Cephalon anteriorly weakly to distinctly<br />
truncate. Antennular bases well separated. Antennules and antennae some<br />
what flattened, antennules usually broader and longer than antennae. Per-<br />
eonites 5—7 with posterolateral corners more or less strongly produced.<br />
Pleonites not laterally incised.<br />
REMARKS Williams and Williams (1980) provide a key to nine species of<br />
Renocila.<br />
Key to species of Renocila<br />
1. Posteroventral angle of pereonite 7 reaching pleonite 1 colini<br />
Posteroventral angle of pereonite 7 reaching beyond pleonite 1 2<br />
2. Dorsal surface of body brown; posteroventral angle of pereonite 7<br />
reaching pleonite 2 waldneri<br />
Dorsal surface of body black; posteroventral angle of pereonite 7<br />
reaching pleonite 3 bowmani<br />
Renocila bowmani Williams and Williams, 1980<br />
Figure 85A<br />
DIAGNOSIS 9 18.0 mm, 6 11.5 mm. Posteroventral angles of pereonites 5—<br />
7 produced, that of pereonite 7 overlapping pleonites 1-3. Pereopods 1—3<br />
lacking swelling on dactylus. Pereopods 6-7 subequal in length. Uropodal<br />
exopod longer than endopod. Pleotelson length 3 A basal width. Color: dorsal<br />
surface of body and appendages uniform black. Attached to dorsum of body<br />
close to dorsal fin.<br />
RECORDS Harlequin bass Serranus tigrinus: Dominican Republic.<br />
Renocila colini Williams and Williams, 1980<br />
Figure 85B,C<br />
DIAGNOSIS Ovigerous 9 12.0-17.5 mm, 6 7.5-13.0 mm. Pereonites 5-7<br />
with posteroventral angle produced, that of pereonite 7 overlapping pleonite<br />
1 only. Pereopods 1—3 lacking swelling on dactyli; pereopods 6—7 subequal in<br />
length. Uropod reaching beyond pleotelson, endopod more than half length
192 FLABELLIFERA • CYMOTHOIDAE<br />
c<br />
Figure 85. A, Renocila bowmani. Renocila colini: B, 9; C, d. D> Renocila waldneri.<br />
of exopod. Pleotelson l /i to 72 wider than long, with slight rounded apex<br />
Color: dorsal surface of body and appendages uniformly yellowish brown<br />
Attached to dorsum of body, close to dorsal fin.
FLABELLIFERA • L1MNOR11DAE 193<br />
RECORDS Flamefish Apogon maculatus: Puerto Rico. Belted cardinalfish Ap-<br />
ogon townsendi: Puerto Rico.<br />
Renocila waldneri Williams and Williams, 1980<br />
Figure 85D<br />
DIAGNOSIS Ovigerous 9 15.3-19.3 mm, 6 5.0-10.8 mm. Posteroventral<br />
angle of pereonite 5 moderately produced, of pereonites 6-7 more strongly<br />
produced, that of pereonite 7 overlapping pleonites 1 and 2. Pereopods 1-3<br />
without swelling on dactyli. Pereopods 6 and 7 subequal in length. Uropodal<br />
exopod slightly longer than endopod. Pleotelson basally wider than long;<br />
posterior margin broadly and evenly rounded. Color: dorsal surface of body<br />
uniform brown; appendages yellowish brown. Attached to dorsum of body<br />
close to dorsal fin.<br />
RECORDS Harlequin bass Serranus tigrinus: Dominican Republic.<br />
Family Limnoriidae Harger, 1879<br />
DIAGNOSIS Body ovate in cross section, often becoming more setose posteri<br />
orly. Cephalon subspherical, freely articulating with pereonite 1; eyes lateral.<br />
Antennules and antennae well separated at bases. Mandible with strong in<br />
cisor; lacking molar and well-defined lacinia mobilis, but with species-<br />
distinctive lacinioid bristle or seta; palp usually of three articles. Maxillipe-<br />
dal palp of five articles; endite well developed. Coxae present on pereonites<br />
2-7. Pleon consisting of five free pleonites plus pleotelson; latter subcircular,<br />
set obliquely to axis of body, usually with anterolateral crests. Uropod with<br />
strong protopod inserted ventrolaterally.<br />
Key to genera of Limnoriidae<br />
1. Uropodal rami very unequal 2<br />
Uropodal rami subequal Paralimnoria<br />
2. Mandibular incisors possessing rasp and file Limnoria<br />
Mandibular incisors lacking rasp and file Phycolimnoria
194 FLABELLIFERA • LIMNORIIDAE<br />
REMARKS This family includes a number of species that are of considerable<br />
economic importance. Given that species of Limnoria are wood borers,<br />
wooden structures such as wharf pilings that are immersed in sea water and<br />
even in water of reduced salinity are vulnerable to attack by these gribbles.<br />
Prolonged exposure can lead to weakening and eventual collapse of these<br />
structures (see Ray, 1959). Even creosote-treated wood is not fully protected;<br />
Limnoria tuberculata will bore into such wood to where the creosote has not<br />
penetrated.<br />
The isopods rasp at the wood fibres with the rasp and file structures of the<br />
mandibles, usually following the grain of the wood. With this boring activity,<br />
saprophytic fungi and bacteria invade the wood and assist in the breakdown<br />
process. Limnoria lack cellulase-secreting microflora in their gut, but proba<br />
bly secrete a cellulase themselves (Boyle and Mitchell, 1978). It is also prob<br />
able that the fungi and bacteria, the latter often densely aggregated on the<br />
setae of the isopod, form part of the animals' diet. In the natural environ<br />
ment, Limnoria perform an important role in the breakdown of dead wood,<br />
especially in mangrove areas.<br />
Sexual dimorphism of the pleotelson does occur in some species. This as<br />
pect of the morphology, however, has hardly been investigated.<br />
Limnoria Leach, 1814<br />
DIAGNOSIS Antennular flagellum of four articles. Antennal flagellum of<br />
three to five articles. Incisor of right mandible equipped with filelike struc<br />
ture on upper surface; incisor of left mandible with rasplike structure. Rami<br />
of pleopod 5 lacking marginal setae. Uropodal exopod much shorter than<br />
endopod, bearing terminal claw. Pleotelson smooth, or variously ornamented<br />
with tubercles and ridges.<br />
Limnoria indica Becker and Kampf, 1958<br />
Figure 86A,B<br />
DIAGNOSIS 6 3.0 mm, ovigerous 9 3.0 mm. Pleonite 5 with submedian pair<br />
of strong rounded ridges, converging slightly posteriorly. Pleotelson basally<br />
with two pairs of submedian tubercles and pair of lateral tubercles.<br />
RECORDS Cozumel, Mexico; Man o'War Cay, Belize.<br />
India; Hong Kong; Philippines; east coast of Australia.
Key to species of Limnoria<br />
Limnoria insulae 195<br />
1. Dorsal surface of pleotelson lacking prominent tubercles, ridges, or<br />
carinae (L. simulata may appear to lack ornamentation; in this<br />
species the tubercles are very small) 2<br />
Dorsal surface of pleotelson bearing tubercles, ridges, or carinae .... 3<br />
2. Pleotelson flat; pleonite 5 with broadly rounded middorsal ridge<br />
Pleotelson cup shaped; pleonite 5 with strong narrowly rounded<br />
platycauda<br />
middorsal ridge insulae<br />
3. Pleotelson with basal tubercles but lacking ridges 4<br />
Pleotelson with ridges but lacking freestanding tubercles 7<br />
4. 6 pleotelson with single strong middorsal tubercle unicornis<br />
Pleotelson with more than one basal tubercle 5<br />
5. Pleotelson with three basal tubercles tuberculata<br />
Pleotelson with more than three basal tubercles 6<br />
6. Pleotelson with four basal tubercles in line (difficult to detect) simulata<br />
6 pleotelson with six basal tubercles indica<br />
7. Pleotelson with single middorsal longitudinal ridge multipunctata<br />
Pleotelson with two rounded basal ridges 8<br />
8. Pleonite 5 with strong Y-shaped ridge Pfeffe<br />
Pleonite 5 with two posteriorly converging ridges saseboensis<br />
Limnoria insulae Menzies, 1957<br />
Figure 86C<br />
DIAGNOSIS 6 3.0 mm, ovigerous 9 3.4 mm. Pleonite 5 with strong middor<br />
sal ridge. Pleotelson cup shaped, lateral crests extended anteromesially, sep<br />
arated basally by distinct gap; posterior margin and lateral crests not<br />
tuberculate.<br />
RECORDS Twin Cays, Belize.
196 FLABELLIFERA • LIMNORIIDAE<br />
200HM<br />
Figure 86. Limnoria indica: A, pi 6;B, pi 9. Limnoria insulae: C<br />
pleotelson. Limnoria multipunctata: D, pleotelson in oblique-lateral view<br />
L i m n o r i a m u l t i p u n c t a t a M e n z i e s , 1 9 5 7<br />
F i g u r e s 8 6 D ; 8 7 A<br />
D I A G N O S I S 6 2 . 8 m m , o v i g e r o u s 9 3 . 0 m m . P l e o n i t e 5 d o r s a l l y s m o o t h .<br />
P l e o t e l s o n w i t h m i d d o r s a l l o n g i t u d i n a l r o u n d e d r i d g e b e a r i n g s e v e r a l
Limnoria multipunctata 197<br />
Figure 87. Limnoria multipunctata: A, pleotelson; Limnoria pfefferi: B} pleotelson<br />
Limnoria platycauda: C, pleotelson; Limnoria saseboensis: D> pleotelson; Limnoria<br />
simulata: E, pleotelson; Limnoria tuberculata: F, pleotelson.<br />
b u t t o n - s h a p e d t u b e r c l e s i n p o s t e r i o r h a l f ; p o s t e r i o r m a r g i n a n d l a t e r a l c r e s t s<br />
t u b e r c u l a t e .<br />
R E C O R D S P u e r t o R i c o ; J a m a i c a ; T w i n C a y s , B e l i z e<br />
J a p a n ; K a i I s l a n d s , S o u t h P a c i f i c .
198 FLABELLIFERA • LIMNORIIDAE<br />
Limnoria pfefferi Stebbing, 1904<br />
Figure 87B<br />
DIAGNOSIS 8 3.8 mm, ovigerous 9 4.0 mm. Pleonite 5 with conspicuous<br />
middorsal Y-shaped carina. Pleotelson basally with pair of submedian<br />
rounded ridges; lateral crests lacking tubercles.<br />
RECORDS Florida Keys; Bahamas; Puerto Rico; U.S. Virgin Islands; Twin<br />
Cays and Man o'War Cay, Belize; Yucatan Peninsula, Mexico.<br />
Minikoi Atoll and Aldabra Atoll, Indian Ocean; Philippines; New Guinea;<br />
Panama.<br />
Limnoria platycauda Menzies, 1957<br />
Figure 87C<br />
DIAGNOSIS 6 2.5 mm, ovigerous 9 2.6 mm. Pleonite 5 with broad middor<br />
sal longitudinal rounded ridge. Pleotelson lacking dorsal ornamentation;<br />
posterior margin and lateral crests bearing tubercles.<br />
RECORDS Cuba; Puerto Rico to Curasao; Cozumel, Mexico; Twin Cays<br />
and Man o'War Cay, Belize.<br />
Aldabra Atoll, Indian Ocean.<br />
Limnoria saseboensis Menzies, 1957<br />
Figure 87D<br />
DIAGNOSIS 6 3.5 mm. Pleonite 5 with submedian pair of ridges, converging<br />
slightly posteriorly. Pleotelson basally with submedian pair of anteriorly tu-<br />
berculate ridges; posterior margin and lateral crests tuberculate.<br />
RECORDS Miami, Florida.<br />
Japan; Fiji.<br />
Limnoria simulata Menzies, 1957<br />
Figure 87E<br />
DIAGNOSIS 6 3.8 mm, ovigerous 9 4.0 mm. Pleonite 5 with obscure me<br />
dian longitudinal groove. Pleotelson basally with submedian pair of tubercles<br />
and small lateral tubercles, latter often difficult to detect; lateral crests<br />
tuberculate.<br />
RECORDS Florida Keys; U.S. Virgin Islands; Gulf of Mexico.
Limnoria tuberculata Sowinsky, 1884<br />
Figure 87F<br />
Paralimnoria andrewsi 199<br />
DIAGNOSIS 6 2.8 mm, ovigerous 9 3.0 mm. Pleonite 5 with two anterior<br />
tubercles, one middorsal posterior tubercle, area between tubercles de<br />
pressed. Pleotelson basally with middorsal tubercle, followed by pair of sub-<br />
median tubercles, all three tubercles having short obscure carina; posterior<br />
margin and lateral crests tuberculate.<br />
RECORDS Rhode Island to Venezuela; Cuba; Man o'War Cay, Belize; Gulf<br />
of Mexico.<br />
Uruguay; West Africa; Mediterranean; Black Sea; India; Hong Kong;<br />
Hawaii; Australia; California.<br />
REMARKS This species has frequently been recorded under the name Lim<br />
noria tripunctata Menzies, 1951a.<br />
Limnoria unicornis Menzies, 1957<br />
Figure 88A,B<br />
DIAGNOSIS 6 2.6 mm, ovigerous 9 2.6 mm. Mandibular palp of one arti<br />
cle. Pleonite 5 with somewhat obscure Y-shaped ridge middorsally.<br />
Pleotelson in 6 with strong basal slightly curved middorsal tubercle; lateral<br />
crests lacking tubercles.<br />
RECORDS Bahamas; Man o'War Cay and Twin Cays, Belize.<br />
Caroline Islands; Palau; Society Islands.<br />
Paralimnoria Menzies, 1957<br />
DIAGNOSIS Antennular flagellum of five articles. Antennal flagellum of five<br />
or six articles. Mandibular incisor with rasp and file. Pleopod 5, rami bearing<br />
marginal setae. Uropodal rami subequal in length, each with clawlike apex.<br />
Paralimnoria andrewsi (Caiman, 1910)<br />
Figure 88C,D<br />
DIAGNOSIS 6 2.6 mm, 9 2.6 mm. Pleonite 5 with or without triangular<br />
middorsal depressed area. Pleotelson with basal submedian pair of tubercles<br />
either obscurely or strongly carinate; lateral crest tubercles of variable<br />
strength.
200 FLABELLIFERA • LIMNORIIDAE<br />
Figure 88. Limnoria unicornis: A, pleotelson, 8; B, pleon, 6*, in lateral view.<br />
Paralimnoria andrewsi: C, p l e o n i t e 5 a n d p l e o t e l s o n ; D , u r o p o d . Phycolimnoria clarkae:<br />
E , p l e o n i t e 5 a n d p l e o t e l s o n ; F , u r o p o d a n d p l e o t e l s o n i n l a t e r a l v i e w .<br />
R E C O R D S F l o r i d a K e y s ; P u e r t o R i c o ; T w i n C a y s , B e l i z e ; C u r a g a o<br />
C h r i s t m a s I s l a n d s , I n d i a n O c e a n ; S a m o a ; H a w a i i ; J a p a n .<br />
R E M A R K S M e n z i e s ( 1 9 5 7 ) d i s c u s s e s t h r e e f o r m s o f t h i s s p e c i e s : F o r m a typ-<br />
i c a , w h i c h l a c k s a c e n t r a l d e p r e s s e d a r e a d o r s a l l y o n p l e o n i t e 5 a n d h a s a p a i r<br />
o f s u b m e d i a n o b s c u r e l y c a r i n a t e t u b e r c l e s o n t h e p l e o t e l s o n ; F o r m a A , w h i c h<br />
h a s a t r i a n g u l a r d e p r e s s e d a r e a d o r s a l l y o n p l e o n i t e 5 a n d a p a i r o f s u b m e -
FLABELLIFERA • SEROLIDAE 201<br />
dian tubercles supported by strong carinae on the pleotelson; Forma B5 hav<br />
ing a triangular depressed area dorsally on pleonite 5 and an obscurely cari-<br />
nate pair of tubercles on the pleotelson. Given that at least two of these forms<br />
have been recorded occurring together, it would seem that this is merely a<br />
highly variable species.<br />
Phycolimnoria Menzies, 1957<br />
DIAGNOSIS Mandibular incisor lacking rasp and file. Uropodal rami une<br />
qual, exopod longer than endopod, latter usually with clawlike apex.<br />
REMARKS Most species of Phycolimnoria are algal borers, frequently encoun<br />
tered in the holdfasts of brown algae such as Macrocystis, Laminaria, and<br />
Sargassum, The one species recorded from the Caribbean, P. clarkae, however,<br />
has only been taken from decaying wood.<br />
Phycolimnoria clarkae Kensley and Schotte, 1987<br />
Figure 88E,F<br />
DIAGNOSIS 6 4.3 mm, ovigerous 9 3.3-4.4 mm. Uropodal exopod less<br />
than half length of endopod, straight, tipped with short squat claw. Pleonite<br />
5 with broad raised middorsal region having irregular bumps. Pleotelson<br />
wider than long, with two rounded submedian ridges basally, becoming ob<br />
solete posteriorly.<br />
RECORDS Bahamas; Twin Cays, Belize.<br />
Aldabra Atoll, Indian Ocean.<br />
Family Serolidae Dana, 1852<br />
DIAGNOSIS Body dorsoventrally depressed. Eyes present or absent.<br />
Cephalon fused with pereonite 1 dorsally. Mandible bearing palp. Maxillipe-<br />
dal palp of one to four articles. Pereonites 2—4 with coxae demarked; per-<br />
eonites 5 and 6 with coxae not demarked; pereonite 7 narrow, lacking free<br />
lateral margins. Pereopod 1 in 6 and 9 subchelate, pereopod 2 subchelate or<br />
ambulatory in 6, ambulatory in 9. Pleonites 1 and 2 free, articulated, re<br />
mainder of pleonites fused with telson. Pleopods 1—3 small, natatory;<br />
pleopods 4 and 5 large, operculate. Uropods lateral, biramous.<br />
REMARKS The serolids reach their greatest diversity (and their greatest size<br />
of up to 80 mm in length) in the southern oceans, with few species extending
202 FLABELLIFERA • SPHAEROMATIDAE<br />
into the subtropics and tropics. The deep- and abyssal-dwelling species usu<br />
ally lack eyes. The animals are epibenthic, living in the upper few centime<br />
ters of the bottom sediment, where they are scavengers and carnivores.<br />
Serolis Leach, 1818<br />
DIAGNOSIS Body markedly dorsoventrally flattened. Coxal plates produced<br />
laterally. Mandible having lacinia mobilis and single spine. Maxillipedal<br />
palp of three articles (rarely two to four). Pereopod 2 exhibiting sexual di<br />
morphism, subchelate in cJ, ambulatory in 9. Pleopods 1-3, peduncles elon<br />
gate, rami subelliptical. Pleopod 3, exopod uniarticulate.<br />
Serolis mgrayi Menzies and Frankenberg, 1966<br />
Figure 89<br />
DIAGNOSIS S 4.5 mm, ovigerous 9 4.7 mm. Eyes present. Cephalon with<br />
two middorsal tubercles. Pereonites 2—4 each with faint rounded tubercle<br />
just mesial to coxal suture. Pereon and pleon with faint middorsal longitudi<br />
nal carina bearing small blunt tubercle on posterior margin of each segment.<br />
Pleonites 1 and 2 with lateral margins not contributing to body outline, over<br />
lapped by pereonite 6. Pleotelson broadly triangular, with lateral carina in<br />
anterior half; apex truncate. Uropodal rami reaching to or slightly beyond<br />
pleotelsonic apex.<br />
RECORDS Off North Carolina, 18-34 m; off South Carolina, 22 m; off<br />
Georgia, 18-47 m; Florida Keys, 18-88 m; Trinidad; Venezuela, 95 m; Flor<br />
ida, Gulf of Mexico, 11-88 m.<br />
Family <strong>Sphaeromatidae</strong> H. Milne Edwards, 1840<br />
DIAGNOSIS Antennular peduncle of three articles, antennal peduncle of five<br />
articles. Mandible stout, lacinia mobilis and molar usually well developed,<br />
palp of three articles. Maxillipedal palp of five articles. Mouthparts in some<br />
genera metamorphosed and somewhat reduced in ovigerous 9 . Pleon of five<br />
partially or completely fused pleonites, often indicated by lateral sutures,<br />
plus dorsally convex and sometimes inflated pleotelson. Uropods lateral, ex<br />
opod free if present, endopod fused with sympod. Sexual dimorphism often<br />
marked, especially in pleotelsonal structure. Animal often capable of con<br />
globating or folding over. Young brooded in internal pouches or anterior or<br />
posterior pockets; oostegites variable in number, if present.
FLABELLIFERA • SPHAEROMATIDAE 203<br />
Figure 89. Serolis mgrayi: A, 6; B, pereopod 1; C, pereopod 2, 8.<br />
REMARKS Right into the 1980s this family was routinely divided into three<br />
groups, based on the structure of the two posterior pairs of pleopods:<br />
Platybranchiatae—pleopods 4 and 5 with both rami membranous and lack<br />
ing branchial pleats; Hemibranchiatae—pleopods 4 and 5 with branchial<br />
pleats on endopods only; Eubranchiatae—pleopods 4 and 5 with branchial<br />
pleats on both rami. These three "groups" were recognized formally as sub<br />
families by Hurley and Jansen (1977) but the names were not based on con-
204 FLABELL1FERA • SPHAEROMATIDAE<br />
tained genera and were replaced with current subfamily names by Bowman<br />
(1981) and Iverson (1982), the latter providing diagnoses for all five sub<br />
families. Four of these are represented in the Caribbean area; the fifth, the<br />
Tecticipitinac, contains only the single primarily Pacific genus Tecticeps.<br />
While the subfamilial status now appears to be resolved, many of the gen<br />
era still require unambiguous diagnoses. The work of Harrison (1984) on the<br />
structure of the female broodpouch, with its various components of<br />
oostegites, internal pouches, and anterior and posterior pockets (Figure 90),<br />
along with the metamorphosis of the female mouthparts (see Figure 96) has<br />
helped enormously to standardize the genera. Nevertheless, these features of<br />
the female remain unknown in several genera. Further, with this stabilization<br />
based on females, many problems of incorrect generic designation have been<br />
uncovered. In this work, Harrison's generic diagnoses are followed as far as<br />
possible. Where uncertainty exists, this is indicated. In some cases, we may<br />
still be unaware of existing problems: future work will without doubt result in<br />
the shifting of species to different genera, as well as in the creation of new<br />
genera.<br />
Key to subfamilies of <strong>Sphaeromatidae</strong><br />
1. Pereopod 1 prehensile in both sexes; pereopod 2 prehensile only in 6<br />
Ancininae<br />
Pereopods 1 and 2 ambulatory 2<br />
2. Pleopods 4 and 5 lacking branchial pleats Cassidininae<br />
Pleopods 4 and 5 with branchial pleats on endopods 3<br />
3. Pleopods 4 and 5 with branchial pleats on both rami . . . Dynameninae<br />
Pleopods 4 and 5 with branchial pleats on endopods only<br />
Subfamily Ancininae Tattersall, 1905<br />
Sphaeromatinae<br />
DIAGNOSIS Body markedly dorsoventrally depressed. Cephalon fused me<br />
dially with pereonite 1. Pereopod 1 prehensile in 8 and 9. Pereopod 2 pre<br />
hensile in 6 only. Pleopods 4 and 5 similar, lacking branchial pleats.<br />
Uropods uniramous.
internal pouch<br />
Ancinus belizensis 205<br />
Figure 90. Diagrammatic representation of 9 sphaeromatid, showing marsupial<br />
structures (adapted from Harrison, 1984).<br />
Ancinus H. Milne Edwards, 1840<br />
DIAGNOSIS Eyes dorsal. 9 mouthparts not metamorphosed. Mandibular<br />
molar absent; palp of three articles. Maxilla 1 of single ramus, endite rudi<br />
mentary. Maxilla 2 of two rami. 9 with oostegites absent; brood held in two<br />
opposing pockets, opening as narrow ventral slit between pereopods 4. Pleon<br />
consisting of short anterior pleonite with free lateral margin, plus broadly<br />
triangular pleotelson. Pleopod 1 uniramous, endopod absent. Pleopod 2 op-<br />
erculiform. Pleopod 3, exopod of single article. Uropod lacking exopod, sym-<br />
pod not laterally expanded.<br />
Key to species of Ancinus<br />
1. Pleotelson as long as basal width, apex narrowly rounded . . . brasiliensis<br />
Pleotelson with basal width greater than length, apex subtruncate<br />
Ancinus belizensis Kensley and Schotte, 1987<br />
Figure 91A-C<br />
belizensis<br />
DIAGNOSIS 8 4.1 mm, 9 2.8 mm. Body oval, about twice longer than wide.<br />
Dorsal integument strongly pitted. Antennular flagellum of 12 articles; an-
206 FLABELLIFERA • SPHAEROMATIDAE<br />
Figure 91. Ancinus belizensis: A, 9; B, pereopod 1 cJ; C, pereopod 2 8. Ancinus<br />
braziliensis: D, adult (from Glynn and Glynn, 1974).<br />
tennal flagellum of 10 articles. 8 pereopod 2, dactylus strongly curved,<br />
reaching to proximal lobe of propodus. Pleopod 2 about 2.5 times longer than<br />
basal width.<br />
RECORDS Carlson Point, Belize, in seagrass flats, 0.5 m.<br />
Ancinus brasiliensis Lemos de Castro, 1959<br />
Figure 91D<br />
DIAGNOSIS 8 7.0 mm, 9 6.0 mm. Body about twice longer than wide. Dor<br />
sal integument smooth. Antennular flagellum of 17 articles; antennal<br />
flagellum of 10 articles. 8 pereopod 2, dactylus strongly curved, reaching to<br />
midlength of posterior margin of carpus. Pleopod 2 almost three times longer<br />
than basal width.
Cass idin idea 207<br />
RECORDS Brazilian coast from Rio de Janeiro northward, 1.5 m; Costa<br />
Rica, Panama; shallow infratidal below sandy beaches.<br />
REMARKS Glynn and Glynn (1974) discussed color polymorphism in this<br />
species.<br />
Subfamily Cassidininae Iverson, 1982<br />
DIAGNOSIS Cephalon not medially fused with pereonite 1. Pereopod 1 am<br />
bulatory. Pleopods 4 and 5, both rami lacking transverse pleats, outer rami<br />
unsegmented. Pleopod 5, outer ramus with low subapical squamiferous pro<br />
tuberances. Pleotelsonic apex entire. Uropods with exopods reduced.<br />
REMARKS The genus Dies has twice been recorded from the Caribbean: D.<br />
arndti Ortiz and Lalana, 1980, from Cuba, and D. barnardi Carvacho, 1977,<br />
from Guadeloupe. This genus is distinguished from Cassidinidea solely on the<br />
basis of the penial structure: biramous in Cassidinidea, uniramous in Dies.<br />
Harrison (1984) has pointed out that the separation of these two genera has<br />
not been satisfactorily resolved. The penis of neither the Cuban nor the<br />
Guadeloupan species has been illustrated, but the whole-animal illustrations<br />
of both look suspiciously like Cassidinidea ovalis. Examination of material of D.<br />
barnardi from the Paris Museum supports the view that this species was based<br />
on immature material of C. ovalis. Neither of the so-called species oi Dies are<br />
dealt with in this work, both being regarded as junior synonyms of C. ovalis.<br />
Key to genera of Cassidininae<br />
1. Frontal lamina visible dorsally between antennular bases; two basal<br />
articles of antennular peduncle not expanded Cassinidinea<br />
Frontal lamina not visible between antennular bases; two basal articles<br />
of antennular peduncle broadly expanded Paraleptosphaeroma<br />
Cassidinidea Hansen, 1905b<br />
DIAGNOSIS Body strongly dorsoventrally depressed. Eyes dorsal, situated<br />
at posterolateral corners of cephalon. Latter somewhat sunken into pereonite<br />
1. Frontal lamina expanded, visible dorsally between antennular bases. An<br />
tenna directed laterally. Pleon consisting of one free pleonite having short<br />
free lateral margin, plus broadly triangular pleotelson. Uropodal endopod
208 FLABELL1FERA • SPHAEROMATIDAE<br />
well developed, fused with sympod; exopod markedly reduced. Penial rami<br />
elongate, separate. 9 mouthparts not metamorphosed. Oostegites absent.<br />
Brood housed in pouch formed by opposing pockets overhanging ventrum,<br />
opening by slit between fourth pereopods.<br />
Key to species of Cassidinidea<br />
1. Posterior margin of pleotelson truncate ovalis<br />
Posterior margin of pleotelson rounded mosaica<br />
Cassidinidea mosaica Kensley and Schotte, 1987<br />
Figure 92A<br />
DIAGNOSIS 6 1.8 mm, ovigerous 9 1.6 mm. Body twice longer than wide.<br />
Dorsal integument bearing close-packed flattened tubercles. Pleotelson tri<br />
angular, with posterior margin narrowly rounded, dorsally convex, basally<br />
inflated.<br />
RECORDS Carrie Bow Cay, Belize, 1.5-10 m; in silty sand and rubble be<br />
tween patch reefs and coral buttresses.<br />
Cassidinidea ovalis (Say, 1818)<br />
Figure 92B-E<br />
DIAGNOSIS 6 and 9 3.6 mm. Body width slightly less than half length.<br />
Dorsal integument smooth. Pleotelson with raised anteromesial area, but<br />
lacking sculpture; posterior margin truncate.<br />
RECORDS New Jersey to Florida, in marsh mud and among dead leaves, 0—<br />
1 m; Trinidad; Belize; Panama; Dominica; Louisiana and Vera Cruz, Gulf of<br />
Mexico. Known from waters of less than l%o to 35%o.<br />
Paraleptosphaeroma Buss and Iverson, 1981<br />
DIAGNOSIS Body oval in outline, entire circumference with transparent<br />
flange of fused setae on two expanded basal articles of antennule, on per-<br />
conites, pleonite 1, and uropods. Expanded basal articles of antennules con-
Paraleptosphaeroma 209<br />
Figure 92. Cassidinidea mosaica: A, 6. Cassidinidea ovalis: B, 6] C, pereopod 1; Dy<br />
pleopod 4; E, pleopod 5. Paraleptosphaeroma glynni: Fy 6.<br />
tiguous in midline. Single articulated pleonite with short free lateral margin.<br />
Uropodal sympod and endopod fused; exopod articulated, much shorter<br />
than fused endopod.
210 FLABELLIFERA • SPHAEROMATIDAE<br />
Paraleptosphaeroma glynni Buss and Iverson, 1981<br />
Figure 92F<br />
DIAGNOSIS 6 2.58 mm, ovigerous 9 2.38 mm. Pleotelson basally broad,<br />
tapering to notched posterior margin. Fused uropodal endopod and sympod<br />
of each side almost touching posterior to pleotelsonic apex.<br />
RECORDS Portsmouth, Dominica, intertidal rock pools.<br />
Punta Paitilla, Pacific Panama.<br />
REMARKS Buss and Iverson (1981) demonstrated that this species displays<br />
sequential protogynous hermaphroditism, and that the change from female<br />
to male seems to be mediated by social conditions, especially the proportion<br />
of males to females. The principal food source for this species was shown to<br />
be abascan bryozoans.<br />
Key to genera of Dynameninae<br />
1. Pleotelson very similar in both sexes 2<br />
Pleotelson showing marked sexual dimorphism 3<br />
2. Cephalon and pleotelson smooth, lacking ridges Ischyromene<br />
Pleotelson and cephalon with ridges Cerceis<br />
3. Uropods lamellar in both sexes 4<br />
Uropods lamellar in 9, endopod reduced, exopod elongate-cylindrical<br />
in 6 5<br />
4. Ovigerous 9 lacking oostegites;
Subfamily Dynameninae Bowman, 1981<br />
Discerceis 211<br />
DIAGNOSIS Cephalon not fused with pereonite 1. Pereopods 1 and 2 am<br />
bulatory. Pleopods 4 and 5, both rami having branchial pleats. Pleopod 4,<br />
exopod unjointed, usually lacking setae, endopod with few setae at most.<br />
Pleotelsonic apex often with terminal notch or foramen, especially in
212 FLABELLIFERA • SPHAEROMATIDAE<br />
Figure 93. "Cerceis" carinata: A, 6. Discerceis linguicauda: B, d. Dynamenella<br />
acutitelson: C, 9; D, pleon 8. Dynamenella angulata: E, 9.<br />
internal pouches (number unknown). Pockets absent. 6 with uropodal endo-<br />
pod and sympod fused, very short; exopod elongate, cylindrical.
Discerceis linguicauda (Richardson, 1901)<br />
Figure 93B<br />
Dynamenella 213<br />
DIAGNOSIS 6 7.2 mm. Dorsal integument, especially of posterior half, with<br />
numerous scattered granular tubercles. Uropodal endopod and sympod<br />
fused, very short, exopod elongate, subcylindrical and slightly bowed. Ante<br />
rior half of pleotelson inflated, with three elongate rounded ridges (each com<br />
posed of two contiguous tubercles) ending posteriorly in subacute tubercle;<br />
posterior margin trilobed, median lobe broadly rounded, with subacute tu<br />
bercle at base, lateral lobe truncate, well separated from median lobe. Head<br />
and pereonite 1 not fused. Frontal lamina visible dorsally between antennal<br />
bases. Penes short, separate. Copulatory stylet basally relatively broad, dis-<br />
tally broadly rounded.<br />
RECORDS Cape Catoche, Yucatan, Mexico, 48-50 m.<br />
REMARKS This species is known only from the four male syntypes.<br />
Dynamenella Hansen, 1905b<br />
DIAGNOSIS Species exhibiting obvious sexual dimorphism. Both sexes lack<br />
ing processes on pereon and pleon, Uropodal rami lamellar. Exopod of<br />
pleopod 3 with or without articulation. 9: Mouthparts not metamorphosed.<br />
Broodpouch lacking oostegites, but formed by two opposing ventral pockets<br />
opening in midline between fourth pereopods. Apex of pleotelson with notch,<br />
Key to species of Dynamenella<br />
1. 6 with pleotelsonic foramen 2<br />
6 lacking foramen but with notch, or appearing entire; 9 pleotelson<br />
with faint notch visible acutitelson<br />
2. 6 with four strong pleotelsonic ridges; 9 with subcircular pleotelsonic<br />
foramen quadrilirata<br />
6 lacking pleotelsonic ridges; 9 with posterior margin of pleotelson<br />
entire perforata
214 FLABELLIFERA • SPHAEROMATIDAE<br />
groove, or foramen. 6: Penes basally fused, rami long, tapering. Copulatory<br />
stylet proximally broad, tapering to acute tip, reaching to or just beyond<br />
apex of endopod. Uropods broader than in $. Posterior pleotelson with dor-<br />
sally directed foramen connected to apex by narrow slit.<br />
REMARKS The species described by Richardson (1901) as Dynamene angulata<br />
from No Name Key, Florida, and referred to by some authors as a Dyna-<br />
menella, while figured here (Figure 93E), is not included in the present key.<br />
The species is known only from immature females; correct generic placement<br />
is thus not possible.<br />
Dynamenella acutitelson Menzies and Glynn, 1968<br />
Figure 93C,D<br />
DIAGNOSIS 6 3.5 mm, 9 2.3 mm. 6: Pereonites 4-6 with transverse ridge<br />
over dorsum, ridge interrupted to form short median section. Pleotelson with<br />
two submedian and two lateral rounded tubercles basally, two submedian,<br />
poorly defined ridges in central area; posterior margin tapering in dorsal<br />
view, with slit eitherjust visible or appearing entire. In lateral view, posterior<br />
pleotelson seen to be laterally compressed, forming narrow groove.<br />
RECORDS Puerto Rico, intertidal rocks and algae.<br />
REMARKS Menzies and Glynn (1968) described this species with two vari<br />
eties, the holotype as D. acutitelson var. typica, and 11 paratypes as D. acu<br />
titelson var. glabrothorax. The major difference between these varieties lay in<br />
the presence of transverse ridges on pereonites 4—6 in typica and their absence<br />
xnglabrothorax. The holotype, however, at 3.5 mm, would seem to be a mature<br />
male, while all the paratypes are smaller. The differences described by Men<br />
zies and Glynn (1968) may thus be due to immaturity. <strong>As</strong> further compara<br />
tive material is lacking, these varieties (or whatever their true status) are not<br />
recognized here.<br />
Menzies and Glynn (1968, fig. 30a) illustrate D. acutitelson var. glabrothorax<br />
as having scattered tiny granules over the dorsal integument. These were not<br />
seen when the type material was reexamined.<br />
Harrison and Holdich (1982) placed this species in Paradella, based on the<br />
literature. However, the penes for both varieties are shown as short and sepa<br />
rate, as in Ischyromene. Again, until further mature males and ovigerous<br />
females are seen, the generic placement of this species must remain in doubt.
Dynamenella perforata (Moore, 1901)<br />
Figure 94A,B<br />
Geocerceis barbarae 215<br />
DIAGNOSIS 6 3.2 mm, 9 2.6 mm. 6: Pleon bearing two low rounded sub-<br />
median "mounds." Pleotelson with strongly convex anterior two-thirds, with<br />
T-shaped foramen. Pleon and pleotelson with numerous scattered small tu<br />
bercles. Uropodal rami broadly ovate, outer margins crenulate. 9:<br />
Pleotelson broadly rounded in dorsal view, posterior margin entire. Inner<br />
uropodal ramus distally subacute.<br />
RECORDS Bermuda to Puerto Rico, intertidal coral rubble and algae, and<br />
under chiton Acanthopleura granulate; Dominican Republic; Cuba.<br />
Dynamenella quadrilirata Kensley, 1984<br />
Figure 94C-H<br />
DIAGNOSIS 6 2.6 mm, 9 2.5 mm. 6: Two low rounded submedian tuber<br />
cles on last pleonite. Anterior half of pleotelson inflated, with four rounded<br />
longitudinal ridges; posterior half tapered, somewhat dorsally flexed, with<br />
cordate foramen. Uropodal rami distally rounded, outer margins crenulate<br />
to dentate. 9: Lacking pleonal tubercles. Pleotelson inflated, unornamented,<br />
posterior margin forming subcircular foramen.<br />
RECORDS Carrie Bow Cay, and Twin Cays, Belize; intertidal to 3 m.<br />
Geocerceis Menzies and Glynn, 1968<br />
DIAGNOSIS Ovigerous 9 with mouthparts metamorphosed. Broodpouch<br />
with three pairs of oostegites, on pereonites 2—4, just overlapping in midline.<br />
Brood held in internal pouches (number unknown). Pockets absent. Uropo<br />
dal rami lamellar, shorter than pleotelson. 6 uropodal endopod fused with<br />
sympod, very short; exopod elongate, club shaped. Pleopod 2 with copula-<br />
tory stylet articulating distally on endopod.<br />
Geocerceis barbarae Menzies and Glynn, 1968<br />
Figure 95A-C<br />
DIAGNOSIS 8 3.3 mm, 9 2.5 mm. Pleopod 3 exopod of single article. Pleon<br />
with two elongate sutures reaching lateral pleon margin. 6: Frontal lamina<br />
expanded into ventrally directed beaklike process. Penes separate, relatively
216 FLABELLIFERA • SPHAEROMATIDAE<br />
Figure 94. Dynamenella perforata: A, 6; B> pleon 9. Dynamenella quadrilirata: C, 6;<br />
D, pleon 9; E, pleon
Ischyromene 217<br />
Figure 95. Geocerceis barbarae: A, 8; B, pleon 9; C,
218 FLABELLIFERA • SPHAEROMATIDAE<br />
nounced. Uropodal rami lamellar. 8 pleopod 2 with copulatory stylet<br />
basally narrow, reaching to or just beyond distal margin of endopod.<br />
Ischyromene barnardi (Menzies and Glynn, 1968)<br />
Figure 95D<br />
DIAGNOSIS 6 4.5 mm, 9 3.7 mm. Both sexes lacking processes on pereon<br />
and pleon. Accessory unguis of pereopods often bifid. Pleopod 3, exopod of<br />
single article. Uropodal rami lamellar. 6: Pereonite 7, posterior margin<br />
bilobed. Penes short, separate to base.<br />
RECORDS Puerto Rico, intertidal.<br />
Paracerceis Hansen, 1905b<br />
DIAGNOSIS Pleopod 3 exopod with transverse suture in distal half. Pleon<br />
with two long sutures reaching to posterolateral margin. 6: Penial rami<br />
short, separate. Pleotelson with basal area strongly vaulted; deep posterior<br />
notch sometimes having denticles on inner margins, and/or median tooth at<br />
base of notch. Uropodal endopod short, fused with sympod; exopod elongate,<br />
club shaped. 9: Mouthparts metamorphosed. Mandible fused with<br />
cephalon. Broodpouch of four pairs of oostegites, three posterior pairs over<br />
lapping. Brood retained in internal pouches. Uropodal rami subequal, lamel<br />
lar. Pleon usually less ornamented than in
Paracerceis caudata 219<br />
3. 6, pleotelsonic notch deep, margins usually with two teeth on each<br />
side; strong median tubercle on anterior pleotelson bluntly bifid; 9,<br />
pleotelson with one or two rounded median tubercles and 2 smaller<br />
tubercles on each side caudata<br />
6, pleotelsonic notch shallow, with tiny lateral denticles; median<br />
tubercle of pleotelson conical, acute; 9, pleotelson with three large<br />
conical acute tubercles and several smaller scattered tubercles in<br />
anterior half cohenae<br />
Paracerceis caudata (Say, 1818)<br />
Figure 96<br />
DIAGNOSIS 6 8.1 mm, 9 6.4 mm. 8: Pleotelson with blunt median bifid<br />
tubercle, with two smaller tubercles on each side. Pleotelsonic notch usually<br />
with two strong denticles on each margin, basal median tooth lacking.<br />
Uropodal exopod reaching well beyond pleotelson, slightly bowed, with 2—4<br />
setose bumps on outer margin. 9: Pleonite 5 with three low tubercles.<br />
Pleotelsonic apex broadly rounded in dorsal view, with two rounded median<br />
tubercles and two smaller tubercles on each side. Uropodal rami subequal,<br />
lamellar, outer distal angle of each acute.<br />
RECORDS Bermuda; New Jersey to Florida Keys; Yucatan to Venezuela;<br />
Turks and Caicos Islands; Cuba; Puerto Rico; Bahamas; Jamaica; Haiti; St.<br />
Maartens, 0.2-127 m; St. Lucia; Gulf of Mexico. Found in the following<br />
algae: Caulerpa, Halimeda, Turbinaria> Amphiroa, Laurencia, Dictyota; between<br />
sponges and tunicates on red mangrove roots; in coral rubble; in spur and<br />
groove zone of reefs, lagoon, back reef, seagrass flats, and fringing<br />
mangroves.<br />
REMARKS Menzies and Glynn (1968:55, fig. 22f) named and figured P.<br />
caudata var. brevipes from Puerto Rico. This variant was characterized as hav<br />
ing the margins of the pleotelsonic notch lacking denticles. Given the con<br />
siderable variation in ornamentation in this species, we feel that no validity<br />
can be given to the name "brevipes."<br />
This is the commonest sphaeromatid in the Caribbean, and it has very<br />
broad ecological requirements, being found in a wide range of habitats and<br />
depths.
220 FLABELL1FERA • SPHAEROMATIDAE<br />
Figure 96. Paracerceis caudata: A, 8,B, pleon 9; C, mandible 6] D, maxilla 1 6; E,<br />
maxilla 2 6; F, maxilliped 6; G> mandible 9; Hy maxilla 19;/, maxilla 1 9; J,<br />
maxilliped 9.<br />
Paracerceis cohenae Kensley, 1984<br />
Figure 97A,B<br />
DIAGNOSIS 6 10.0 mm, 9 7.9 mm. 6: Pereonites each with median tuber<br />
cle and several smaller lateral tubercles near posterior margin of somite.<br />
Pleonite 5 with large median conical tubercle. Anterior two-thirds of
Paracerceis glynni 221<br />
pleotelson inflated, faintly tripartite, with strong median conical tubercle;<br />
notch in posterior margin shallow, with low median tooth and tiny lateral<br />
denticles; posterolateral margins finely dentate. Uropodal exopod cylindri<br />
cal, distally denticulate, six to seven times longer than basal width. 9: Per-<br />
eon and pleon much as in 6 y but pleotelsonic notch shallower and<br />
posterolateral margins not denticulate. Uropodal rami subequal, lamellar,<br />
exopod with distolateral angle acute.<br />
RECORDS Carrie Bow Cay, Belize, 15-16 m. Only known from sponge<br />
Callispongia plicifera growing on outer reef slope.<br />
Paracerceis edithae Boone, 1930<br />
Figure 97C-E<br />
DIAGNOSIS 6 4.0 mm, 9 3.1 mm. 6: Posterior three pereonites and<br />
pleonites each with irregular row of small tubercles near posterior margin,<br />
densely setulose tubercles becoming spinose more posteriorly. Pleotelson<br />
with strong median conical tooth in anterior half, flanked by convex spinose<br />
mound. Pleotelsonic notch deep, with elongate median basal tooth bearing<br />
strong acute tooth at its base. Lobes of posterior pleotelsonic margin broad,<br />
flattened, margins denticulate. Uropodal exopod tuberculate, tapering, api-<br />
cally acute. 9: Integument much less tuberculate-spinose than in 6. Imma<br />
ture 9, posterior margin of pleotelson with faintly rounded median lobe. In<br />
mature 9, posterior margin distinctly trilobed. Uropodal rami subequal,<br />
lamellar, distally rounded, with tiny distolateral spine on exopod.<br />
RECORDS Bahamas, 60-66 m, in vase sponge; Haiti; Puerto Rico, 20—25 m.<br />
Paracerceis glynni Kensley, 1984<br />
Figure 97F,G<br />
DIAGNOSIS 6 6.4 mm, 9 5.2 mm. 6: Integument becoming strongly setose<br />
and tuberculate posteriorly from about pereonite 5. Posterior margin of inf<br />
lated anterior area of pleotelson bearing strong median conical tubercle and<br />
smaller acute lateral tubercle, with low swelling beneath each lateral tuber<br />
cle. Posterior notch deep, narrow, with small basal median tooth, lobes form<br />
ing notch tricuspid, outer cusps recurved dorsally. Uropodal exopod fairly<br />
straight, cylindrical, apically acute. 9: Body far less setose and tuberculate<br />
than 6. Pleotelson with strongly inflated anterior area having very faint mid-<br />
dorsal tubercle; notch well marked, formed by triangular lobes of
222<br />
FLABELLIFERA • SPHAEROMATIDAE<br />
E<br />
Figure 97. Paracerceis cohenae: A, 6\ B, pleotelson, 9. Paracerceis edithae: C,<br />
pleotelson, 9; D, mature 6] E, immature S. Paracerceis glynni: F, 8\ G} pleotelson,<br />
9. Paracerceis nuttingi: H, 9.<br />
pleotelsonic margin. Uropodal rami subequal, flattened, endopod with distal<br />
margin faintly trituberculate; exopod with few distal tubercles.<br />
RECORDS Alligator Light, Florida, 11 m; Carrie Bow Cay, Belize, 11-15.2<br />
B<br />
G
Paradella 223<br />
m, from green alga Halimeda sp. on forereef, and from sponge Aphysina<br />
jistularis.<br />
Paracerceis nuttingi (Boone, 1921)<br />
Figure 97H<br />
RECORDS Barbados; Puerto Rico, 1.5 m, from Cymodocea seagrass, and coral<br />
rubble and sponges.<br />
REMARKS The types of this species from Barbados consist only of females<br />
(total length 4.1 mm). Menzies and Glynn (1968) record an immature male<br />
i<br />
from Puerto Rico with an incipient pleotelsonic notch. This specimen, how<br />
ever, still has the subequal lamellar uropodal rami. The mature male, with<br />
the characteristically reduced uropodal endopod and cylindrical exopod, is<br />
unknown. The possibility exists that this is not a true Paracerceis.<br />
Paradella Harrison and Holdich, 1982<br />
DIAGNOSIS Marked<br />
ual dimorphism. Accessory unguis of pereopods simple, not bifid. Pleopod 3<br />
Key to species of Paradella<br />
1. Pereonite 7 with projecting bilobed flange; pleon and pleotelson finely<br />
but distinctly granulate 2<br />
Pleon and pleotelson smooth 3<br />
2. 8 with pleotelsonic foramen distinctly heart shaped, with median<br />
point; four submedian tubercles of pleotelson in 8 and 9 somewhat<br />
elongate; 9 pleotelson posteriorly narrowed, slit visible dorsally<br />
8 with pleotelsonic foramen wider than long, but lacking median<br />
point; four submedian tubercles of pleotelson in 8 small, rounded,<br />
obscure in 9; 9 pleotelson posteriorly truncate, slit not visible<br />
dianae<br />
dorsally plicatura<br />
3. Tubercles on pleotelson in 8 and 9 small to obscure; 8 with<br />
pleotelsonic foramen subcircular quadripunctata<br />
Tubercles on pleotelson broadly rounded mounds in 8 and 9; 8 with<br />
pleotelsonic foramen wider than long tumidicauda
224 FLABELL1FERA • SPHAEROMAT1DAE<br />
exopod with articulation. Uropodal rami lamellar. 9: Mouthparts not meta<br />
morphosed. One pair of oostegites arising from pereonite 4, short, not reach<br />
ing midline. Brood held in pouch formed by two opposing pockets covering<br />
entire ventrum, opening by transverse slit between 4th pereopods.
Paradella quadripunctata 225<br />
Figure 98. Paradella dianae: A, 8; B, pleopod 2 8; C, pleon 9. Paradella plicatura:<br />
Dy 8; E, pleon 9. Paradella quadripunctata: F, 8; G, pleon 9. Paradella tumidicauda;<br />
H, pleon 9 (from Glynn, 1970); /,
226 FLABELL1FERA • SPHAEROMAT1DAE<br />
Paradella tumidicauda (Glynn, 1970)<br />
Figure 98H,I<br />
DIAGNOSIS 8 6.7 mm, 9 6.5 mm. 6: Last pleonite with two submedian<br />
swellings. Pleotelson with four submedian broadly rounded swellinglike tu<br />
bercles and two pairs of lateral tubercles. Foramen wider than long, posterior<br />
contiguous borders of foramen each bearing rounded swelling. 9: Pleotelson<br />
with four submedian swellinglike tubercles, sometimes with two obscure lat<br />
eral tubercles; posterior slit not visible dorsally, area surrounding slit<br />
swollen, horseshoe shaped.<br />
RECORDS Margarita Island, Venezuela, from among intertidal barnacles.<br />
Subfamily Sphaeromatinae H. Milne Edwards, 1840<br />
DIAGNOSIS Cephalon not fused with pereonite 1. Pereopods 1 and 2 am<br />
bulatory. Pleopods 4 and 5, endopods having branchial pleats, exopods un-<br />
pleated, membranous, of two articles. Uropods biramous.<br />
Key to genera of Sphaeromatinae<br />
1. Uropodal exopod with outer margin serrate Sphaeroma<br />
Uropodal exopod with outer margin entire or faintly crenulate 2<br />
2.
Cymodoce ruetzleri 227<br />
Pleopod 5, exopod of two articles, distal article with apex and internal mar<br />
gin covered with fine teeth, anterior surface with long distally toothed boss;<br />
proximal article with two small toothed bosses at internodistal angle. 6:<br />
Maxillipedal palp articles 2—4 bearing setigerous lobes. Penial rami elongate,<br />
separate. Pleon usually more tuberculate than in 9. Uropodal exopod lamel<br />
lar, shorter than endopod. 9: Mouthparts metamorphosed. Broodpouch<br />
formed by four pairs of oostegites arising from pereonites 1-4, overlapping in<br />
midline. Brood housed in five pairs of internal pouches.<br />
"Cymodoce" barrerae (Boone, 1918)<br />
Figure 99A,B<br />
DIAGNOSIS 9 7.5 mm. 9: Body dorsally strongly vaulted, unornamented.<br />
Frontal lamina distally broadly rounded, lateral shoulders rounded.<br />
Mouthparts not metamorphosed. Pleotelson anteriorly strongly inflated with<br />
barest indication of two submedian swellings; posterior margin trilobed, with<br />
median lobe strong, narrowly rounded, outer lobes much smaller and ventral<br />
to median lobe. Uropodal endopod distally obliquely truncate; exopod dis<br />
tally acute.<br />
RECORDS Cabanas, Cuba.<br />
REMARKS This species is known only from the nonovigerous female<br />
holotype. Loyola e Silva (1960) placed the species in Cymodoce, based on a<br />
female specimen from Brazil. <strong>As</strong> the mouthparts are not metamorphosed,<br />
this does not agree with the present concept of Cymodoce, but with neither<br />
ovigerous females nor males available, the correct generic placement cannot<br />
be determined.<br />
Cymodoce ruetzleri Kensley, 1984<br />
Figure 99C-G<br />
DIAGNOSIS 6 5.0 mm, 9 4.2 mm. 6: Integument with numerous small<br />
tubercles, becoming densely setose posteriorly. Pleonite 4, posterior margin<br />
broadly bilobed. Pleotelson bearing pair of strong conical tubercles with<br />
acute tips, each tubercle flanked by low rounded tubercle; apex trilobed,<br />
outer lobes triangular, acute, sharp spine at base of incision, median lobe<br />
apically blunt. Uropodal exopod apically acute, oval in cross section, endo<br />
pod and sympod fused, somewhat flattened, apex triangular with strong<br />
tooth. 9: Pleotelson with two conical apically acute tubercles, apex barely<br />
notched, with short rounded lobe slightly offset from posterior margin. Both
228 FLABELLIFERA • SPHAEROMATIDAE<br />
A<br />
Figure 99. "Qymodoce" barrerae: A, $; B, frontal lamina. Cymodoce ruetzleri: C, pleopod 4; G, pleopod 5.<br />
uropodal rami flattened; exopod with tiny apical tooth, endopod distally<br />
truncate-rounded, with small mediodistal tooth.<br />
RECORDS Carrie Bow Cay, Belize, 0.5-13 m; in algal clumps, reef crest<br />
rubble, and seagrass flats.<br />
E
Exosphaeroma Stebbing, 1900<br />
Exosphaeroma alba 229<br />
DIAGNOSIS Maxillipedal palp articles 2-4 produced medially into lobes.<br />
Pereonites 6 and 7 dorsally unarmed. Pleopod 3, exopod biarticulate. 6:<br />
Penes short, separate. Copulatory stylet of pleopod 2 elongate, slender.<br />
Pleotclson lacking strong apical notch. 9: Mouthparts not metamorphosed.<br />
Broodpouch of three pairs of oostegites on pereonites 2-4; oostegites short,<br />
not reaching midline. Brood held in four pairs of internal pouches.<br />
Key to species of Exosphaeroma<br />
1. Pleotelson with posterior margin entire, evenly convex 2<br />
Pleotelson with posterior margin faintly notched or trilobed 3<br />
2. Frontal lamina with length less than 1.5 times greatest width diminuta<br />
Frontal lamina with length almost two times greatest width<br />
productatelson<br />
3. Pleotelson with posterior margin faintly trilobed, and with three low<br />
rounded tubercles anteriorly yucatanum<br />
Pleotelson with posterior margin faintly notched 4<br />
4. Pleotelson posteriorly broadly notched; two rounded submedian<br />
tubercles on inflated midregion antillense<br />
Pleotelson with faint narrow notch posteriorly; lacking dorsal tubercles<br />
Exosphaeroma alba Menzies and Glynn, 1968<br />
Figure 100A-C<br />
DIAGNOSIS 8 2.0 mm, 9 2.3 mm. Frontal lamina anteriorly broadly<br />
rounded, basally slightly wider than midlength. Pleotelson similar in 8 and<br />
9; anterodorsally inflated and unornamented, posteriorly tapering to slight<br />
median notch, seen in dorsal view. Uropodal rami distally shallowly serrate,<br />
exopod 2.5 times longer than wide.<br />
RECORDS Puerto Rico, intertidal to 0.5 m; in algae on rocks, and under<br />
Chiton tuberculatus and C. marmoratus.<br />
alba
230 FLABELLIFERA • SPHAEROMATIDAE<br />
Figure 100. Exosphaeroma alba: A; B, frontal lamina; Cy uropod. Exosphaeroma<br />
antillense: D; E, frontal lamina; F, uropod. Exosphaeroma diminuta: G; H, frontal<br />
lamina; /, uropod. Exosphaeroma productatelson: J; K> frontal lamina; L, uropod.<br />
Exosphaeroma yucatanum: My pleon (from Richardson, 1905).<br />
Exosphaeroma antillense Richardson, 1912d<br />
Figure 100D,F<br />
DIAGNOSIS 9 5.0 mm. Frontal lamina anteriorly tapering to subacute apex.<br />
Pleotelson with two broadly subconical submedian tubercles on inflated an-
Exosphaeroma yucatanum 231<br />
terior area; posterior margin subtruncate to very faintly emarginate. Uropo<br />
dal exopod distally crenulate, length slightly more than twice greatest width;<br />
endopod with faint distal notch.<br />
RECORDS Montego Bay, Jamaica.<br />
REMARKS The single ovigerous female holotype is the only known specimen<br />
of this species. The overlapping oostegites suggest that this may not be an<br />
Exosphaeroma.<br />
Exosphaeroma diminuta Menzies and Frankenberg, 1966<br />
Figure 100G-I<br />
DIAGNOSIS 8 2.2 mm. Frontal lamina widest at midlength, anteriorly<br />
truncate-rounded. Pleotelson with posterior margin broadly rounded.<br />
Uropodal rami not quite reaching pleotelsonic apex; exopod margin distally<br />
crenulate.<br />
RECORDS Chesapeake Bay to Florida; Venezuela; sand dwelling, intertidal<br />
and shallow subtidal.<br />
Exosphaeroma productatelson Menzies and Glynn, 1968<br />
Figure 100J-L<br />
DIAGNOSIS 6 2.5 mm, 9 1.5 mm. Sexes essentially similar. Frontal lamina<br />
widest at midlength, where slight shoulder apparent, anteriorly broadly<br />
rounded, 1.6 times longer than wide. Pleotelson unornamented, anteriorly<br />
inflated, posterior margin entire, evenly convex. Uropodal exopod distally<br />
shallowly serrate, almost four times longer than wide; endopod wider than<br />
exopod. Broad lateral patches of pigment on pleotelson in both sexes.<br />
RECORDS Puerto Rico, intertidal to 0,5 m, in algae on rocks; Texas, Gulf of<br />
Mexico.<br />
Exosphaeroma yucatanum (Richardson, 1901)<br />
Figure 100M<br />
DIAGNOSIS Frontal lamina anteriorly tapering from widest point to sub<br />
acute apex, proximally narrower than at midlength. Pleotelson posteriorly<br />
obscurely trilobed, median lobe narrowly rounded, longest; three low<br />
rounded tubercles on pleotelson in anterior region.
232 FLABELL1FERA • SPHAEROMAT1DAE<br />
RECORDS Cape Catoche, Yucatan, Mexico, 48 m.<br />
REMARKS This species was described from a single specimen which has<br />
since been lost. The true generic placement of this species is thus undeter<br />
mined and full description awaits the finding of more material.<br />
Harrieta Kensley, 1987c<br />
DIAGNOSIS 9 with mouthparts metamorphosed. Broodpouch of three pairs<br />
of oostegites on pereonites 2—4, overlapping in midline; brood held in five<br />
pairs of internal pouches. Uropodal rami subequal, lamellar in 9, exopod<br />
twice length of endopod and oval in cross section in 8. Pleopod 2 in 6 with<br />
copulatory stylet articulating basally on endopod, curved, barely reaching<br />
apex of endopod. Penes basally fused, rami slender, elongate, tapering.<br />
Harrieta faxoni (Richardson, 1905)<br />
Figure 101 A,B<br />
DIAGNOSIS 6 6.0 mm, 9 6.5 mm. 6: Frontal lamina with broad slightly<br />
convex anterior margin. Two low rounded submedian tubercles on cephalon<br />
near posterior margin. Two rounded submedian tubercles on last pleonite.<br />
Pleotelson anteriorly inflated with two submedian tubercles; posterior mar<br />
gin trilobed. 9: Essentially similar to 69 but posterior margin of pleotelson<br />
less markedly trilobed, with median lobe longer, and uropodal rami subequal<br />
in length.<br />
RECORDS Florida to Texas, Gulf of Mexico, intertidal and subtidal in<br />
Thalassia, Halodule, and Syringodium seagrass beds, in salinities of 7%o to 36%o.<br />
Sphaeroma Bosc, 1802<br />
DIAGNOSIS Maxillipedal palp with three distal articles poorly developed,<br />
lacking lobes; fringe of robust plumose setae with swollen bases on internal<br />
margin of endite; distal margin of endite with simple setae. Pereopods 1—3<br />
with plumose setae on ischium and merus. Posterior margin of pleotelson<br />
entire, similar in 6 and 9. Pleopod 3, exopod uniarticulate. Uropodal ex<br />
opod with outer margin serrate. Able to conglobate. 6: Penes short,<br />
rounded, separate. Pleopod 2, copulatory stylet articulating basally on endo<br />
pod, slender, reaching well beyond rami. 9: Mouthparts not meta-
Figure 101. Harrieta faxoni: A, 6; B, pie<br />
Sphaeroma terebrans; E, Sphaeroma walkeri.<br />
B<br />
E
234 FLABELL1FERA • SPHAEROMAT1DAE<br />
morphosed. Three pairs of overlapping oostegites arising from pereonites 2-<br />
4 (but S. terebrans has anterior pair rudimentary).<br />
REMARKS The genus Sphaeroma is one of the few sphaeromatids in which the<br />
number of oostegites varies, from the diagnostic three pairs, through two<br />
normal pairs (as in S. terebrans), to having the oostegites completely absent<br />
(as in S. annandalei).<br />
Jacobs (1987) has provided a useful reevaluation of the European, Medi<br />
terranean, and northwest African species of Sphaeroma and related genera.<br />
Key to species of Sphaeroma<br />
1. Pleotelson posteriorly bluntly triangular, with 4 strong anterior<br />
tubercles<br />
terebrans<br />
Pleotelson posteriorly broadly rounded 2<br />
2. Pleotelson dorsally smooth or with few low tubercles quadridentata<br />
Pleotelson dorsally with numerous strong tubercules walkeri<br />
Sphaeroma quadridentata Say, 1818<br />
Figure 101C<br />
DIAGNOSIS 6 11.0 mm, 9 8.0 mm. Pleotelson anteriorly inflated, some<br />
times with few low rounded tubercles, posteriorly flattened to concave; pos<br />
terior margin entire, broadly rounded.<br />
RECORDS New England to Florida; Gulf of Mexico, intertidal to 1 m, often<br />
in pilings and partially submerged dead tree trunks, and commonly associ<br />
ated with barnacles.<br />
Sphaeroma terebrans Bate, 1866<br />
Figure 10 ID<br />
DIAGNOSIS 6 10.0 mm, 9 11.5 mm. Pereonite 7 with pair of submedian<br />
and pair of lateral tubercles. Dorsal pleon densely tuberculate. Posterior<br />
pleonite with pair of submedian acute tubercles. Pleotelson anteriorly with<br />
submedian pair and lateral pair of tubercles, posteriorly rounded-triangular.
FLABELLIFERA • TRIDENTELLIDAE 235<br />
RECORDS Virginia to Florida; Belize; Cuba; Venezuela to Brazil; Gulf of<br />
Mexico.<br />
Nigeria, east coast of southern Africa, India, Sri Lanka, Thailand, Indo<br />
nesia, Philippines, Australia.<br />
REMARKS There is no agreement on whether this species is synonymous<br />
with S. destructor Richardson, 1897. This latter (if distinct) bores into wood<br />
pilings in estuarine waters, while S. terebrans is found in the prop roots of the<br />
red mangrove tree, Rhizophora mangle. In this habitat, the isopods are inter<br />
preted either as being destructive agents (e.g., Rehm and Humm, 1973) or as<br />
promoting increased root growth (Simberloff et al., 1978). It is unlikely that<br />
the bored wood itself is a source of food for the isopods; rather, as with the<br />
genus Limnoria, the food is probably detritus or fungi and bacteria growing on<br />
the wood fragments in the burrows or on the setae of the appendages.<br />
Sphaeroma walkeri Stebbing, 1905<br />
Figure 101E<br />
DIAGNOSIS 6 9.5 mm, 9 10.0 mm. Pereonites 3—7 with transverse row of<br />
large rounded tubercles. Last pleonite with row of prominent tubercles and<br />
smaller scattered tubercles laterally. Pleotelson anteriorly inflated, posteri<br />
orly concave and cuplike, with four irregular longitudinal rows of large tu<br />
bercles plus many small scattered tubercles. Posterior margin rounded, en<br />
tire to irregularly crenulate. Uropodal endopod with several rounded<br />
tubercles on dorsal surface; exopod with row of smaller tubercles on ventral<br />
surface.<br />
RECORDS Probably pan-tropical. Florida to Puerto Rico, intertidal.<br />
Family Tridentellidae Bruce, 1984<br />
DIAGNOSIS Eyes well developed. Pereonites 2—7 with distinct coxae. Pleon<br />
consisting of five free pleonites plus pleotelson. Mandible with acute incisor;<br />
lacinia mobilis absent; molar present; palp of three articles. Maxilla 1, outer<br />
ramus styliform with three to five strong terminal spines, and several short<br />
recurved subapical spines. Maxilla 2 uniramous, biarticulate, bearing small<br />
sometimes tridentate spines or scales distally. Maxillipedal palp of five arti<br />
cles; endite slender, lamellar, usually lacking coupling hooks.
236 FLABELLIFERA • TRIDENTELLIDAE<br />
Tridentella Richardson, 1905<br />
DIAGNOSIS Body dorsally often bearing spines, tubercles, or carinae, more<br />
developed in 8 than in 9. Frontal lamina narrow, pentagonal. Antcnnular<br />
peduncle of three articles; antennal peduncle of five articles. Mandibular<br />
molar weakly sclerotized. Pereopods 1-3 weakly prehensile; pereopods 4-7<br />
ambulatory. Copulatory stylet of pleopod 2 rodlike, arising proximally on<br />
mesial margin of endopod. Pleopod 5 endopod lacking marginal setae.<br />
REMARKS Delaney and Brusca (1985) provide useful taxonomic and dis<br />
tributional comments on the family Tridentellidae.<br />
Tridentella virginiana (Richardson, 1900b)<br />
Figure 102<br />
DIAGNOSIS 8 9.5 mm, ovigerous 9 9.5-11.0 mm. Cephalon and pereon<br />
dorsally smooth, pleon minutely granular. Uropodal rami with distal mar<br />
gins faintly dentate, apically narrowly rounded, endopod wider and slightly<br />
longer than exopod. Pleotelson basally wider than middorsal length; pos<br />
terior margin broadly rounded to subtruncate.<br />
RECORDS Nova Scotia to Florida; off Georgia, 550 m; Gulf Stream off Key<br />
West, 220 m.<br />
Suborder Gnathiidea Leach, 1814<br />
DIAGNOSIS Eyes usually well developed, rarely on short lateral processes,<br />
occasionally absent. Mandibles in 8 greatly enlarged, projecting anteriorly<br />
from cephalon, not used in feeding. Mandibles lacking in 9. Mouthparts of<br />
praniza larva styliform, with acute mandibles projecting anteriorly (see Fig<br />
ure 103D). Pereopod 1 modified, forming second pair of broad opercular<br />
maxillipeds covering mouthparts, referred to as pylopods. Pereopods 2—6<br />
ambulatory. Pereonite 7 reduced, lacking pereopod. Pleonites separate, nar<br />
rower than pereon. Uropods lateral, rami lamellar, forming tailfan with<br />
telson. Praniza larva with pereonites 4-6 enlarged, sometimes inflated. 9<br />
with pereonites 4-6 greatly inflated, forming broodpouch for internally<br />
brooded eggs (see Figure 103E).<br />
REMARKS The gnathiideans are entirely marine, most described species<br />
being from shallow waters. The males and females are frequently found in<br />
association with sponges and do not feed. The praniza larva is an efficient<br />
swimmer and has been recorded from shallow-water plankton, but is more
GNATHIIDEA 237<br />
Figure 102. Tridentella virginiana: A, 9; B, pereopod 1; C, maxilla 1; D, mandible;<br />
Ey maxilliped; F, maxilla 2.<br />
frequently encountered as a fish parasite, the favored site for sucking the<br />
host's blood being in the nares. Upton (1987a, 1987b) has shed light on the<br />
unusual life history of at least one gnathiid genus, Paragnathia.<br />
The taxonomy of the Gnathiidae is based almost entirely on males, the<br />
praniza and females of most species being remarkably similar.
238 GNATHIIDEA • GNATHIIDAE<br />
Family Gnathiidae Harger, 1879<br />
DIAGNOSIS <strong>As</strong> for the suborder Gnathiidea.<br />
Gnathia Leach, 1814<br />
DIAGNOSIS In addition to features mentioned in diagnosis of suborder: Eyes<br />
present in most species. Pylopod with two small articles distal to broad oper<br />
cular article 2, terminal article minute.<br />
Key to species of Gnathia (6 only)<br />
1. Anterior margin of cephalon with medial process or slightly convex . . 2<br />
Anterior margin of cephalon concave or lacking medial process 9<br />
2. Anterior margin of cephalon broadly triangular, projecting, with small<br />
lateral teeth triospathiona<br />
Anterior margin of cephalon not triangular and projecting 3<br />
3. Cephalon and two free anterior pereonites dorsally granular 4<br />
Cephalon and two free anterior pereonites smooth 5<br />
4. Lobe of outer margin of mandible notched; pereonite 5 twice wider<br />
than middorsal length velosa<br />
Lobe of outer margin of mandible rounded; pereonite 5 1.5 times wider<br />
than middorsal length 6<br />
5. Anterior margin of cephalon with distinct medial process virginalis<br />
Anterior margin of cephalon barely convex, lacking medial process<br />
rat hi<br />
6. Inner proximal lobe of mandible distinct 7<br />
Inner proximal lobe of mandible indistinct samariensis<br />
7. Inner proximal lobe of mandible entire 8<br />
Inner proximal lobe of mandible with rounded toothlike marginal<br />
structures Johanna<br />
8. Pereonites 3-5 poorly defined puertoricensis<br />
Pereonites 3-5 clearly defined magdalenensis<br />
9. Anterior margin of cephalon concave, lacking projections gonzalezi<br />
Anterior margin of cephalon with four projections beethoveni
Gnathia beethoveni Paul and Menzies, 1971<br />
Figure 103 A<br />
Gnathia magdalenensis 239<br />
DIAGNOSIS 6 3.0 mm. Anterior margin of cephalon with two low tubercles<br />
flanking shallow medial notch plus slightly larger pair of lateral tubercles.<br />
Cephalon lacking dorsal tubercles. Pereonite 5 1.5 times wider than middor-<br />
sal length. Uropodal endopods reaching beyond telsonic apex.<br />
RECORDS Off Venezuela, 95 m. Colombia.<br />
Gnathia gonzalezi Miiller, 1988<br />
Figure 103B<br />
DIAGNOSIS 6 2.0 mm. Body smooth. Anterior margin of cephalon concave.<br />
Pereonites 3—5 distinct; pereonite 5 2.5 times wider than middorsal length.<br />
Cutting margin of mandible with four or five low rounded teeth.<br />
RECORDS Colombia, 30 m.<br />
Gnathia Johanna Monod, 1926<br />
Figure 103C<br />
DIAGNOSIS 8 2.1 mm. Anterior margin of cephalon medially convex be<br />
tween pair of submedian tubercles. Pereonites 4 and 5 poorly separated.<br />
Proximomedial lobe of mandible having four or five rounded crenulations,<br />
with tiny seta between adjacent crenulations.<br />
RECORDS U.S. Virgin Islands, 29-46 m; Colombia.<br />
Gnathia magdalenensis Miiller, 1988<br />
Figure 103D<br />
DIAGNOSIS 6 3.0 mm. Anterior margin of cephalon with three tubercles,<br />
median tubercle slightly shorter than submedian pair. Cephalon with few<br />
scattered low granulations dorsally. Pereonite 5 about 1.5 times wider than<br />
middorsal length. Proximomedial lobe of mandible entire.<br />
RECORDS Carrie Bow Cay, Belize, intertidal; Colombia, 18 m.
Figure 103. Gnathia beethhoveni: A, 6. Gnathia gonzalezi: B, 6 (after Miiller, 1988).<br />
Gnathia Johanna: C,
Gnathia puertoricensis Menzies and Glynn, 1968<br />
Figure 103E-G<br />
Gnathia triospathiona 241<br />
DIAGNOSIS 6 3.0 mm, ovigerous $ 1.8 mm. Anterior margin of cephalon<br />
having three tubercles between mandibular bases, medial tubercle narrower<br />
than submedian pair. Dorsal integument finely granular, with coarser gran<br />
ules mediodorsal to eye. Pereonites 4 and 5 indistinctly separated. Mandible<br />
lacking proximomedial lobe.<br />
RECORDS Carrie Bow Cay, Belize, intertidal to 2 m; Puerto Rico, intertidal;<br />
Cuba.<br />
Gnathia rathi Kensley, 1984<br />
Figure 104 A<br />
DIAGNOSIS 6 1.9 mm, ovigerous 9 2.2 mm. Frontal margin faintly convex<br />
to straight between single low lateral tubercle mesial to mandibular bases.<br />
Dorsal integument of cephalon and anterior two free pereonites coarsely<br />
granular. Lateral margins of telson faintly denticulate. Pereonites 4 and 5<br />
poorly separated.<br />
RECORDS Carrie Bow Cay, Belize, 1-36 m.<br />
Gnathia samariensis Miiller, 1988<br />
Figure 104B<br />
DIAGNOSIS 6 2.0 mm. Anterior margin of cephalon with three tubercles,<br />
median tubercle slightly shorter than submedian pair; dorsal integument<br />
smooth. Pereonites 4 and 5 well differentiated; pereonite 5 about 2.2 times<br />
wider than middorsal length. Mandible lacking proximomedial lobe.<br />
RECORDS Colombia.<br />
Gnathia triospathiona Boone, 1918<br />
Figure 104C<br />
DIAGNOSIS 6 8.8 mm. Anterior margin of cephalon with broad-based tri<br />
angular projection bearing three low teeth; deep V-shaped depression pos<br />
terior to anterior margin, with low flanking granulations.<br />
RECORDS Off Key West, in Gulf Stream, 218 m.
Figure 104. Gnathia rathi: A, 6; Gnathia samariensis: B, 6 (after Miiller, 1988);<br />
Gnathia triospathiona; C, 6; Gnathia virginalis: D, 6; Gnathia velosa: E, 6 (after<br />
Miiller, 1988).<br />
E
Gnathia velosa Muller, 1988<br />
Figure 104E<br />
MIGROGERBERIDEA • MIGROGERBERIDAE 243<br />
DIAGNOSIS 6 1.5 mm. Anterior margin of cephalon with three tubercles,<br />
median tubercles slightly shorter and narrower than submedian pair. Dorsal<br />
integument of cephalon and anterior three pereonites granular. Pereonite 5<br />
about 2.5 times wider than middorsal length. Lateral lobe of mandible<br />
notched.<br />
RECORDS Colombia.<br />
Gnathia virginalis Monod, 1926<br />
Figure 104D<br />
DIAGNOSIS 6 2.2 mm. Anterior margin of cephalon with three tubercles,<br />
median tubercles slightly longer than submedian pair. Dorsal integument of<br />
cephalon and anterior three pereonites granular. Pereonite 5 about 1.7 times<br />
wider than middorsal length. Lateral lobe of mandible rounded.<br />
RECORDS U.S. Virgin Islands, 29 m; Colombia.<br />
Suborder Microcerberidea Lang, 1961<br />
DIAGNOSIS Cephalon free. Mandibles with reduced palp, or lacking palp.<br />
Maxillipedal palp of five articles. Pereon of seven free segments. Pereopod 1<br />
subchelate; pereopods 2—7 ambulatory. Pleon of two free pleonites plus<br />
pleotelson. Pleopod 1 in 6 usually absent. Pleopod 2 modified for copulation.<br />
Pleopod 3 uniramous, opercular. Pleopod 4 biramous. Pleopod 5 reduced.<br />
Uropods usually uniramous or biramous.<br />
Family Microcerberidae Karaman, 1933b<br />
DIAGNOSIS Eyes absent. Body elongate, slender. Antennular peduncle of<br />
three articles; antennal peduncle of six to eight articles. Mandibular palp of<br />
single article; molar reduced to single stout fringed spine. Maxilla 2 reduced<br />
to single ramus bearing two distal fringed lobes. Pereopods 2—7 ambulatory,<br />
dactyli biunguiculate.<br />
REMARKS The species of the Microcerberidae are all very small (less than 2<br />
mm total length) and are most often found in interstitial habitats. They have<br />
been recorded from marine, brackish, and freshwater environments.
244 MICROCERBERIDEA • MICROCERBERIDAE<br />
The microcerbcrideans were often classified with the Anthuridea, mainly<br />
because of the similarity in body shape. Wagele (1983) however, has con<br />
vincingly demonstrated the asellotan affinities of the group.<br />
Key to genera of Microcerberidae<br />
1. MaxilHpedal palp articles 2 and 3 enlarged; basis of pereopods lacking<br />
spinous process Yvesia<br />
MaxilHpedal palp articles 2 and 3 not markedly enlarged; basis of<br />
pereopods with spinous process Microcerberus<br />
Microcerberus Karaman, 1933b<br />
DIAGNOSIS MaxilHpedal palp articles 2 and 3 not enlarged. Articles 2 and 3<br />
of antennal peduncle with spinous process. Basis of pereopods with spinous<br />
process. Propodus of pereopod 2 with two denticulate proximal spines.<br />
Microcerberus syrticus Kensley, 1984<br />
Figure 105A-E<br />
DIAGNOSIS (5 1.1 mm, 9 1.1 mm. Tergal lobes of pereonites 2—4 rounded.<br />
Apical lobe of 6 pleopod 2 acute.<br />
RECORDS Carrie Bow Cay, Belize, interstitial in intertidal sand bar.<br />
REMARKS In addition to M. syrticus, six species of Microcerberus have been<br />
recorded from the Caribbean area: M. littoralis Chappuis and Delamare De-<br />
boutteville, 1956, from the Bahamas; M. minutus Coineau and Botosaneanu,<br />
1973, from Cuba; M. mirabilis Chappuis and Delamare Deboutteville, 1956,<br />
from the Bahamas; M. nunezi Coineau and Botosaneanu, 1973, from Cuba;<br />
M. renaudi Chappuis and Delamare Deboutteville, 1956, from the Bahamas;<br />
M. simplex Coineau and Botosaneanu, 1973, from Cuba. The reader is re<br />
ferred to the original descriptions for separation of the species.
\<br />
Figure 105. Microcerberus syrticus: A, 8; B, pereopod 1; C, maxilliped; D, pereopod<br />
2; Ey pleopod 2 8. Yvesia striata (from Coineau and Botosaneanu, 1973): F9<br />
maxilliped; G, pereopod 1; H, pereopod 2.<br />
G
246 MICROCERBER1DEA • ONISCIDEA<br />
Yvesia Coineau and Botosaneanu, 1973<br />
Yvesia striata Coineau and Botosaneanu, 1973<br />
Figure 105F-H<br />
DIAGNOSIS 9 1.6 mm. Antennal peduncular articles 2 and 3 smooth, lack<br />
ing spinous processes. Maxillipedal palp articles 2 and 3 enlarged. Bases of<br />
pereopods unarmed, lacking spinous processes. Propodus of percopod 1 with<br />
single smooth proximal spine. Body having longitudinal ventrolateral striae.<br />
RECORDS Oriente, Cuba, interstitial on beach.<br />
Suborder Oniscidea Latreille, 1803<br />
DIAGNOSIS Compound eyes usually present. Antennules usually very short.<br />
Antennae with 4- or 5-articulate peduncle; flagellum varying from few arti<br />
cles to multiarticulate. Mandibular palp present. Distal articles of maxillipe<br />
dal palp often reduced. Coxae of pereonites 1—7 usually distinct, expanded.<br />
Pleopods respiratory, often with pseudotrachea; 6 with pleopod 2, and<br />
sometimes pleopod 1 as well, modified for copulation. Uropods terminal or<br />
subterminal with terete rami, or ventral and opercular, with reduced rami.<br />
REMARKS The Oniscidea includes all the isopods that have successfully in<br />
vaded the terrestrial environment. While still in some degree reliant on exter<br />
nal moisture, their morphological and behavioral adaptations have allowed<br />
them to live in almost all terrestrial habitats, from hot, dry deserts, through<br />
tropical rainforests and grasslands, to cold-temperate niches. Several forms<br />
have successfully inveigled themselves into termite or ant colonies, where<br />
with varying degrees of morphological adaptations they take advantage of<br />
the security of these habitats. A small number of species have evolved to live<br />
in more constantly wet habitats. Several species may be found in the marine<br />
intertidal, either living in and under piles of decomposing litter along the<br />
high-tide line, digging into beach sand, or sheltering in the damp cracks and<br />
crevices of rocky shores. A few may also be found in mangrove swamps.<br />
A breakdown of families, genera, and species is not provided for this sub<br />
order, but those few species that are commonly encountered in intertidal<br />
habitats are dealt with individually. Schultz (1974, 1984) records several<br />
oniscidean isopods from the Caribbean area.
Key to genera and species of littoral Oniscidea<br />
ONISCIDEA 247<br />
1. At least one uropodal ramus reaching well beyond outline of body ... 5<br />
Uropodal rami very short, not reaching beyond outline of body 2<br />
2. Uropods ventral, not visible in dorsal view 3 (Tylos)<br />
Uropods visible in dorsal view Armadilloniscus ninae<br />
3. Ventral extensions of pleonite 5 meeting in midline Tylos niveus<br />
Ventral extensions of pleonite 5 not meeting in midline 4<br />
4. Ventral extensions of pleonite 5 very short, obsolete Tylos wegeneri<br />
Ventral extensions of pleonite 5 well separated Tylos marcuzzii<br />
Ventral extensions of pleonite 5 just falling short of meeting in midline<br />
Tylos latreillei<br />
5. Uropodal rami both elongate, subequal 6 (Ligia)<br />
Uropodal rami very unequal in length 8<br />
6. Propodus of 6 pereopod 1 with distal rounded lobe Ligia exotica<br />
Propodus of 6 pereopod 1 lacking rounded lobe 7<br />
7. Apex of 6 pleopod 2 club shaped Ligia olfersii<br />
Apex of 6 pleopod 2 bifid Ligia baudiniana<br />
8. Antennal flagellum of two articles Rhyscotus texensis<br />
Antennal flagellum of three articles 9 (Vandeloscia)<br />
9. Endopod of 6 pleopod 1 with large scalelike subapical process<br />
Endopod of 6 pleopod 1 with small scalelike subapical process<br />
Armadilloniscus Ul'yanin, 1875<br />
Armadilloniscus ninae Schultz, 1984<br />
Figure 106A<br />
Vandeloscia riedli<br />
Vandeloscia culebrae<br />
- - - - - ,<br />
DIAGNOSIS 8 3.2 mm, 9 4.1 mm. Uropodal sympod expanded to form part<br />
of body outline; rami set mesial to expanded base, with exopod half length of<br />
endopod.<br />
RECORDS Ambergris Cay, Belize; under damp objects along beach drift<br />
line.
Figure 106. A, Armadilloniscus ninae. Ligia baudiniana: B; C} 6 pleopod 2 endopod.<br />
Ligia exotica: D, dactylus and propodus of pereopod 1; E, 6 pleopod 2 endopod.<br />
Ligia olfersii: F, 6 pleopod 2 endopod. G, Rhyscotus texensis. Tylos latreillei: H,<br />
ventral pleon. Tylos marcuzzi: I, ventral pleon (from Schultz, 1984). Tylos niveus: J,<br />
lateral view; K} ventral pleon. Tylos wegeneri: L} ventral pleon. Vandeloscia culebrae:<br />
M, apex of pleopod 1 endopod. N, Vandeloscia riedli.
Ligia Fabricius, 1798<br />
Ligia baudiniana H. Milne Edwards, 1840<br />
Figure 106B,C<br />
Rhyscotus texensis 249<br />
DIAGNOSIS 6 and 9 up to 22 mm. Antennal flagellum elongate, multiar-<br />
ticulate. Apex of 6 pleopod 2 bifid, with lateral lobe longer and more slender<br />
than mesial lobe. Uropods inserted terminally on pleotelson; sympods<br />
elongate-cylindrical; rami slender, elongate, subequal.<br />
RECORDS Bermuda; Bahamas; U.S. Virgin Islands; Antigua; Carrie Bow<br />
Cay, Belize; Bonaire; Aruba; Trinidad; Tobago; Gulf of Mexico.<br />
REMARKS <strong>As</strong> is typical in the genus Ligia, this species may be seen on rocks<br />
i<br />
and sea walls, as well as piles of drift debris at low tide. When disturbed, they<br />
run rapidly, to shelter in damp crevices and hollows.<br />
Ligia exotica Roux, 1828<br />
Figure 106D,E<br />
DIAGNOSIS 6 28.5 mm, ovigerous 9 32.0 mm. Propodus of 8 pereopod 1<br />
with rounded lobe on inner distal surface. Apex of 6 pleopod 2 club shaped,<br />
convoluted.<br />
RECORDS New Jersey to Uruguay; Indo-Pacific.<br />
Ligia olfersii Brandt, 1833<br />
Figure 106F<br />
DIAGNOSIS 6 20.0 mm, ovigerous 9 24.0 mm. Apex of 6 pleopod 2 simple,<br />
club shaped.<br />
RECORDS South Florida to Rio de Janeiro, Brazil; Texas, Gulf of Mexico.<br />
Rhyscotus Budde-Lund, 1885<br />
Rhyscotus texensis (Richardson, 1905)<br />
Figure 106G<br />
DIAGNOSIS 6 and 9 6.0 mm. Antennal flagellum of two unequal articles.<br />
Uropodal endopod at least twice length of exopod, inserted distally on base,<br />
exopod inserted distally on base. Pleotelson broadly triangular.<br />
RECORDS Carrie Bow Cay, Belize; Texas, Gulf of Mexico.
250 ONISCIDEA<br />
Tylos Latreille, 1826<br />
Tylos latreillei Audouin, 1826<br />
Figure 106H<br />
DIAGNOSIS 6 12.8 mm, 9 13.0 mm. Ventral extensions of pleonite 5 not<br />
meeting in midline.<br />
RECORDS Bermuda; Cuba; Puerto Rico; Honduras.<br />
Mediterranean.<br />
Tylos marcuzzii Soika, 1954<br />
Figure 1061<br />
DIAGNOSIS 6 6.6 mm. Antennal flagellum of four articles. Ventral exten<br />
sions of pleonite 5 well separated.<br />
RECORDS Florida Keys; Bahamas; Leeward Islands; Ambergris Cay, Be<br />
lize; under debris on sand beach drift line.<br />
Tylos niveus Budde-Lund, 1885<br />
Figure 106J,K<br />
DIAGNOSIS 8 11.0 mm., 9 12.0 mm. Antennal flagellum of four articles.<br />
Ventral extensions of pleonite 5 expanded, medially contiguous.<br />
RECORDS Bahamas; Florida Keys; Cuba; Dominica; Lesser Antilles; Bo<br />
naire; Curasao, under piles of decaying mangrove leaves at beach drift line;<br />
Carrie Bow Cay, Ambergris Cay, Belize, under deep piles of dead plant ma<br />
terial on beach drift line; Tobago; Panama.<br />
Rio de Janeiro, Brazil.<br />
Tylos wegeneri Vandel, 1952<br />
Figure 106L<br />
DIAGNOSIS 6 10.5 mm, 9 15 mm. Antennal flagellum of three articles.<br />
Ventral extensions of pleonites short or nearly absent. Pleonite 5 lacking free<br />
lateral margins.<br />
RECORDS Tobago; Venezuela, under decaying beach debris on drift line;<br />
Trinidad.
Vandeloscia Roman, 1977<br />
Vandeloscia culebrae (Moore, 1901)<br />
Figure 106M<br />
VALVIFERA 251<br />
DIAGNOSIS 8 5.0 mm, 9 6.1 mm. Tiny lateral tubercles present on per<br />
eonites. Endopod of pleopod 1 in 8 with small scalelike subapical process on<br />
laterally folded tip.<br />
RECORDS Florida Keys; U.S. Virgin Islands; Puerto Rico; under decaying<br />
plant material, especially Thalassia testudinea accumulated along beach drift<br />
line,<br />
Vandeloscia riedli (Strouhal, 1966)<br />
Figure 106N<br />
DIAGNOSIS 8 5.9 mm, 9 6.0 mm. Tiny obsolete tubercles present on all<br />
pereonites. Endopod of 8 pleopod 1 with large scalelike subapical process on<br />
laterally folded tip.<br />
RECORDS Yucatan Peninsula, Mexico; Ambergris Cay, Belize; Barbuda;<br />
Venezuela; Brazil.<br />
Gulf of Aqaba; Red Sea; northeastern coast of Africa; Madagascar; Bay of<br />
Bengal; St. Helena Is.<br />
Suborder Valvifera Sars, 1882<br />
DIAGNOSIS Pereopodal coxae, in addition to usual dorsal coxal plates, ex<br />
panded ventrally to form plates. Penes situated ventrally on articulation be<br />
tween pereon and pleon, or on pleonite 1. Pleonites and pleotelson variously<br />
fused. Uropods forming operculum covering over pleopods.<br />
Key to families of Valvifera<br />
1. Body often geniculate, flexed between pereonites 4 and 5; anterior<br />
pereopods setose for feeding, posterior pereopods ambulatory<br />
Arcturidae<br />
Body never geniculate; all pereopods ambulatory Idoteidae
252 VALV1FERA • ARGTUR1DAE<br />
REMARKS Of the six families in the suborder, only two have been recorded<br />
in the Caribbean area, the Idoteidae and the Arcturidae.<br />
Family Arcturidae Sars, 1897<br />
DIAGNOSIS Pereonite 1 either distinct, or completely or imcompletely fused<br />
with cephalon. Anterior four pairs of pereopods directed anteriorly, usually<br />
strongly setose; posterior three pairs of pereopods ambulatory, used for cling<br />
ing to substrate. Body often bent between pereonites 4 and 5. Uropods usu<br />
ally biramous, with minute endopod concealed by larger exopod. Pleonites<br />
variously fused with pleotelson. Sexual dimorphism often marked.<br />
REMARKS Menzies and Kruczynski (1983) described three species of<br />
arcturids from the west coast of Florida, in depths of 55—73 m: Arcturella<br />
spinata, Arcturella bispinata, and Edwinjoycea horologium. These species are not<br />
covered here.<br />
Key to genera of Arcturidae<br />
1. Pereonite 1 not fused with cephalon; at least one free pleonite<br />
Pereonite 1 fused with cephalon; pleonites fused with pleotelson<br />
<strong>As</strong>tacilla Cordiner, 1793<br />
Thermarcturus<br />
<strong>As</strong>tacilla<br />
DIAGNOSIS Antennae at least half length of body. Pereopod 1 with strong<br />
terminal claw on dactylus. Pereopods 2—4 lacking dactyli. Endopod of<br />
Key to species of <strong>As</strong>tacilla<br />
1. Body integument lacking ornamentation cymodocea<br />
Body integument with spines or tubercles 2<br />
2. Pereonite 4 in 6 and 9 with strong middorsal tubercle; pairs of spines<br />
lacking on pereonites regina<br />
Pereonite 4 lacking strong middorsal tubercle; pairs of spines on all<br />
pereonites lasallae
<strong>As</strong>tacilla regina 253<br />
pleopod 1 6 with median notch and three specialized setae; pleopod 2 copu-<br />
latory stylet apically trifid. Pereonite 4 considerably longer than preceeding<br />
or following pereonite.<br />
<strong>As</strong>tacilla cymodocea Menzies and Glynn, 1968<br />
Figure 107A,B<br />
DIAGNOSIS 6 6.4 mm, ovigerous 9 9.0 mm. Body cylindrical, ovigerous 9<br />
with pereonite 4 somewhat bulged, 6 with pereonite 4 elongate-cylindrical.<br />
Shallow groove marking fusion between cephalon and pereonite 1. Pleonites<br />
fused with pleotelson, with two incomplete shallow dorsal grooves marking<br />
lines of fusion anteriorly. Pleotelson lacking any shoulders, posteriorly tape<br />
red to narrowly rounded apex.<br />
RECORDS Florida Keys; Puerto Rico, 1.5 m, on Cymodocea sp. seagrass; Car<br />
rie Bow Cay, Belize, 1—2 m, on Syringodium filiforme seagrass.<br />
REMARKS In life, A. cymodocea is bright green, blending in with its preferred<br />
substrate of seagrasses.<br />
<strong>As</strong>tacilla lasallae Paul and Menzies, 1971<br />
Figure 107C<br />
DIAGNOSIS 9 3.5 mm. Cephalon with large rounded area bearing pair of<br />
spines; all pereonites and two anterior fused pleonites bearing pair of short<br />
submedian spines. Pleotelson with strong anterior shoulder, posteriorly tri<br />
angular, tapering sharply to narrowly rounded apex.<br />
RECORD Off Venezuela, 95 m.<br />
REMARKS This species is known only from the small female holotype, and<br />
until a mature male and ovigerous female are found, it cannot be confidently<br />
diagnosed.<br />
<strong>As</strong>tacilla regina Kensley, 1984<br />
Figure 107D-G<br />
DIAGNOSIS 6 6.5 mm, ovigerous 9 7.1 mm. Body strongly tuberculate,<br />
many tubercles acute. Cephalon with two submedian pairs of acute tuber<br />
cles; fused pereonite 1 and pereonites 2 and 3 each with single middorsal<br />
acute tubercle. Pereonite 4 with strong middorsal tubercle situated in ante-
254 VALVIFERA • ARGTURIDAE<br />
Figure 107. <strong>As</strong>tacilla cymodocea: A, 8; By 9. <strong>As</strong>tacilla lasallae: C, 6. <strong>As</strong>tacilla regina:<br />
D, 6\ E, 9; F, pereopod 4; G, pereopod 1. Thermarcturus venezuelensis: H, 9 (from<br />
Paul and Menzies, 1971).<br />
rior half. Pleotelson with strong lateral shoulder in anterior half, second<br />
shoulder in posterior half, then tapering to rounded apex.<br />
RECORDS Carrie Bow Cay, Belize, on forereef slope, 27—36 m; Barbados,<br />
100—400 m; St. Lucia, 2—3 m, associated with crinoids.
Thermarcturus Paul and Menzies, 1971<br />
VALVIFERA • IDOTEIDAE 255<br />
DIAGNOSIS Pereonite 1 not fused with cephalon. Pereonite 4 subequal in<br />
length to pereonite 3, not markedly elongate. Pereopods 2—4 having dactyli<br />
but lacking elongate setae. Body cylindrical, flexed between pereonites 4 and<br />
5. Pleon consisting of two free pleonites plus pleotelson.<br />
Thermarcturus venezuelensis Paul and Menzies, 1971<br />
Figure 107H<br />
DIAGNOSIS 9 4.5 mm. Cephalon, all pereonites, and anterior two pleonites<br />
each with submedian pair of dorsal tubercles, those on pereonites 2 and 3<br />
broad and expanded. Pleonite 2 with pair of bulbous lateral swellings, pos<br />
terior margin triangular. Pleotelson with lateral shoulder anteriorly, posteri<br />
orly triangular.<br />
RECORDS Off Venezuela, 95 m.<br />
REMARKS Only the holotype (which seems to be lost) is known of this spe<br />
cies. Considerable uncertainty exists regarding some of the features.<br />
Family Idoteidae Fabricius, 1798<br />
Subfamily Idoteinae Dana, 1852<br />
DIAGNOSIS Flagellum of antenna either multiarticulate; clavate, i.e., with<br />
large basal articles and with or without one to four reduced distal articles; or<br />
Key to genera of Idoteinae<br />
1. Antennal flagellum multiarticulate Idotea<br />
Antennal flagellum clavate 2<br />
2. Pereopod 4 reduced, considerably smaller than pereopods 3 or 5 .... 3<br />
Pereopod 4 not reduced, of similar size to pereopods 3 and 5<br />
Erichsonella<br />
3. Pleon consisting of three complete and one incomplete pleonites plus<br />
pleotelson Cleantioides<br />
Pleon consisting of two complete and two incomplete pleonites plus<br />
pleotelson Miratidotea
256 VALVIFERA • IDOTEIDAE<br />
vestigial. Maxillipedal palp consisting of five or fewer articles. Uropods uni-<br />
ramous or biramous, rami usually much smaller than sympod. Pleonites<br />
variously fused with pleotelson; number of fused pleonites often indicated by<br />
lateral sutures or furrows.<br />
REMARKS Brusca (1984) has reviewed the phylogeny, evolution, and bio-<br />
geography of the subfamily Idoteinae, the only one of the five subfamilies<br />
recorded from the Caribbean.<br />
Cleantioides Kensley and Kaufman, 1978<br />
DIAGNOSIS Antennal flagellum a single clavate article. Maxillipedal palp of<br />
four or five articles. Pereopod 4 somewhat reduced. Uropod uniramous.<br />
Pleon consisting of three complete and one incomplete pleonites plus<br />
pleotelson.<br />
Cleantioides planicauda (Benedict, 1899)<br />
Figure 108A<br />
DIAGNOSIS Ovigerous 9 5.5 mm. Body parallel sided. Maxillipedal palp of<br />
five articles. Pleotelson posteriorly broadly rounded, with obliquely truncate<br />
subcircular dorsal area in posterior half.<br />
RECORDS Maryland to Florida; Puerto Rico; Panama; Louisiana, Gulf of<br />
Mexico, intertidal to 44 m; often in hollow stems and roots of seagrasses, and<br />
tubes of the polychaete Diopatra cuprea.<br />
Oaxaca, Pacific Mexico.<br />
REMARKS Cleantioides planicauda has been recorded only once in the eastern<br />
Pacific, where it occurs with the more common C. occidentalis (Richardson,<br />
1899).<br />
Key to species of Erichsonella<br />
1. Pereonites with dorsal spines 2<br />
Pereonites lacking dorsal spines attenuata<br />
2. Pereonites 1—4 with middorsal and lateral spines floridana<br />
Pereonites 1-4 with middorsal spines only Jiliformis
Erichsonella 257<br />
Figure 108. Ay Cleantioides planicauda; B, Erichsonella attenuata 8\ C, Erichsonella<br />
filiformis 6\ Dy Erichsonella Jloridana 9; E, Idotea balthica 6] F, Idotea metallica $; G,<br />
Miratidotea bruscai $.<br />
Erichsonella Richardson, 1901<br />
DIAGNOSIS Antennal flagellum clavate. Maxillipedal palp of four articles.<br />
Uropod uniramous. Pleonites completely fused with pleotelson.<br />
REMARKS Pires (1984) reviewed the genus Erichsonella and did not recog<br />
nize the subspecies E. filiformis tropicalis Menzies and Glynn, 1968.
258 VALVIFERA • IDOTEIDAE<br />
Erichsonella attenuata (Harger, 1873)<br />
Figure 108B<br />
DIAGNOSIS 6 11.4 mm, ovigerous 9 12.0 mm. Body dorsally smooth.<br />
Cephalon lacking middorsal elevation. Antcnnulc reaching only slightly be<br />
yond antennal peduncular article 2. Pleotelson with slight marginal bulge in<br />
anterior half, indicating ventrolateral articulation of uropod.<br />
RECORDS Connecticut to Miami; Florida, Mississippi, Texas, Gulf of Mex<br />
ico; intertidal to 2 m, usually associated with submerged seagrass and algal<br />
beds.<br />
REMARKS While not recorded in the Florida Keys, this species does reach<br />
Miami, and continues into the Gulf of Mexico.<br />
Erichsonella filifor mis (Say, 1818)<br />
Figure 108C<br />
DIAGNOSIS 6 10.5 mm, ovigerous 9 8.2 mm. Body dorsally with bifid tu<br />
bercle on cephalon, and low rounded middorsal tubercle on pereonites. An-<br />
tennule reaching midlength of antennal peduncular article 3. Basis of per-<br />
eopods 2—7 with larges tubercles. Pleotelson with distinct lateral shoulder in<br />
anterior half.<br />
RECORDS Connecticut to Florida, shallow infratidal to 55 m; Bahamas;<br />
Turks and Caicos Islands; Puerto Rico; Quintana Roo, Yucatan Peninsula,<br />
Mexico, 60-109 m; Florida and Texas, Gulf of Mexico.<br />
Brazil.<br />
Erichsonella jloridana Richardson, 1901<br />
Figure 108D<br />
DIAGNOSIS Ovigerous 9 10.0 mm. Antennule reaching distal end of anten<br />
nal peduncular article 3. Cephalon with strong trifid tubercle. Pereonites 1—7<br />
each with posteriorly directed spine near posterior margin; pereonites 1—4<br />
each with lateral spine. Basis of pereopods 2-7 smooth.<br />
RECORDS Florida Keys, intertidal to 2 m; Florida, Gulf of Mexico, interti<br />
dal mud flats.
Idotea Fabricius, 1798<br />
Miratidotea 259<br />
DIAGNOSIS Antennal flagellum multiarticulate. Maxillipedal palp of four or<br />
five articles. Uropod uniramous. Pleon consisting of two complete and one<br />
incomplete pleonites plus pleotelson.<br />
Key to species of Idotea<br />
1. Posterior margin of pleotelson truncate metallica<br />
Posterior margin of pleotelson with distinct median lobe balthica<br />
Idotea balthica (Pallas, 1772)<br />
Figure 108E<br />
DIAGNOSIS 6 24.5 mm, ovigerous 9 13.2-23.5 mm. Anterior margin of<br />
cephalon concave. Cephalon dorsally smooth. Pereonites evenly convex,<br />
smooth. Posterior margin of pleotelson with rounded median lobe.<br />
RECORDS Worldwide in tropical to cold-temperate waters, often on floating<br />
seaweed, from surface to 357 m.<br />
Idotea metallica Bosc, 1802<br />
Figure 108F<br />
DIAGNOSIS 6 30.0 mm, ovigerous 9 22.2 mm. Cephalon with sinuous fur<br />
row in posterior half. Pereonites 2—4 laterally with rounded convex area close<br />
to coxae. Posterior margin of pleotelson truncate.<br />
RECORDS Worldwide in tropical to cold-temperate waters, often on floating<br />
seaweed, from surface to 200 m.<br />
Miratidotea Kensley, 1987a<br />
DIAGNOSIS Antennal flagellum of single clavate article. Maxillipedal palp<br />
of four articles. Uropod uniramous. Pleon consisting of two complete and two<br />
incomplete pleonites plus pleotelson.
260 VALVIFERA • IDOTEIDAE<br />
Miratidotea bruscai Kensley, 1987a<br />
Figure 108G<br />
DIAGNOSIS Ovigerous 9 13.0 mm. Body parallel sided. Maxillipedal palp<br />
of four articles, terminal article very short. Pereopods 1—3 increasing in<br />
length posteriorly, pereopod 4 reduced, shorter than pereopod 5, and with<br />
dactylus spinelike, pereopods 5—7 increasing in length. Pleotelson consisting<br />
of two complete and two incomplete pleonites plus pleotelson; latter with<br />
broadly rounded posterior margin, and with bifid median process situated<br />
dorsal to posterior oblique-concave area.<br />
RECORDS Carrie Bow Cay, Belize, 1.5 m, in hollow root-internodes of sea-<br />
grass Syringodium filiforme.
Zoogeography<br />
FAUNAL PROVINCES<br />
The area under discussion has been divided into several faunal regions or<br />
provinces, of which the Caribbean, West Indian, and Brazilian are the major<br />
ones (Briggs, 1974). The extent and boundaries of the provinces have been<br />
variously defined depending on the group of organisms under discussion.<br />
Inevitably, zones of overlap exist, but for the purposes of this discussion, the<br />
following rough limits have been used.<br />
Brazilian Province: This province stretches from Cape Frio near Rio de<br />
Janeiro in Brazil to the mouth of the Orinoco River in Venezuela. The out<br />
flow of freshwater from the major rivers of this region has probably contrib<br />
uted to the isolation of the Brazilian coral reefs and their associated fauna<br />
from those of the Caribbean. This isolation is demonstrated by the consider<br />
able endemism of the Brazilian reef fauna and that of the Caribbean reef<br />
fauna, with very few species being common to both.<br />
Caribbean Province: This province has two components, a northern part<br />
in peninsular Florida, that stretches from around Cape Kennedy on the east<br />
coast to Tampa or Sanibel Island on the west coast, and a southern compo<br />
nent that runs from the mouth of the Orinoco River to around Cabo Rojo or<br />
Tampico on the gulf coast of Mexico. The northern Gulf of Mexico is ex<br />
cluded from this province and is characterized as being warm-temperate,<br />
rather than subtropical (Briggs, 1974:66).<br />
West Indian Province: This includes all the islands of the West Indian<br />
chain, the Bahamas, and the isolated outrider, Bermuda. The West Indian<br />
Province closely approaches the Caribbean Province in the Yucatan Penin<br />
sula to the north, and between Grenada and Trinidad in the south. There is<br />
also some indication of the isolating effect on the Bahamas of the Florida<br />
Current through the Straits of Florida.<br />
It has been suggested, on the basis of the molluscan fauna, that a relict of<br />
the Neogene Gatunian Province exists around northern Venezuela and Col<br />
ombia (Petuch, 1982). While several isopod species have been recorded only<br />
from this area, these are all described in a single paper that covers a very<br />
small part of this region (Paul and Menzies, 1971). There is as yet too little<br />
evidence to explore the idea of this relict fauna further.<br />
261
262 ZOOGEOGRAPHY<br />
ANALYSIS OF THE ISOPOD FAUNA<br />
In the following discussion, the West Indian and Caribbean provinces are<br />
treated as one, the isopod faunas offering little evidence to warrant a separate<br />
treatment of each.<br />
It is a truism that for any discussion of the zoogeography of an area to have<br />
meaning, the true extent of the fauna must be known. With the area under<br />
review, collecting effort has been uneven, and the true faunal composition of<br />
many regions is still incompletely known. Obviously, any conclusions based<br />
on such incomplete data are approximate and subject to revision. Neverthe<br />
less, certain general patterns or trends emerge when the present isopod fauna<br />
is broken down into its components.<br />
The deepwater isopod fauna of the Caribbean (i.e., from deeper than 200<br />
m) has barely been explored, and little is to be gained from discussing the<br />
relatively few species known. A list of these deeper dwelling species is in<br />
cluded (Table 4).<br />
Although about 280 shallow-water species are covered by this work, cer<br />
tain categories of species must be excluded, for various reasons, before anal<br />
ysis can be attempted. Such excluded groups include the species of Oniscidea<br />
(being essentially terrestrial forms and not part of the marine regime); the<br />
cymothoid species and the species of Aegidae (being fish parasites for at least<br />
part of their life history, and whose distribution is complicated by the dis<br />
tribution and mobility of the hosts); the limnoriids (being wood-borers whose<br />
distribution is more a function of the distribution of floating wood); and the<br />
true cave species (which have a history more reflective of the geological his<br />
tory of the area than of the marine regime). The epicarideans have a distribu<br />
tion somewhat complicated by the distribution of their crustacean hosts and<br />
their pelagic epicaridean and cryptoniscan larvae. Nevertheless, the decapod<br />
hosts of the great majority of species covered here are Caribbean endemics,<br />
and inclusion of the epicarideans changes very little the overall patterns of<br />
distribution, as demonstrated by the two figures provided (Figure 109). After<br />
making these exclusions there remain about 166 species (218 with the epi<br />
carideans) that can be broken down into the following components (figures<br />
in brackets include epicarideans):<br />
1. True Caribbean/Bahamian species—124 species, 74.8% [147, 67.5%].<br />
These are the species recorded only from the Caribbean and the Bahamas.<br />
The term endemic is avoided, as too little is known of the actual distribution<br />
of many species. Of these species, 86 [87] have been recorded from a single<br />
locality.<br />
2. Species occurring south of the discussion area, and extending into<br />
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<br />
SUBORDER ANTHURIDEA<br />
Family Paranthuridae<br />
Neoanthura coeca Menzies, 1956b. South of Jamaica, 1244 m<br />
SUBORDER ASELLOTA<br />
Family Dendrotiidae<br />
Dendrotion hanseni Menzies, 1956b. South of Jamaica, 1244 m<br />
Family Desmosomatidae<br />
Desmosoma magnispina Menzies, 1962a. Bay of Panama, 1906 m<br />
Family Echinothambematidae<br />
Echinothambema ophiuroides Menzies, 1956a. North of Puerto Rico Trench,<br />
5104-5122 m<br />
Family Eurycopidae<br />
Acanthocope spinosissima Menzies, 1956b. South of Jamaica, 1224 m<br />
Storthyngura pulchra caribbea (Benedict, 1901). Off Windward Islands, 1256 m<br />
Storthyngura snanoi Menzies, 1962a. Colombia abyssal plain, 4071 m<br />
Family Haploniscidae<br />
Antennuloniscus dirneroceras (Barnard, 1920). North of Puerto Rico Trench,<br />
5440-5410 m; South Atlantic off South and West Africa, 1400-3921 m;<br />
off Argentina, 5843 m<br />
Haploniscus unicornis Menzies, 1956a. North of Puerto Rico Trench, 5104-<br />
5122 m<br />
Hydroniscus quadrifrons Menzies, 1962a. North of Puerto Rico Trench, 5271-<br />
5684 m<br />
Family Ischnomesidae<br />
Haplomesus tropicalis Menzies, 1962a. Colombia abyssal plain, 4071 m; off<br />
South Africa, 2526 m; Mediterranean<br />
Heteromesus bifurcatus Menzies, 1962a. Colombia abyssal plain, 4071 m<br />
Ischnomesus armatus Hansen, 1916. North of Puerto Rico Trench, 5494-5477<br />
m; Davis Straits, 2702 m<br />
Ischnomesus caribbicus Menzies, 1962a. Off Panama, 1714 m<br />
Ischnomesus multispinis Menzies, 1962a. Off Panama, 975 m<br />
Family Janiridae<br />
Abyssianira dentifrons Menzies, 1956a. North of Puerto Rico Trench, 5104-<br />
5122 m; off Argentina, 5024-5293 m; off southwest Africa, 4588 m<br />
Ianirella caribbica Menzies, 1956b. South of Jamaica, 1244 m<br />
Ianirella vemae Menzies, 1956a. Near Puerto Rico Trench, 5104-5122 m<br />
Spinianirella serrata Kensley and Heard, 1985. Off Puerto Rico, 350 m<br />
Family Macrostylidae<br />
Macrostylis caribbicus Menzies, 1962a. Off Colombia, 2875-2941 m<br />
Macrostylis minutus Menzies, 1962a. North of Puerto Rico Trench, 5163-<br />
5494<br />
{continued)
264 ZOOGEOGRAPHY<br />
TABLE 4. {Continued)<br />
Macrostylis setifer Menzies, 1962a. North of Puerto Rico Trench, 5477-<br />
5494 m<br />
Macrostylis vemae Menzies, 1962a. North of Puerto Rico Trench, 5410—<br />
5684 m<br />
Family Mesosignidae<br />
Mesosignum kohleri Menzies, 1962a. Colombia abyssal plain, 2868-4076 m<br />
Family Nannoniscidae<br />
Nannoniscus camayae Menzies, 1962a. Off Panama, 1714 m<br />
SUBORDER GNATH11DEA<br />
Family Gnathiidae<br />
Akidognathia poteriophora Monod, 1926. OffU. S. Virgin Islands, 914 m.<br />
SUBORDER VALVIFERA<br />
Family Arcturidae<br />
Antarcturus annaoides Menzies, 1956b. South of Jamaica, 1244 m<br />
Arcturus caribbaeus Richardson, 1901. OfTAves Island, 1360 m<br />
Arcturus purpureus Beddard, 1886. Off Leeward Islands, 900 m<br />
Note: Records from deep water around Bermuda are not included.<br />
great area of mixed-salinity waters resulting from the outflow of the Orinoco,<br />
Amazon, Tocantins, and Parnaiba rivers form an effective barrier to the<br />
movement of shallow-water isopod species.<br />
3. Species having an amphi-Panamic distribution—7 species, 4.2% [8,<br />
3.7%] (Table 5). In spite of the history of immergence and emergence of the<br />
Isthmus of Panama, this very small amphi-Panamic component in the Carib<br />
bean isopod fauna suggests that most of this fauna has evolved since the last<br />
emergence of the late Pliocene. Given the limited mobility of most isopod<br />
species, the Panama Canal seems to have played a minimal role in contribut<br />
ing to this component.<br />
4. Species occurring outside of the western Atlantic (but excluding the<br />
amphi-Panamic species)—3 species, 1.8% [7, 3.2%].<br />
5. The role of the Gulf of Mexico isopod fauna (see Clark and Robertson,<br />
1982) in the composition of the Caribbean/Bahamian is complex and diffi<br />
cult to analyze. One hundred and thirteen species of shallow-water isopods<br />
have been recorded from the Gulf of Mexico (Table 6). This number would<br />
indicate that many species remain to be recorded in this region. Of these 113<br />
species, 61 (54%) have also been reported from the Caribbean region. It is<br />
therefore possible that there exists a true Gulf of Mexico fauna, whose evolu<br />
tion was perhaps spurred by the relative isolation and reduction of the Gulf
Car<br />
Car<br />
Sout<br />
South 3.0% %<br />
Panam 4.2%<br />
Epicaridea excluded<br />
Panam 3.7%<br />
Out 3.2%<br />
North<br />
Epicaridea included<br />
5.5%<br />
ZOOGEOGRAPHY 265<br />
GoM/N/Car 7.2%<br />
oM/Car 5.4%<br />
3.6%<br />
GoM/N/Car 9.6%<br />
/Car 6.4%<br />
Figure 109. Relative proportions of the zoogeographic components of the<br />
Caribbean isopod fauna, with and without the parasitic Epicaridea. Car,<br />
Caribbean; Out, extra-western Atlantic; GoM/Car, Gulf of Mexico-Caribbean;<br />
GoM/N/Car, Gulf of Mexico-Northern-Caribbean; North, northern; Panam, amphi-<br />
Panamic; South, southern.<br />
during a low-water stand (100 m below present sea level) during the<br />
Pleistocene. A significant proportion (about 27 species, 28%) of the Gulf of<br />
Mexico isopods are known from the eastern coast of the United States north
266 ZOOGEOGRAPHY<br />
TABLE 5. SPECIES OF ISOPODS OCCURRING ON BOTH SIDES OF THE ISTHMUS<br />
OF PANAMA<br />
*Aega deshaysiana (H. Milne Edwards,<br />
1840)<br />
Anopsilana browni (Van Name, 1936)<br />
Cleantioides planicauda (Benedict,<br />
1899)<br />
Excirolana braiiliensis Richardson,<br />
1912<br />
Excorallana tricornis (Hansen, 1890)<br />
*Nerocila acuminata Schioedte and<br />
Meinert, 1881<br />
* fish parasite or fish predator<br />
of Cape Kennedy, which would indicate a significant cooler-water compo<br />
nent. What proportion of originally Gulf species have spread into the Carib<br />
bean, and what proportion of Caribbean and temperate east coast species<br />
have entered the Gulf, cannot yet be assessed, given our incomplete knowl<br />
edge of the Gulf fauna. Because of this unresolved situation, three categories<br />
of species have been separated: species ranging from north of Cape Kennedy<br />
into the Caribbean—6, 3.6% [12, 5.5%]; species occurring in the Gulf of<br />
Mexico and the Caribbean—9, 5.4% [14, 6.4%]; species occurring north of<br />
Cape Kennedy, in the Gulf, and in the Caribbean—12, 7.2% [21, 9.6%] The<br />
conclusion that the fauna of the Gulf of Mexico contains an endemic compo<br />
nent, a Caribbean component, and a warm-temperate component was also<br />
reached by Topp and HofT (1972), in an analysis of the pleuronectiform<br />
fishes of the Gulf.<br />
THE BAHAMAS<br />
The Florida Current flowing through the Straits of Florida has been sug<br />
gested as a factor in reducing the movement of shallow-water fauna between<br />
peninsular Florida and the Florida Keys on the west and the Bahamas on the<br />
east (Briggs, 1974). Comparison of the number of isopod species on either<br />
side of the Straits of Florida (13 from the Bahamas, 50 from southern penin<br />
sular Florida and the Florida Keys) supports this view. Of the 13 species<br />
from the Bahamas, only four arc "endemic," three of these being interstitial<br />
microcerberideans.<br />
Paradella dianae (Menzies, 1962b)<br />
Paraleptosphaeroma glynni Buss and<br />
Iverson, 1981<br />
Probopyrus pandalicola (Packard,<br />
1879)<br />
*Rocinela oculata Harger, 1883<br />
*Rocinela signata Schioedte and<br />
Meinert, 1879<br />
Uromunna reynoldsi Frankenberg and<br />
Menzies, 1966
TABLE 6. ISOPOD SPECIES OCCURRING IN THE GULF OF MEXICO<br />
SUBORDER ANTHURIDEA<br />
*Accalathura crenulata (Richardson,<br />
1901)<br />
*Amakusantkura magnified (Menzies<br />
and Frankenberg, 1966)<br />
Cyathura polita (Stimpson, 1855)<br />
Horoloanthura irpex Menzies and<br />
Frankenberg, 1966<br />
Kupellonura formosa (Menzies and<br />
Frankenberg, 1966)<br />
*Mesanthura Jloridensis Menzies and<br />
Kruczynski, 1983<br />
Mesanthura hopkinsi Hooker, 1985<br />
Mesanthura pulchra Barnard,<br />
1925<br />
Paranthura Jloridensis Menzies and<br />
Kruczynski, 1983<br />
Ptilanthura tricarina Menzies and<br />
Frankenberg, 1966<br />
Skuphonura lindae Menzies and<br />
Kruczynski, 1983<br />
*Xenanthura brevitelson Barnard,<br />
1925<br />
SUBORDER ASELLOTA<br />
Carpias Jloridensis Menzies and<br />
Kruczynski, 1983<br />
Gnathostenetrioides pugio Hooker,<br />
1985<br />
*Joeropsis coralicola Schultz and<br />
McCloskey, 1967<br />
*Joeropsis rathbunae Richardson,<br />
1902<br />
Mexicope kensleyi Hooker, 1985<br />
Munnogonium wilsoni Hooker, 1985<br />
*Pleurocope Jloridensis Hooker,<br />
1985<br />
*Santia milleri (Menzies and Glynn,<br />
1968)<br />
*Stenetrium stebbingi Richardson,<br />
1902<br />
ZOOGEOGRAPHY 267<br />
Uromunna hayesi Robertson, 1978<br />
*Uromunna reynoldsi Frankenberg<br />
and Menzies, 1966<br />
SUBORDER EPICARIDEA<br />
Allodiplophryxus Jloridanus Markham,<br />
1985<br />
*Aporobopyrina anomala Markham,<br />
1973<br />
*Azygopleon schmitti (Pearse, 1932)<br />
*Bopyrina abbreviata Richardson,<br />
1904<br />
*Bopyrione synalphei Bourdon and<br />
Markham, 1980<br />
*Cancricepon choprae (Nierstrasz and<br />
Brender a Brandis, 1925)<br />
Dactylokepon sulcipes Adkison, 1982<br />
Eophryxus subcaudalis (Hay, 1917)<br />
*Gigantione mortenseni Adkison,<br />
1984b<br />
Gigantione uberlackerae Adkison,<br />
1984b<br />
*Hemiarthrus synalphei (Pearse,<br />
1950)<br />
Hyperphrixus castrensis Markham,<br />
1985<br />
*Munidion longipedis Markham,<br />
1975a<br />
Ovobopyrus alphezemiotes Markham,<br />
1985<br />
Parabopyrella mortenseni (Nierstrasz<br />
and Brender a Brandis, 1929)<br />
*Parabopyrella richardsonae<br />
(Nierstrasz and Brender a<br />
Brandis, 1929)<br />
Parabopyriscus stellatus Markham,<br />
1985<br />
*Probopyria alphei (Richardson,<br />
1900b)<br />
Probopyrinella heardi Adkison,<br />
1984a<br />
{continued)
268<br />
TABLE 6. (Continued)<br />
*Probopyrinella latreuticola (Gissler,<br />
1882)<br />
Prodajus cf. bigelowiensis Schultz and<br />
Allen, 1982<br />
Pseudione cognata Markham, 1985<br />
Pseudione upogebiae Hay, 1917<br />
*Schizobopyrina urocaridis (Richardson,<br />
1904)<br />
*Stegophryxus hyptius Thompson, 1902<br />
*Synsynella choprae (Pearse, 1932)<br />
*Synsynella deformans Hay, 1917<br />
Synsynella Integra Bourdon, 1981<br />
* Urobopyrus processae Richardson, 1904<br />
SUBORDER FLABELLIFERA<br />
*Aega deshaysiana (H. Milne Edwards,<br />
1840)<br />
*Aega ecarinata Richardson, 1898<br />
Aega incisa Schioedte and Meinert,<br />
1879<br />
Alcirona krebsii Hansen, 1890<br />
Ancinus depressus (Say, 1818)<br />
Anilocra acuta Richardson, 1910<br />
Anilocra laticauda H. Milne Edwards,<br />
1840<br />
*Bathynomus giganteus A. Milne<br />
Edwards, 1879<br />
*Cassidinidea ovalis (Say, 1818)<br />
Ceratothoa transversa (Richardson,<br />
1900b)<br />
Cirolana borealis Lilljeborg, 1851<br />
*Cirolana obtruncata Richardson, 1901<br />
* Cirolana parva Hansen, 1890<br />
Conilera cylindracea (Montagu, 1804)<br />
*Cymothoa caraibica Bovallius, 1885<br />
*Cymothoa excisa Perty, 1833<br />
*Cymothoa oestrum (Linnaeus, 1793)<br />
*Cerceis carinata Glynn, 1970<br />
*Eurydice convexa Richardson, 1900b<br />
Eurydice littoralis (Moore, 1901)<br />
*Eurydice piperata Menzies and<br />
Frankenberg, 1966<br />
*Excirolana braziliensis Richardson,<br />
1912a<br />
*Excirolana mayana (Ives, 1891)<br />
^Excorallana antillensis (Hansen,<br />
1890)<br />
Excorallana mexicana Richardson,<br />
1905a<br />
*Excorallana tricornis (Hansen,<br />
1890)<br />
*Harrieta faxoni (Richardson,<br />
1905a)<br />
*Limnoria tuberculata Sowinsky,<br />
1884<br />
Lironeca ovalis (Say, 1818)<br />
* Lironeca redmanni Leach, 1818<br />
Lironeca texana Pearse, 1952<br />
Lironeca tropicalis Menzies and<br />
Kruczynski, 1983<br />
*Nalicora rapax Moore, 1901<br />
*Nerocila acuminata Schioedte and<br />
Meinert, 1881<br />
Olencira praegustator (Latrobe, 1802)<br />
*Paracerceis caudata (Say, 1818)<br />
*Paradella dianae (Menzies, 1962b)<br />
Paradynamene benjamensis<br />
Richardson, 1905<br />
*Politolana polita (Stimpson, 1853)<br />
*Rocinela insularis Schioedte and<br />
Meinert, 1879<br />
*Rocinela oculata Harger, 1883<br />
*Rocinela signata Schioedte and<br />
Meinert, 1879<br />
*Serolis mgrayi Menzies and<br />
Frankenberg, 1966<br />
*Sphaeroma quadridentata Say, 1818<br />
*Sphaeroma terebrans Bate, 1866<br />
SUBORDER GNATHIIDEA<br />
Gnathia Jloridensis Menzies and<br />
Kruczynski, 1983<br />
SUBORDER MICROCERBERIDEA<br />
Microcerberus mexicanus Pennak, 1958
SUBORDER VALVIFERA<br />
Antarcturus jloridanus (Richardson,<br />
1900b)<br />
Arcturella bispinata Menzies and<br />
Kruczynski, 1983<br />
Arcturella spinata Menzies and<br />
Kruczynski, 1983<br />
<strong>As</strong>tacilla lauffi Menzies and<br />
Frankenberg, 1966<br />
Chiridotea excavata Harper,<br />
1974<br />
Cleantioides planicauda (Benedict,<br />
1899)<br />
species also occurring in the Caribbean<br />
ZOOGEOGRAPHY 269<br />
Edotea lyonsi (Menzies and<br />
Kruczynski, 1983)<br />
Edotea montosa (Stimpson, 1853)<br />
Edwinjoycea horologium Menzies and<br />
Kruczynski, 1983<br />
*Erichsonella attenuata (Harger,<br />
1873)<br />
* Erichsonella filiformis (Say, 1818)<br />
* Erichsonella Jloridana Benedict,<br />
1901<br />
Erichsonella isabelensis Menzies,<br />
1951b<br />
*Idotea metallica Bosc, 1802<br />
Note: Records for the Gulf of Mexico have been assembled from published<br />
BERMUDA<br />
literature; in most cases, actual material has not been examined.<br />
Twenty-nine species of isopods have been recorded from Bermuda (Table 7).<br />
Of these, nine are endemics (three being cave forms). The remaining 20<br />
species have all been recorded from the Caribbean region, indicating a strong<br />
subtropical connection, in spite of the relatively high latitude (32°15'N). Al<br />
though Bermuda is of Eocene or Oligocene age, the tropical fauna was prob<br />
ably decimated by the low temperatures of the last Pleistocene glaciation<br />
(Briggs, 1974:76).<br />
GAVE ISOPODS<br />
With the expanding efforts of cave divers, more and more true stygobiont<br />
forms are being found. Concurrently, discussion of the origin of cave fauna<br />
has spurred several theories, all invoking the geological history of the Carib<br />
bean area.<br />
Among the isopods, cave forms have been found in four suborders, the<br />
<strong>As</strong>ellota, Anthuridea, Flabellifera, and Microcerberidea. Two valuable dis<br />
cussions on the origin of cave crustaceans may be found in Stock (1986) and<br />
Wagele (1985).<br />
The only true cave asellote, Atlantasellus cavernicolus Sket, was collected<br />
from Bermuda.
270 ZOOGEOGRAPHY<br />
TABLE 7. ISOPOD SPECIES OCCURRING AT BERMUDA<br />
Alcirona krebsi Hansen, 1890<br />
*Anthomuda stenotelson Schultz, 1979<br />
*Apanthura harringtortiensis Wagele,<br />
1981<br />
*Arubolana aruboides (Bowman and<br />
Iliffc, 1983)<br />
*Atlantasellus cavernicolus Sket, 1979<br />
Bopyrissa wolffi Markham, 1978<br />
Cancricepon choprae (Nierstrasz and<br />
Brender a Brandis, 1925)<br />
Carpias bermudensis Richardson, 1902<br />
*Carpias minutus (Richardson, 1902)<br />
*Colanthura tenuis Richardson, 1902<br />
Colopisthus parvus Richardson, 1902<br />
* Curassanthura bermudensis Wagele,<br />
1985<br />
Dynamenella perforata (Moore, 1901)<br />
Eurydice per sonata Kensley, 1987b<br />
* recorded only from Bermuda<br />
Excorallana quadricornis (Hansen, 1890)<br />
Joeropsis rathbunae Richardson, 1902<br />
Leidya bimini Pearse, 1951<br />
Paracerceis caudata (Say, 1818)<br />
Paranthura infundibulata Richardson,<br />
1902<br />
Pendanthura tanaiformis Menzies and<br />
Glynn, 1968<br />
Para theIges piriformis Markham, 1972b<br />
Parathelges tumidipes Markham, 1972b<br />
Probopyrinella latreuticola (Gissler,<br />
1882)<br />
Pseudione affinis (Sars, 1882)<br />
*Stegias clibanarii Richardson, 1904<br />
Stenetrium stebbingi Richardson, 1902<br />
Stenobermuda acutirostrata Schultz, 1979<br />
Synsynella choprae (Pearse, 1932)<br />
Synsynella deformans Hay, 1917<br />
The anthuridean cave representatives are found in two families: the genus<br />
Curassanthura Kensley in the Paranthuridae, and the genus Cyathura subgenus<br />
Stygocyathura Botosaneanu and Stock in the Anthuridae (see Figure 110).<br />
Three species of Curassanthura are known, one each from Curagao, Ber<br />
muda, and Lanzarote in the Canary Islands. Curassanthura halma Kensley,<br />
from Curagao, is an interstitial form found in hypersaline waters. Curas<br />
santhura bermudensis Wagele was found in water of about 26%o salinity. The<br />
Lanzarote species, C. canadensis Wagele, came from seawater in a lava cave.<br />
Wagele (1985) suggests that this amphi-Atlantic distribution of Curassanthura<br />
is the result of plate tectonics separating an ancestral hypogean progenitor<br />
that had a Tethyan distribution.<br />
The genus Cyathura has representatives in the sea, in estuarine-brackish<br />
habitats, and in freshwater caves, and is found in the Atlantic, Indian, and<br />
Pacific oceans. This widespread distribution suggests a very long history for<br />
the genus. Using the morphology of the male copulatory stylet, Wagele<br />
(1985) suggests that marine ancestors, having a Tethyan distribution, en<br />
tered freshwater interstitial habitats. The series of regressions of sea level<br />
forms.<br />
serv
Figure 110. Map showing distribution of cave anthurideans.
* cave cirolanids<br />
Figure 111. Map showing distrib
ZOOGEOGRAPHY 273<br />
The flabelliferan family Cirolanidae contains five stygobiont genera in the<br />
Caribbean: Anopsilana, Arubolana, Bahalana, Creaseriella, and Haptolana (Figure<br />
111). Six other genera are known from the North American continent:<br />
Antrolana, Cirolanidesy Mexilana, Speocirolana, Sphaerolana, and Troglocirolana, all<br />
of which, except Antrolana from the Appalachian Valley of Virginia, occur in<br />
Mexico and Texas (see Notenboom, 1981). A few of these forms occur in<br />
brackish water, but most are found in freshwater of caves. Cave cirolanids<br />
are also known from Palau, North and East Africa, Madagascar, Bulgaria,<br />
Greece, Jugoslavia, Israel, France, and Spain. This widespread distribution<br />
again suggests a Tethyan marine origin, with dispersal and isolation due to<br />
sea regressions.<br />
The suborder Microcerberidea and the asellotan family Microparasellidae<br />
contain almost entirely interstitial forms, although few occur in caves. At<br />
least two genera, Microcerberus and Angliera, have very widespread distribu<br />
tions and are known from marine, brackish-water, and freshwater habitats,<br />
and may well have a history similar to that of Cyathura.
Appendix<br />
Since the manuscript of this work was completed and sent to press, a few<br />
papers have appeared either describing new species, mentioning new<br />
records for the Caribbean and associated areas, or instituting a major<br />
new taxon. It was thought useful to include these, if only in an appendix,<br />
to make the work as current as possible. The relevant taxa are listed<br />
alphabetically, with the full citation given below.<br />
Antheluridae Poore and Lew Ton, 1988<br />
Poore, G. C. B., and H. M. Lew Ton. 1988. Antheluridae, a new<br />
family of Crustacea (Isopoda: Anthuridea) with new species from<br />
Australia. Journal of Natural History 22:489—506.<br />
Within the geographical area covered by this work, only Anthomuda<br />
belongs to this new family.<br />
Aporobopyrus collardi Adkison, 1988<br />
Adkison, D. L. 1988. Pseudione parviramus and Aporobopyrus collardi,<br />
two new species of Bopyridae (Isopoda: Epicaridea) from the Gulf<br />
of Mexico. Proceedings of the Biological Society of Washington<br />
101(3):576-584.<br />
Booralana tricarinata Camp and Heard, 1988<br />
Camp, D. K., and R. W. Heard. 1988. Booralana tricarinata, a new<br />
species of isopod from the western Atlantic Ocean (Crustacea: Iso<br />
poda: Cirolanidae). Proceedings of the Biological Society of Washington<br />
101 (3):603-613.<br />
Originally recorded from the outer shelf and upper slope off the<br />
Little Bahama Bank and the Antilles Islands in 110-610 m, this<br />
species has since been recorded off Haiti in 620 m.<br />
Bythognathia yucatanensis Camp, 1988<br />
Camp, D. K. 1988. Bythognathia yucatanensis, new genus, new species,<br />
from abyssal depths in the Caribbean Sea, with a list of gnathiid<br />
275
276 APPENDIX<br />
species described since 1926 (Isopoda: Gnathiidae). Journal of Crust<br />
acean Biology 8(4):668-678.<br />
Edotea samariensis Muller, 1988<br />
Muller, H. G. 1988. Idoteidae aus N-Kolumbien mit Beschreibung<br />
von Edotea samariensis n. sp. (Crustacea: Isopoda: Valvifera).<br />
Senckenbergiana biologica 68(4/6):407-412.<br />
Gnathia Johanna Monod, 1926<br />
Miiller, H. G. 1988. Redescription of Gnathia Johanna, 1926 (Iso<br />
poda) from St. John, Virgin Islands. Bulletin Zoologisch Museum, Uni-<br />
versiteit van Amsterdam 11(15): 129—133.<br />
Phycolimnoria bacescui Ortiz and Lalana, 1988<br />
Ortiz, M., and R. Lalana. 1988. Una nueva especie del genero Phy<br />
colimnoria (Isopoda, Limnoriidae) de aguas cubanas. Revista de Inves<br />
tigations Marinas, La Habana 9(2): 37—42.<br />
Pseudione parviramus Adkison, 1988<br />
Adkison, D. L. 1988. Pseudione parviramus and Aporobopyrus collardi,<br />
two new species of Bopyridae (Isopoda: Epicaridea) from the Gulf<br />
of Mexico. Proceedings of the Biological Society of Washington<br />
101(3):576-584.
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. 1925. A revision of the family Anthuridae (Crustacea Isopoda), with remarks on<br />
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Bate, C. S. 1866. Carcinological gleanings, 2. Annals and Magazine of Natural History (3)17:24-<br />
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Bijdragen tot de Dierkunde 52(l):43-48.<br />
. 1983. On the origin of the Microcerberidae (Crustacea: Isopoda). Zeitschrift fur<br />
zoologische Systematik und Evolutionsforschung 21 (4):249—262.<br />
. 1985. New west Atlantic localities for the stygobiont paranthurid Curassanthura<br />
(Crustacea, Isopoda, Anthuridea) with description of C. bermudensis n. sp. Bijdragen tot de<br />
Dierkunde 55(2):324-330.<br />
Walker, A. O. 1901. Contributions to the Malacostracan fauna of the Mediterranean. Journal<br />
of the Linnean Society of London, Zoology 28:290-307.<br />
Waterman, T. H., ed. 1960. The Physiology of Crustacea, vols. 1, 2, 670, 681 pp. New York &<br />
London: Academic Press.<br />
Williams, E. H., and L. B. Williams. 1980. Four new species of Renocila (Isopoda:<br />
Cymothoidae), the first reported from the New World. Proceedings of the Biological Society of<br />
Washington 93(3):573-592.<br />
. 1982. Mothocya bohlkeorum, new species (Isopoda: Cymothoidae) from West Indian<br />
cardinalfishes (Apogonidae). Journal of Crustacean Biology 2(4):570-577.
292 LITERATURE CITED<br />
. 1985a. A new cymothoid isopod, Glossobius hemiramphi, from the mouth of the<br />
ballyhoo, Hemiramphus brasiliensis (Linnaeus) (Exocoetidae), in the Caribbean Sea.<br />
Crustaceana 48(2): 147-152.<br />
. 1985b. Cuna insularis n. gen. and n. sp. (Isopoda: Cymothoidae) from the gill<br />
chamber of the sergeant major Abudefduf saxatilis (Linnaeus) (Osteichthyes) in the West<br />
Indies. Journal of Parasitology 71 (2):209-214.<br />
. 1986. Kuna Nomen Novum for Cuna Williams and Williams, 1985, preoccupied by Cuna<br />
Hedley, 1902. Journal of Parasitology 72(6):879.<br />
Williams, L. B., and E. H. Williams. 1981. Nine new species of A nilocra (Crustacea: Isopoda:<br />
Cymothoidae) external parasites of West Indian coral reef fishes. Proceedings of the Biological<br />
Society of Washington 94(4): 1005-1047.<br />
Wilson, G. D. F. 1987. The road to the Janiroidea: Comparative morphology and evolution<br />
of the asellote isopod crustaceans. Zeitschrifi fir Zfiologische Systematik und Evolutionsforschung<br />
25(4):257-280.
Index<br />
abbreviata, Bopyrina, 110, 267<br />
Abudefduf saxatilis, 171, 175, 186<br />
abudefdufi, Anilocra, 171, 175<br />
Abyssianira dentifrons, 263<br />
Acanthocope spinosissima, 263<br />
acanthopkora, Domecia, 111<br />
Acanthopleura granulata, 215<br />
acanthura, Anopsilana, 124, 125<br />
acanthuri, Anilocra, 171, 175, 176<br />
Acanthurus<br />
bahianus, 171, 176<br />
chirurgus, 171, 176<br />
acanthurus, Macrobrachium, 112<br />
crenulata, 64, 65, 267<br />
setosa, 64, 65<br />
Achelion occidentalism 110<br />
acuminata, Nerocila, 171, 172, 173, 190, 266<br />
268<br />
forma acuminata, 190<br />
forma &yter, 190<br />
acuta, Anilocra, 268<br />
acutirostrata, Stenobermuda, 106, 270<br />
acutirostris, Processa, 113<br />
acutitelson, Dynamenella, 213, 214<br />
ilagfl, 116, 117<br />
antillensiSy 117<br />
incisa, 268<br />
-4i?£fl (Aega), 116, 117<br />
deshaysiana, 117, 266, 268<br />
ecarinata, 117, 268<br />
-dtfgfl (Rhamphion), 116, 117<br />
^wtata, 117, 119<br />
tenuipes, 117, 119<br />
Aegidae, 115, 116, 262<br />
Aetobatus narinari, 166<br />
Pseudione, 113, 270<br />
Upogebia, 112<br />
Agaricia, 88, 166<br />
agaricicola, Metacirolana, 153, 154<br />
A£
294 INDEX<br />
angulata<br />
Dynamene, 214<br />
Dynamenella, 214<br />
angustifrons, Hexapanopeus, 111<br />
Anilocra, 170, 174, 175<br />
abudefdufi, 171, 175<br />
acanthuri, 171, 175, 176<br />
acuta, 268<br />
chaetodontis, 171, 175, 177<br />
fAnwiw, 172, 175, 177<br />
haemuli, 172, 173, 175, 177<br />
kolacantki, 172, 175, 179<br />
holocentri, 175, 179<br />
laticauda, 268<br />
myripristis, 172, 175, 180<br />
/wr/i/i, 173, 175, 180<br />
annandalei, Sphaeroma, 234<br />
annaoides, Antarcturus, 264<br />
annulicornis, Pandalus, 113<br />
annulipes, Pagurus, 113<br />
anomala, Aporobopyrina, 110, 267<br />
anops, Creaseriella, 137<br />
Anopsilana, 124, 273<br />
acanthura, 124, 125<br />
ArMWH, 124, 125, 266<br />
owiata, 124, 125<br />
cubensis, 124, 126<br />
;m, 124, 127<br />
radicicola, 124, 127<br />
annaoides, 264<br />
jloridanus, 269<br />
Antennuloniscus dimeroceras, 263<br />
Antheluridae, 275<br />
Anthomuda, 17, 23, 291<br />
stenotehon, 23, 270<br />
Anthuridae, 16, 270<br />
Anthuridea, 2, 14, 15, 16, 244, 263, 269<br />
antiguai, Plesionika, 113<br />
antillense, Exosphaeroma, 229, 230<br />
antillensis<br />
Aega, 117<br />
Dromidia, 111<br />
Excorallana, 161, 162, 268<br />
Paranthura, 69<br />
Antrolana, 273<br />
Apanthura? 17, 25<br />
cracenta, 25<br />
cram, 25, 26<br />
harringtoniensis3 25, 26, 270<br />
Apanthuroides? 17, 26<br />
millae, 27<br />
Aphysina fistulariSy 223<br />
lachneri, 171, 188<br />
maculatus, 171, 193<br />
townsendi, 171, 193<br />
Apogonidae, 187<br />
Aporobopyrina, 110<br />
anomala, 110, 267<br />
Aporobopyrus, 110<br />
collardi, 275<br />
curtatus, 110<br />
arbuscula, Oculina, 88<br />
Archosargus probatocephalus, 122<br />
Arcturella, 252<br />
bispinata, 252, 269<br />
#wiflte, 252, 269<br />
Arcturidae, 251, 252, 264<br />
caribbaeus, 264<br />
purpureus, 264<br />
arenatus, Priacanthus, 173, 183<br />
Argeia, 110<br />
atlantica, 110<br />
Argeiinae, 107<br />
Ariusfelis, 171, 190<br />
Armadilloniscus; 247<br />
m'/zai, 247<br />
Iscfinomesus, 263<br />
Petrolisthes, 110<br />
armillatuSy Alpheus, 112<br />
arndti, Dies, 207<br />
aruboidesy Arubolana, 144, 145, 270<br />
Arubolana, 144, 273<br />
aruboides, 144, 145, 270<br />
/ma/a, 144, 145<br />
parvioculata, 144, 145<br />
ascensionis, Holocentrus, 172, 179<br />
<strong>As</strong>elloidea, 75<br />
<strong>As</strong>ellota, 15, 15, 73, 263, 267, 269<br />
<strong>As</strong>tacilla, 252<br />
cymodocea, 252, 253<br />
/ajtf//
Atlantasellidae, 75<br />
Atlantasellus, 75<br />
cavernicolus, 75, 269, 270<br />
atlantica<br />
Argeia, 110<br />
Megalops, 172, 183<br />
attenuata, Erichsonella, 256, 258, 269<br />
aurolineatum, Haemulon, 172, 178<br />
Azygopleon, 110<br />
schmitti, 110, 267<br />
bacescui, Phycolimnoria, 276<br />
Bagatus, 83<br />
Bahalana, 124, 127, 128, 273<br />
cardiopus, 128<br />
geracei, 128<br />
mayana, 128<br />
bahianus, Acanthurus, 171, 176<br />
bajonado, Calamus, 122<br />
Balanopleon, 110<br />
tortuganus, 110<br />
Balistes vetula, 122<br />
balthica, Idotea, 259<br />
barbadensis, Micropanope, 111<br />
barbarae, Geogerceis, 215<br />
6anzar
296 INDEX<br />
Cancrion carolinus, 111<br />
capistratus, Chaetodon, 171, 177<br />
caraibica, Cymothoa, 182, 268<br />
Caranx, 122, 171<br />
hippos, 171, 183<br />
latus, 171, 183<br />
ruber, 171, 183<br />
carbonarium, Haemulon, 172, 178<br />
carcinus, Macrobrachium, 112<br />
cardiopus, Bahalana, 128<br />
caribaeus, Dactylokepon, 111<br />
caribbaeus, Arcturus, 264<br />
caribbea, Storthyngura pulchra, 263<br />
Ianirella, 263<br />
Malacanthura, 45<br />
Ischnomesus, 263<br />
Macrostylis, 263<br />
caribea, Uromunna, 94<br />
carinata, 'Cerceis,' 211, 268<br />
carolii, Metaphrixus, 111<br />
carolinense, Tozeuma, 112<br />
carolinus, Cancrion, 111<br />
Gzr/u
Colopisthus, 144, 146<br />
parvus, 147, 270<br />
confixa, Cortezura, 31<br />
Conilera cylindracea, 268<br />
Conilerinae, 123, 139<br />
conklini, Phaeoptyx, 173, 188<br />
constricta, Munida, 112<br />
convexa, Eurydice, 147, 148, 149, 268<br />
coralicolay Joeropsis, 88, 267<br />
Corallanidae, 115, 157<br />
corallicola, Minyanthura, 53<br />
corallinus, Pylopagurus, 112<br />
Cortezura, 17, 29<br />
confixa, 31<br />
penascoensis, 31<br />
cracenta, Apanthura, 25<br />
crassa, Virganthura, 73<br />
Creaseriella, 124, 137, 273<br />
anops, 137<br />
crenata, Anopsilana, 124, 125<br />
crenulata, Accalathura, 64, 65, 267<br />
crenulitelson, Cirolana, 132, 133<br />
cromis, Pogonias, 173, 190<br />
cruciaria, <strong>As</strong>talione, 110<br />
cruris, Apanthura, 25, 26<br />
cruentatus, Epinephelus, 172, 179<br />
crumenophthalmus, Selar, 173, 183<br />
Cryptoniscoidea, 107<br />
Crytoniscidae, 107, 109<br />
cubana, Cyathura (Cyathura), 33<br />
cubense, Munidion, 111<br />
Anopsilana, 124, 126<br />
tfocme/a, 119, 120<br />
cuborientalis, Cyathura (Stygocyathura), 34, 35<br />
culebrae, Vandeloscia, 247, 251<br />
cumanensis, Malacanthura, 45<br />
cumulus, Agarna, 173<br />
cuprea, Diopatra, 256<br />
curacaoensis, Hippolyte, 110<br />
Curassanthura, 64, 67, 270<br />
bermudensis, 67, 270<br />
canariensis, 270<br />
Afl/mfl, 67, 68, 270<br />
curassavica, Cyathura (Stygocyathura), 35<br />
o/rn, Haliophasma, 41<br />
curtatus, Aporobopyrus, 110<br />
cuvieri, Galeocerdo, 122<br />
cyaneus, Chromis, 172, 177<br />
Cyathura, 17, 31, 270, 273<br />
/>o/z7a? 267<br />
(Cyathura), 31<br />
cubana, 33<br />
(Stygocyathura), 31, 33, 35, 270<br />
cuborientalis, 34, 35<br />
curassavica, 35<br />
hummelincki, 35<br />
motasi, 35, 36<br />
orghidani, 35, 36<br />
parapotamica, 35, 36<br />
salpiscinalis, 35, 38<br />
sbordonii, 35, 38<br />
specus, 35, 38<br />
univam, 35, 38<br />
Cyclograpsus interger, 111<br />
cylindracea, Conilera, 268<br />
Cymodoce, 226, 227<br />
barrerae, 227<br />
ruetzleri, 227<br />
Cymodocea, 223, 253<br />
cymododea, <strong>As</strong>tacilla, 252, 253<br />
Cymothoa, 170, 172, 182<br />
caraibica, 182, 268<br />
«OM, 172, 173, 182, 268<br />
INDEX 297<br />
wrfrem, 171, 172, 173, 182, 183, 268<br />
Cymothoidae, 115, 169, 170<br />
Cynoscion, 172, 183<br />
nebulosus, 172, 183<br />
Dactylokepon<br />
caribaeus, 111<br />
sulcipes, 267<br />
Dajidae, 107<br />
Dardanus jucosus, 1122<br />
Dasyatis americana, 122, 166<br />
decorata, Mesanthura, 53<br />
deformans, Synsynella, 110, 113, 268, 270<br />
delaneyi, Excorallana, 160, 161<br />
Dendrotiidae, 263<br />
Dendrotion hanseni, 263<br />
dentata, Aega, 117, 119<br />
dentifrons, Abyssianira, 263<br />
deplanata, Ceratothoa, 180<br />
depressa, Panoplax, 111<br />
depressus, Ancinus, 268<br />
deshaysiana, Aega, 117, 266, 268<br />
Desmosoma magnispina, 263<br />
Desmosomatidae, 263<br />
destructor, Sphaeroma, 235<br />
desultor, <strong>As</strong>ymmetrione, 110<br />
rfifl/M*, Paradella, 224, 266, 268<br />
Dicropleon<br />
periclimenis, 111<br />
Dictyota, 219<br />
Z)r>j, 207<br />
flnw/rf, 207<br />
barnardi, 207
298 INDEX<br />
dimeroceras, Antennuloniscus, 263<br />
diminuta, Exosphaeroma, 229, 231<br />
diogenes, Petrochirus, 110<br />
Diopatra cuprea, 256<br />
Diplophryxus, 111<br />
Discerceis, 210, 211<br />
linguicauda, 213<br />
dispar, Paraliomera, 111<br />
distorta, Leidya, 111<br />
Domecia<br />
acanthophora, 111<br />
hispida, 111<br />
Dromidia antillensis, 111<br />
dubitans, Angliera, 91<br />
Dynamene angulata, 214<br />
Dynamenella, 210, 213, 214, 224<br />
acutitelson, 213, 214<br />
var. glabrothorax, 214<br />
var. typica, 214<br />
angulata, 214<br />
perforata, 213, 215, 270<br />
quadrilirata, 213, 215<br />
Dynameninae, 204, 210, 211<br />
ecarinata, Aega, 117, 268<br />
Echinothambema ophiuroides, 263<br />
Echinothambematidae, 263<br />
edithae, Paracerceis, 218, 221<br />
Edo tea<br />
lyonsi, 269<br />
montosa, 269<br />
samariensis, 276<br />
edulis, Processa, 113<br />
edwardsi, Plesionika, 113<br />
Edwinjoycea horologium, 252, 269<br />
eglanteria, Raja, 122<br />
Eisothistos, 16, 38, 39<br />
petrensis, 39<br />
ten, 39<br />
mm, Plesionika, 113<br />
Entoniscidae, 107, 109<br />
Entophilinae, 107<br />
Eophrixus subcaudalis, 111, 267<br />
Epicaridea, 4, 14, 107, 267<br />
Epinephelus, 122, 172, 183<br />
cruentatus, 172, 179<br />
>/z^, 172, 179<br />
guttatus, 172, 179<br />
*7, 122, 172, 190<br />
morio, 122<br />
Erichsonella, 255, 256, 257<br />
attenuata, 256, 258, 269<br />
Jiliformis, 256, 258, 269<br />
tropicalis, 257<br />
floridana, 256, 258, 269<br />
isabelensis, 269<br />
Eriphia gonagra, 111<br />
Eubranchiatae, 203<br />
Eurycopidae, 263<br />
Eurydice, 143, 147<br />
con^xfl, 147, 148, 149, 268<br />
littoralis, 148, 149, 268<br />
personata, 147, 149, 270<br />
piperata, 147, 149, 268<br />
Eurydicinae, 123, 139, 143<br />
excavata, Chiridotea, 269<br />
Excirolana, 144, 149, 150<br />
braziliensis, 150, 266, 268<br />
mayana, 150, 153, 268<br />
Atrial, Cymothoa, 172, 173, 182, 268<br />
Excorallana, 157, 159, 161<br />
antillensis, 161, 162, 268<br />
berbicensis, 161, 162<br />
delaneyi, 160, 161<br />
fissicauda, 161, 162<br />
mexicana, 160, 161, 268<br />
oculata, 161, 163<br />
quadricornis, 161, 165, 270<br />
sexticornis, 161, 165<br />
subtilis, 160<br />
tricornis, 266, 268<br />
occidentalis, 167<br />
tricornis, 161, 165<br />
warmingii, 161, 167<br />
exilipes, Palaemonetes, 113<br />
Exocoettus, 172, 184<br />
Exosphaeroma, 226, 229, 231<br />
fl/Afl, 229<br />
antillense, 229, 230<br />
diminuta, 229, 231<br />
productatelson, 229, 231<br />
yucatanum, 229, 231<br />
exotica, Ligia, 247, 249<br />
faber, Chaetodipterus, 171, 190<br />
fasciata, Mesanthura, 47, 49<br />
faustinum, Macrobrachium, 112<br />
/flxwii, Harrieta, 232, 268<br />
felis,Arius, 171, 190<br />
filiforme, Syringodium, 253, 260<br />
Jiliformis, Erichsonella, 256, 258, 269<br />
fimbriata<br />
Pleurocryptella, 112<br />
Processa, 113<br />
fissicauda, Excorallana, 161, 162<br />
fistularis, Aphysina, 223<br />
Flabellifera, 2, 14, 114, 115, 268, 269<br />
Jlavolineatum, Haemulon, 122, 172, 178
Jlinti, Munida, 111<br />
jloridana<br />
Erichsonella, 256, 258, 269<br />
Pleurocrypta, 112<br />
florid anus<br />
Allodiplophryxus, 267<br />
Antarcturus, 269<br />
Thor, 110, 111<br />
jloridensis<br />
Carpias, 267<br />
Gnathic, 268<br />
Mesanthura, 53, 267<br />
Paranthura, 69, 71, 267<br />
Pleurocope, 98, 267<br />
joetens, Synodus, 173, 183<br />
foliatus, Parathelges, 112<br />
formosa, Kupellonura, 267<br />
formosus, Alpheus, 111, 112<br />
fritzmuelleri, Synalpheus, 110, 111<br />
jucorum, Latreutes, 112<br />
Jucosus, Dardanus, 112<br />
Julvus, Epinephelus, 172, 179<br />
Jurcifer, Paranthias, 173, 179<br />
Galathea rostrata, 112<br />
galathinus, Petrolisthes, 110<br />
Galeocerdo cuvieri, 122<br />
geminsula, Amakusanthura, 18<br />
Geocerceis, 210, 215<br />
barbarae, 215<br />
geracei, Bahalana, 128<br />
Genes rhombeus, 172, 187<br />
giardi, Synalpheion, 113<br />
giganteus, Bathynomus, 131, 268<br />
Gigantione<br />
mortenseni, 111, 267<br />
uberlackerae, 267<br />
Ginglymostoma cirratum, 122, 158<br />
glabrothorax, var., Dynamenella acutitelson, 214<br />
Glossobius, 170, 172, 183, 184<br />
hemiramphi, 172, 184<br />
impressus, 172, 184<br />
Paracerceis, 218, 221<br />
Paraleptosphaeroma, 210, 266<br />
Cwato, 238<br />
beethoveni, 238, 239<br />
Jloridensis, 268<br />
gonzalezi, 238, 239<br />
Johanna, 238, 239, 276<br />
magdalenensis, 238, 239<br />
puertoricensis, 238, 239<br />
ra^z, 238, 241<br />
samariensis, 238, 241<br />
triospathiona, 238, 241<br />
ye/
300 INDEX<br />
hemiramphi3 Glossobius, 172, 184<br />
Hemiramphidae, 187<br />
Hemiramphus<br />
bermudensis, 188<br />
brasiliensis, 172, 184<br />
hemphilli, Synalpheus, 110, 111<br />
hendleri, Pendanthura, 56<br />
herbstii, Panopeus, 111<br />
herrerai, Microcharon, 93<br />
heterocarpus, Plesionika, 113<br />
heterochaelis, A Ipheus, 112<br />
Heteromesus bijurcatus, 263<br />
Hexapanopeus angustifrons, 111<br />
Hippolyte<br />
curacao ensis 3 110<br />
pleutacanthus, 110, 111<br />
Zostericola, 110<br />
hippos, Caranx, 171, 183<br />
Hirundichthys speculifer, 172, 184<br />
hispida, Domecia, 111<br />
holacanthi, Anilocra, 172, 175, 179<br />
Holacanthus tricolor, 172, 179<br />
holocentri, Anilocra, 175, 179<br />
Holocentrus ascensionis, 172, 179<br />
hopkinsi, Mesanthura, 47, 51, 267<br />
Horoloanthura irpex, 267<br />
horologium, Edwinjoycea, 252, 269<br />
hummelincki, Cyathura (Stygocyathura), 35<br />
Hydroniscus quadrifrons, 263<br />
Hyperphrixus castrensis, 267<br />
Hypoconcha<br />
sabulosa, 111<br />
spinosissima, 111<br />
Hyporhamphus unifasciatus, 172, 188<br />
hyptius, Stegophryxus, 113, 268<br />
Hyssuridae, 16, 58, 60<br />
fanirella<br />
caribbica3 263<br />
vemae, 263<br />
Afotai, 255, 259<br />
balthica, 259<br />
metallicay 259, 269<br />
Idoteidae, 251, 252, 255<br />
Idoteinae, 255, 256<br />
Iliacantha<br />
liodactyla, 111<br />
subglobosa3 111<br />
imbricata, Parapagurion, 112<br />
impressa, Politolana, 140<br />
impressus, Glossobius, 172, 184<br />
imswe, Kupellonura, 60<br />
imw/a, Arubolana, 144, 145<br />
/nriftZj 4*£A, 268<br />
indica, Limnoria, 194, 195<br />
infundibulata, Paranthura, 69, 71, 270<br />
insulae, Limnoria, 195<br />
insular is<br />
Alcirona, 158<br />
A'ttrtfl, 170<br />
Rocinela, 119, 120, 268<br />
integra, Synsynella, 268<br />
interger, Cyclograpsus, 111<br />
intermedius, Palaemonetes, 113<br />
loninae, 107<br />
Iridopagurus, 112, 113<br />
zm, 113<br />
z'm, Iridopagurus, 113<br />
irmae, Haliophasma, 43<br />
fr/fcx, Horoloanthura, 267<br />
irrasa, Munida, 111<br />
irritans, Munidion, 111<br />
isabelensis, Erichsonella, 269<br />
Ischnomesidae, 263<br />
Ischnomesus<br />
armatus, 263<br />
caribbicus, 263<br />
multispinis, 263<br />
Ischyromene, 210, 214, 217<br />
bamardi, 218<br />
iVfljflrfl, Epinephelus, 122, 172, 190<br />
jacobus, Myripristis, 172, 180<br />
jacqueti, Sclerocrangon, 110<br />
Janiridae, 80, 81, 263<br />
Janiroidea, 75, 79, 80, 81<br />
Joeropsidae, 80, 87<br />
Joeropsis, 87, 88<br />
bifasciatus, 88<br />
coralicola, 88, 267<br />
personatus, 88, 90<br />
rathbunae, 88, 90, 267, 270<br />
JOAHHM, G/ifltfifl, 238, 239, 292<br />
jonesi, Anopsilana, 124, 127<br />
kadiakensis, Palaemonetes, 113<br />
kensleyi, Mexicope, 81, 267<br />
kohleri, Mesosignum, 264<br />
AWAJI, Alcirona, 158, 268, 270<br />
tfiwifl, 170, 184<br />
insularis, 171, 186<br />
Kupellonura, 60<br />
formosa, 267<br />
imswe, 60<br />
lachneri, Apogon, 171, 188<br />
Lachnolaimus maximus, 122<br />
Laminaria, 201<br />
lamprotaenia, Anchoa, 171, 187<br />
laodicense, Gnathostenetroides, 78
lasallae, <strong>As</strong>tacilla, 252, 253<br />
lata, Parabopyrella, 112<br />
lathridia, Amakusanthura, 18, 20<br />
laticauda, Anilocra, 268<br />
laticeps, Skuphonura, 58<br />
latreillei, Tylos, 247, 250<br />
Latreutes Jucorum, 112<br />
latreuticola, Probopyrinella, 112, 268, 270<br />
latus, Caranx, 171, 183<br />
/flw#?, <strong>As</strong>tacilla, 269<br />
Laurencia, 219<br />
Aimini, 111, 270<br />
distorta, 111<br />
Leiostomus xanthurus, 172, 183, 187,<br />
190<br />
Lepisosteus spatula, 172, 190<br />
leptorhynchus, Pandalus, 113<br />
lewisi, Chalixanthura, 27, 29<br />
Licranthura, 16, 43<br />
amyle, 43<br />
Lfcifl, 247, 249<br />
baudiniana, 247, 249<br />
wrrtiai, 247, 249<br />
302 INDEX<br />
mayana<br />
Bahalana, 128<br />
Excirolana, 150, 153, 268<br />
mcclendoni, Synalpheus, 110, 111<br />
Megalops atlantica, 172, 183<br />
menziesi, Metacirolana, 153, 154<br />
Mesantkura, 17, 45<br />
bivittata, 47<br />
decorata, 53<br />
fasciata, 47, 49<br />
JloridensiSy 53, 267<br />
hopkinsit479 51, 267<br />
looensis, 47, 51<br />
paucidens, 47, 51<br />
pulchra, 47, 52, 267<br />
punctillata, 47, 53<br />
reticulata, 47, 53<br />
Mesosignidae, 264<br />
Mesosignum kohleri, 264<br />
Metacirolana, 144, 153<br />
agaricicola, 153, 154<br />
Afl/ifl, 153, 154<br />
menziesi, 153, 154<br />
sphaeromiformis, 153, 154<br />
metallica, Idotea, 259, 269<br />
Metaphrixus carolii, 111<br />
mexicana, Excorallana, 160, 161, 268<br />
mexicanus, Microcerberus t 269<br />
Mexicope, 80, 81<br />
kensleyiy 81, 267<br />
Mexilana, 273<br />
mgrayiy Serolis, 202, 268<br />
Microcerberidae, 243, 244<br />
Microcerbendea, 14, 243, 269, 273<br />
Microcerberus, 244, 273<br />
littoralis, 244<br />
mexicanuSy 269<br />
minutuSy 244<br />
mirabiliSy 244<br />
nunezi, 244<br />
renaudiy 244<br />
simplex, 244<br />
syrtiaiSy 244<br />
Microcharon, 90, 91<br />
herrerai, 93<br />
phreaticus, 93<br />
sabulurrty 91<br />
Micropanope barbadensis, 111<br />
Microparasellidae, 80, 90, 273<br />
Microphrys bicornutus, 110<br />
//nV^r, Munida} 112<br />
millae, Apanthuroides, 27<br />
milleri, Santiay 99, 267<br />
minocule, Stenetrium, 100, 102<br />
minusy Synalpheus, 110, 113<br />
minuta, Cirolana, 132, 135<br />
Cfl#wj, 82, 84, 270<br />
MacrostyliSy 263<br />
Microcerberusy 244<br />
Minyanthura, 16, 53<br />
corallicola, 53<br />
mirabilis<br />
MadraciSy 29, 41<br />
Microcerberusy 244<br />
Miratidoteay 255, 259<br />
bruscai> 260<br />
Monacanthus ciliatuSy 172, 190<br />
montaguiy Pandalus, 113<br />
montosa, Edoteat 269<br />
morioy Epinephelus, 122<br />
Gigantione, 111, 267<br />
Parabopyrellay 112, 267<br />
mosaica, Cassidinidea, 208<br />
motasiy Cyathura (Stygocyathura), 35, 36<br />
Mothocya, 170, 187<br />
bohlkeorum, 171, 173, 187, 188<br />
wflwfl, 172, 187, 188<br />
Mugil cephalus, 172, 190<br />
multilineatus, Chromis, 172, 177<br />
multipunctata, Limnoria, 195, 196<br />
multispinis, Ischnomesus, 263<br />
constricta, 112<br />
TKnrt, 111<br />
irrasas 111<br />
longipeSy 112<br />
772 £/w, 112<br />
schroederiy 112<br />
simplex, 110<br />
stimpsoni, 111<br />
valida, 110<br />
Munidion<br />
cubensCy 111<br />
irritans, 111<br />
longipedisy 112, 267<br />
Munna, 93<br />
petronastes, 94<br />
Munnidae, 80, 93<br />
Munnogoniumy 96<br />
wilsoni, 96<br />
Mycteroperca<br />
bonaci, 122<br />
venenosa, 122<br />
Myripristis jacobus, 172, 180
myripristis, Anilocra, 172, 175, 180<br />
Nalicora, 157, 168<br />
rapax, 169, 268<br />
nana, Mothocya, 172, 187, 188<br />
Nannoniscidae, 264<br />
Nannoniscus camayae, 264<br />
narinari, Aetobatus, 166<br />
Natatolana, 139<br />
gracilis, 140<br />
nebulosus, Cynoscion, 172, 183<br />
Nemanthura, 43<br />
Neoanthura coeca, 263<br />
Neopanope<br />
packardii, 111<br />
ttxana sayi, 111<br />
Neostenetroides, 77, 78<br />
stocki, 78<br />
Nerocila, 170, 172, 188<br />
acuminata, 171, 172, 173, 190, 266, 268<br />
forma acuminata, 190<br />
forma aj/er, 190<br />
ninae, Armidilloniscus, 246, 247<br />
nfzwtf, T^/ar, 247, 250<br />
normanni, Alpheus, 112<br />
northropi, Palaemon, 113<br />
nunezi, Microcerberus, 244<br />
nuttingi, Paracerceis, 218, 223<br />
obtruncata, Cirolana, 132, 135, 268<br />
occiden talis<br />
Achelion, 110<br />
Cleantioides, 256<br />
Excorallana tricornis, 167<br />
Parathelges, 112<br />
ocellatus, Chaetodon, 171, 177 I<br />
Excorallana, 161, 163<br />
Rocinela, 119, 120, 266, 268<br />
Oculina arbuscula, 88<br />
Ocyurus chrysurus, 172, 183<br />
owfram, Cymothoa, 171, 172, 173, 182, 183,<br />
268<br />
ohione, Macrobrachium, 112<br />
Olencira praegustator, 268<br />
Zigia, 247, 249<br />
Macrobrachium, 112<br />
Ontilorpheus, 124, 139<br />
stebbingi, 139<br />
Oniscidea, 1, 4, 14, 15, 246, 247, 262<br />
ophiuroides, Echinothambema, 263<br />
Orbioninae, 107<br />
orghidani, Cyathura (Stygocyathura), 35,<br />
36<br />
Orthopristis<br />
chrysoptera, 172, 183<br />
raA^r, 122, 173, 179<br />
ovalis<br />
Cassidinidea, 207, 208, 268<br />
Lironeca, 268<br />
Ovobopyrus alphezemiotes, 267<br />
oxyophthalmus, Paguristes, 112<br />
Pachygrapsus transversus, 111<br />
packardii, Neopanope, 111<br />
Paguristes<br />
oxyophthalmus, 112<br />
tortugae, 112<br />
Pagurus<br />
annulipes, 113<br />
bonairensis, 110, 113<br />
brevidactylus, 112, 113<br />
longicarpus, 110, 113<br />
provenzanoi, 110, 112, 113<br />
Palaemon<br />
northropi, 113<br />
pandaliformis, 113<br />
Palaemonetes<br />
exilipes, 113<br />
intermedius, 113<br />
kadiakensis, 113<br />
paludosus, 113<br />
pugio, 113<br />
vulgaris, 113<br />
paludosus, Palaemonetes, 113<br />
pandalicola, Probopyrus, 112, 266<br />
pandaliformis, Palaemon, 113<br />
Pandalus<br />
annul ico rni s, 113<br />
Ari, 113<br />
leptorhynchus, 113<br />
montagui, 113<br />
pandionis, Synalpheus, 111, 113<br />
Panopeus herbstii, 111<br />
Panoplax depressa, 111<br />
Parabopyrella<br />
lata, 112<br />
mortenseni, 112, 267<br />
richardsonae, 112, 267<br />
thomasi, 112<br />
Parabopyriscus stellatus, 267<br />
Paracerceis, 210, 218, 223<br />
cflurfate, 219, 268, 270<br />
var. brevipes, 219<br />
roA«ifl*, 219, 220<br />
wfifAae, 218, 221<br />
£/>wii, 218, 221<br />
Hatting, 218, 223
304 INDEX<br />
Paradella, 210, 214, 223<br />
dianae, 224, 266, 268<br />
plicatura, 223, 224<br />
quadripunctata, 223, 224<br />
tumidicauda, 223, 226<br />
Paradynamene benjamensis, 268<br />
Paragnathia, 237<br />
Paraleptosphaeroma, 207, 208<br />
^iww, 210, 266<br />
Paralimnoria, 193, 199<br />
andrewsi, 199<br />
forma A, 200<br />
forma B, 201<br />
forma typica, 200<br />
Paraliomera dispar, 111<br />
Paramunnidae, 80, 96<br />
Paranthias farcifer, 173, 179<br />
Paranthura, 64, 69<br />
antillensis, 69<br />
barnardiy 69, 71<br />
floridensis, 69, 71, 267<br />
infundibulatay 69, 71, 270<br />
Paranthuridae, 16, 64, 263, 270<br />
Parapagurion imbricata, 112<br />
Parapagurus, 112<br />
parapotamica, Cyathura (Stygocyathura), 35,<br />
Para theIges<br />
foliatus, 112<br />
occidentalism 112<br />
piriformis, 112, 270<br />
tumidipes, 112, 270<br />
partiti, Anilocra, 173, 175, 180<br />
partitus, Pomacentrus3 173, 180<br />
parva, Cirolana, 132, 135, 268<br />
parvioculata, Arubolana, 144, 145<br />
parviramus, Pseudione, 276<br />
parvus, Colopisthus, 147, 270<br />
patulipalma, Stenetrium, 100, 102<br />
paucidens, Mesantkura, 47, 51<br />
pectiniger, Synalpheus, 110, 111, 113<br />
penascoensis; Cortezura, 31<br />
Pendanthura, 17, 56<br />
hendleri, 56<br />
tanaiformis, 56, 270<br />
penna, Calamus, 122<br />
perforata, Dynamenella, 213, 215, 270<br />
Periclimenes<br />
americanus, 111, 113<br />
longicaudatus, 113<br />
periclimenis, Dicropleon, 111<br />
personata, Eurydice, 147, 149, 270<br />
personatus, Joeropsis, 88, 90<br />
petrensis, Eisotkistos, 39<br />
Petrochirus diogenes, 110<br />
Petrolisthes<br />
armatus, 110<br />
galathinus, 110<br />
marginatus, 110<br />
petronastesy Munna, 94<br />
pfefferi, Limnoria, 195, 198<br />
Phaeoptyx<br />
conklini, 173, 188<br />
pigmentaria, 173, 188<br />
phreaticuSy Microcharoriy 93<br />
Phreatocoidea, 14<br />
Phycolimnoria, 193, 201<br />
bacescuiy 276<br />
clarkae, 201<br />
Phyllodurinae, 107<br />
pigmentariay Phaeoptyx, 173, 188<br />
piperata, Eurydice, 147, 149, 268<br />
piriformis, Parathelges, 112, 270<br />
planicauda, Cleantioides, 256, 266, 268<br />
Platybranchiatae, 203<br />
platycauda, Limnoria, 195, 198<br />
Plesionika<br />
antiguai, 113<br />
edwardsiy 113<br />
mw, 113<br />
heterocarpus, 113<br />
martia, 113<br />
p leuracanthus, Hip poly te, 110, 111<br />
Pleurocope, 97<br />
floridensisy 98, 267<br />
Pleurocopidae, 80, 81, 96<br />
Pleurocryptella fimbriata, 112<br />
plicatura, Paradella, 223, 224<br />
plicifera, Callispongia, 221<br />
plumieri, Haemulon, 172, 179<br />
Pogonias cromis, 173, 190<br />
polita<br />
Cyathura, 267<br />
Politolana, 140, 143, 268<br />
Politolana, 139, 140<br />
impressa, 140<br />
^o/ito, 140, 143, 268<br />
Pomacentrus partitus, 173, 180<br />
Pontonia margarita, 113<br />
Porce liana say ana, 110<br />
A>n7^, 88, 90<br />
poteriophora, Akidognathia, 264<br />
praegustator, Olencira, 268<br />
Priacanthus arenatus, 173, 183<br />
probatocephalus, Archosargus, 122<br />
Probopyria alphei, 112, 267
Probopyrinella<br />
heardiy 267<br />
latreuticola, 112, 268, 270<br />
Probopyrus pandalicola, 112, 266<br />
Processa<br />
acutirostris y 113<br />
canaliculata y 113<br />
eduliSy 113<br />
Jimbriatay 113<br />
tenuipes, 113<br />
processaey Urobopyrusy 113, 268<br />
Prodajus cf. bigelowiensis, 268<br />
productatelson, Exosphaeroma3 229, 231<br />
provenzanoiy Pagurus> 110, 112, 113<br />
psamathus, Anglieray 91<br />
Pseudasymmetrione, 113<br />
Pseudione<br />
qffinis, 113, 270<br />
cognata, 268<br />
parviramuSy 276<br />
upogebiae, 268<br />
Pseudioninae, 107<br />
Ptilanthura tricarina, 267<br />
puertoricensis, Gnathia} 238, 241<br />
pugilatoty Uca, 111<br />
GnathostenetroideSy 77', 267<br />
PalaemoneteSy 113<br />
pulchra<br />
caribbedy Storthyngura, 263<br />
Mesanthura, 47, 52, 267<br />
punctatuSy Carpias, 82, 85<br />
punctillatdy Mesanthura, 47, 53<br />
purpureus, Arcturus3 264<br />
PylopaguruSy 110<br />
corallinus, 112<br />
quadricornisy Excorallana, 161, 165, 270<br />
quadridentata, Sphaeroma, 234, 268<br />
quadrifrons, Hydroniscus, 263<br />
quadriliratdy Dynamenella, 213, 215<br />
quadripunctata, Paradella, 223, 223<br />
racovitzai, Angliera, 91<br />
radicicola<br />
Anopsilana, 124, 127<br />
Xylolana, 157<br />
i?a/a eglanteridy 122<br />
rapaXy Nalicora, 169, 268<br />
rathbunae<br />
JoeropsiSy 88, 90, 267, 270<br />
Lysmata, 112<br />
rafAi, Gnathic, 238, 241<br />
redmanni, Lironecay 172, 173, 186, 268<br />
regalis, Scomberomorus, 173, 187<br />
regina, <strong>As</strong>tacilla, 252, 253<br />
renaudiy Microcerberus, 244<br />
Renocila, 170, 191<br />
bowmani, 173, 191<br />
«/!/!!, 171, 191<br />
waldneriy 173, 191, 193<br />
reticulatas Mesanthura, 47, 53<br />
INDEX 305<br />
reynoldsiy Uromunnay 94, 95, 266, 267<br />
RhamphioUy see i4«ga (Ramphion)<br />
Rhizophora mangle, 235<br />
rhombeus, GerreSy 172, 187<br />
Rhyscotus, 247, 249<br />
fexwuij, 247, 249<br />
richardsonae, Parabopyrella, 112, 267<br />
ricordiy Sesarma, 111<br />
nVrf/z, Vandeloscia, 247, 251<br />
Ritkropanopeus karrisiiy 111<br />
Rocinela, 116, 119<br />
cubensisy 119, 120<br />
insularis, 119, 120, 268<br />
oca/ate, 119, 120, 266, 268<br />
jignafa, 119, 120, 266, 268<br />
rostratay Galathea, 112<br />
Caranx, 171, 183<br />
Orthopristisy 122, 173, 179<br />
ruetzleri, Cymodoce, 227<br />
sabulosa, Hypoconcha, 111<br />
sabulum, Microcharony 91<br />
salpiscinalis, Cyathura (Stygocyathura), 35, 38<br />
samariensis<br />
Edotea, 276<br />
GnafAia, 238, 251<br />
Santiay 98<br />
ffiiV/m, 99, 267<br />
Santiidae, 80, 98<br />
Sargassum, 84, 201<br />
saseboensiSy Limnoria, 195, 198<br />
saxatilis, Abudefdufy 171, 175, 186<br />
sayana, Porcellana, 110<br />
ja>^", Neopanope texana, 111<br />
sbordoniiy Cyathura (Stygocyathura), 35,<br />
38<br />
Schizobopyrina urocaridis, 113, 268<br />
schmittiy Azygopleon, 110, 267<br />
schoepfi<br />
Alutera, 171, 190<br />
Chilomycterusy 172, 190<br />
schroederi, Munida, 112<br />
sciuruSy Haemulon, 172, 179<br />
Sclerocrangon jacqueti, 110<br />
Scomberomorus<br />
cavalla, 173, 187
306 INDEX<br />
Scomberomorus (cont.)<br />
maculatus, 173, 187<br />
regain, 173, 187<br />
scopulosa, Chalixanthura, 27, 29<br />
sedentarius, Chaetodon, 171, 177<br />
Selar crumenophthalmuSy 173, 183<br />
Serolidae, 114, 115, 201<br />
Serolis, 202<br />
mgrayi, 202, 268<br />
Serranus tigrinus, 173, 191, 193<br />
serrata, Spinianirella, 263<br />
serratum, Stenetrium, 100, 102<br />
serricaudus, Carpias, 82, 87<br />
Sesarma ricordi, 111<br />
seticornis, Stenorhynchus, 110<br />
setifer, Macrostylis, 264<br />
setosa, Accalathura, 64, 65<br />
sexticorniSy Excorallana, 161, 165<br />
Amakusanthura, 18, 21<br />
Rocinela, 119, 120, 266, 268<br />
signijica, Amakusanthura, 18, 23<br />
Microcerberus, 244<br />
Munida, 110<br />
simulata, Limnoria, 195, 198<br />
Skuphonura, 17, 58<br />
laticeps, 58<br />
lindae, 267<br />
snanoi, Storthyngura, 263<br />
somala, Haptolana, 138<br />
Sparisoma viride, 122<br />
spathulicarpusy Stenetrium, 100, 104<br />
spatula, Lepisosteus, 172, 190<br />
speculifer, Hirundichthys, 172, 184<br />
specus, Cyathura (Stygocyathura), 35, 38<br />
Speocirolana, 273<br />
Sphaerolana> 273<br />
Sphaeroma, 226, 232, 234<br />
annandalei, 234<br />
destructor, 235<br />
quadridentata, 234, 268<br />
terebrans, 234, 235, 268<br />
ttWfen, 234, 235<br />
<strong>Sphaeromatidae</strong>, 114, 115, 202, 204<br />
Sphaeromatinae, 204, 226<br />
sphaeromiformis, Metacirolana, 153,<br />
154<br />
Sphoeroides maculatus, 173, 190<br />
Sphyraena barracuda, 122<br />
j/>i»ata, Arcturella, 252, 269<br />
Spinianirella serrata, 263<br />
spinosissima<br />
Acanthocope, 263<br />
Hypoconcha, 111<br />
Oncilorpheus, 139<br />
Stenetrium, 100, 104, 267, 270<br />
Stegias clibanarii, 113, 270<br />
Stegophryxus hyptius, 113, 268<br />
steindachneri, Haemulon, 122<br />
<strong>As</strong>trapogon, 171, 188<br />
Parabopyriscus, 267<br />
Stenetriidae, 99<br />
Stenetrioidea, 99<br />
Stenetrium, 99, 100<br />
bowmani, 100<br />
minocule, 100, 102<br />
patulipalma, 100, 102<br />
serratum, 100, 102<br />
spathulicarpus, 100, 104<br />
stebbingi, 100, 104, 267, 270<br />
Stenobermuda, 99, 106<br />
acutirostrata, 106, 270<br />
Stenorhynchus seticornis, 110<br />
stenotelson, Anthomuda, 23, 270<br />
stimpsoni, Munida, 111<br />
stocki, Neostenetroides, 78<br />
pulchra caribbea, 263<br />
snanoi, 263<br />
striata, Yvesia, 246<br />
striatus, Chaetodon, 171, 177<br />
Stygocyathura, see Cyathura (Stygocyathura)<br />
subcaudalis, Eophrixus, 111, 267<br />
subglobosa, Iliacantha, 111<br />
subtilis, Excorallana, 160<br />
sulcipes, Dactylokepon, 267<br />
surinamensis, Batrachoides, 171, 190<br />
surinamicum, Macrobrachium, 112<br />
symmetricus, Ambidexter, 113<br />
synagris, Lutjanus, 172, 183<br />
synalphei<br />
Bopyrione, 110, 267<br />
Hemiarthrus, 111, 267<br />
Synalpheion giardi, 113<br />
Synalpheus<br />
bousfieldi, 110<br />
brevicarpus, I ] 0<br />
AiwwW, 110, 111, 113<br />
fritzmuelleri, 110, 111<br />
goodei, 110, 111<br />
hemphilli, 110, 111
longicarpus, 110, 111, 113<br />
mcclendoni, 110, 111<br />
minus, 110, 113<br />
pandionis, 111, 113<br />
pectiniger, 110, 111, 113<br />
Synodus foe tens, 173, 183<br />
Synsynella, 110, 113<br />
choprae, 113, 268, 270<br />
deformans, 110, 113, 268, 270<br />
Integra, 268<br />
Syringodium, 166, 232<br />
filiforme, 253, 260<br />
syrticus, Microcerberus, 244<br />
tanaiformis, Pendanthura, 56, 270<br />
Tecticeps, 204<br />
Tecticipitinae, 204<br />
/l^, 117, 119<br />
Processa, 113<br />
tenuis, Colanthura, 65, 270<br />
tenuistylis, Lironeca, 171, 186, 187<br />
terebrans, Sphaeroma, 234, 235, 268<br />
ten, Eisothistos, 39<br />
testudinea, Thalassia, 251<br />
texana, Lironeca, 268<br />
texensis, Rhyscotus, 247, 249<br />
Thalassia, 166, 232<br />
testudinea, 251<br />
Thermarcturus, 252, 255<br />
venezuelensis, 255<br />
thomasi, Parabopyrella, 112<br />
floridanus, 110, 111<br />
manningi, 111<br />
f/Wn, Bopyrinella, 110<br />
tigrinus, Serranus, 173, 191, 193<br />
tortugae, Paguristes, 112<br />
tortuganus, Balanopleon, 110<br />
Tozeuma carolinense, 112<br />
transversa, Ceratothoa, 268<br />
transversus, Pachygrapsus, 111<br />
tricarina, Ptilanthura, 267<br />
tricarinata, Booralana, 275<br />
trichostoma, Haptolana, 138<br />
Clibanarius, 110, 112, 113<br />
Holacanthus, 172, 179<br />
tricornis, Excorallana, 161, 165, 266, 268<br />
occidentalis, 167<br />
tricornis, 161, 165<br />
Tridentella, 236<br />
virginiana, 236<br />
Tridentellidae, 115, 235, 236<br />
triospathiona, Gnathia, 238, 241<br />
tripunctata, Limnoria, 199<br />
/n/ow, Carpias, 82, 87<br />
Troglocirolana, 273<br />
tropicalis<br />
Erichsonella filiformis, 2bl<br />
Haplomesus, 263<br />
Lironeca, 268<br />
INDEX 307<br />
tuberculata, Limnoria, 194, 195, 199, 268<br />
tuberculatus, Chiton, 229<br />
tumidicauda, Paradella, 223, 226<br />
tumidipes, Parathelges, 112, 270<br />
Turbinaria, 166, 219<br />
7>/Wj 247, 250<br />
/
308 INDEX<br />
Virganthura, 64, 73<br />
crassa, 73<br />
virginalis, Gnathia, 238, 243<br />
virginiana, Tridentella, 236<br />
viridari, Alpheus, 112<br />
viride, Sparisoma, 122<br />
vittatus, Clibanarius, 110, 112<br />
vulgaris, Palaemonetes, 113<br />
waldneri, Renocila, 173, 191, 193<br />
walkeri, Sphaeroma, 234, 235<br />
warmingii, Excorallana, 161, 167<br />
wegeneri, Tylos, 247, 250<br />
wilsoni, Munnogonium, 96<br />
wolffi, Bopyrissa, 110<br />
wurdemanni, Lysmata, 112<br />
xanthurus, Leiostomus, 172, 183, 187, 190<br />
Xenanthura, 60<br />
brevitelson, 62, 267<br />
A>/