An Updated Classification of the Recent Crustacea

An Updated Classification 

of the Recent Crustacea 

By Joel W. Martin and George E. Davis 

Natural History Museum of Los Angeles County 

Science Series 39 December 14, 2001

AN UPDATED CLASSIFICATION 

OF THE RECENT CRUSTACEA

Cover Illustration: Lepidurus packardi, a notostracan branchiopod from an ephemeral pool in the Central 

Valley of California. Original illustration by Joel W. Marin.

AN UPDATED CLASSIFICATION 

OF THE RECENT CRUSTACEA 

BY 

JOEL W. MARTIN 

AND 

GEORGE E. DAVIS 

NO. 39 

SCIENCE SERIES 

NATURAL HISTORY MUSEUM 

OF LOS ANGELES COUNTY

SCIENTIFIC PUBLICATIONS COMMITTEE 

NATURAL HISTORY MUSEUM 

OF LOS ANGELES COUNTY 

John Heyning, Deputy Director 

for Research and Collections 

John M. Harris, Committee Chairman 

Brian V. Brown 

Kenneth E. Campbell 

Kirk Fitzhugh 

Karen Wise 

K. Victoria Brown, Managing Editor 


Los Angeles, California 90007 

ISSN 1-891276-27-1 

Published on 14 December 2001 

Printed in the United States of America

For anyone with interests in a group of organisms 

as large and diverse as the Crustacea, it is difficult 

to grasp the enormity of the entire taxon at one 

time. Those who work on crustaceans usually specialize 

in only one small corner of the field. Even 

though I am sometimes considered a specialist on 

crabs, the truth is I can profess some special knowledge 

about only a relatively few species in one or 

two families, with forays into other groups of crabs 

and other crustaceans. Crabs are but a small picture 

of the overall diversity of the Crustacea. They represent 

only one infraorder [Brachyura] within one 

order [Decapoda] within one superorder [Eucarida] 

within one subclass [Eumalacostraca] within one 

class [Malacostraca] of the six currently recognized 

classes of the Crustacea (as depicted herein). I am 

certain that this situation is similar for all other 

crustacean systematists, with the result that there 

are no living specialists who can truly claim to have 

an in-depth understanding of the Crustacea as a 

whole. 

This volume is an attempt to provide the reader, 

whether a seasoned systematist or a beginning student, 

with a glimpse into the enormous variety of 

extant crustaceans. The sheer number of categories 

that humans have constructed to contain and order 

this group is some indication of the incredible 

amount of morphological diversity they exhibit. 

But this is only a small part of the overall picture. 

Even if one were to grasp the full range of taxonomic 

diversity as presented in this classification, 

PREFACE 

such knowledge would shed no light on the actual 

biology of these fascinating animals: their behavior, 

feeding, locomotion, reproduction; their relationships 

to other organisms; their adaptations to the 

environment; and other facets of their existence 

that fall under the heading of biodiversity. 

By producing this volume we are attempting to 

update an existing classification, produced by Tom 

Bowman and Larry Abele (1982), in order to arrange 

and update the Crustacea collection of the 

Natural History Museum of Los Angeles County. 

This enormous and diverse collection contains an 

estimated four to five million specimens, making it 

the second largest collection of Crustacea in the 

Americas. While undertaking this task, it occurred 

to us that others might benefit from our efforts, and 

that perhaps a general update on the number and 

arrangement of the living crustacean families, along 

with an explanation of the systematic and classificatory 

changes suggested during the last two decades, 

might be a welcome addition to the literature. 

I hope this volume is seen as nothing more 

than the briefest of introductions into an understanding 

of crustaceans and that it might lead to 

further work not only on the relationships among 

crustaceans but also toward understanding the 

overall picture of crustacean biodiversity and natural 

history. 

Joel W. Martin 

June 2001 

Los Angeles, California

We sincerely thank the many carcinologists to 

whom we sent earlier versions of the classification 

(all of whom are listed in Appendix II). Although 

not all of these persons responded to our queries 

(we had a response rate of approximately 60% to 

the first mailing and approximately 70% to the second) 

and some saw only later versions, we felt it 

appropriate to list all persons from whom comments 

were solicited. Drs. Rodney Feldmann and 

Geoffrey Boxshall, in addition to commenting on 

sections of the classification, served as external referees 

for the entire manuscript, and to both we are 

extremely grateful. We mourn the loss of Erik Dahl 

in January 1999, of Mihai Băcescu in August 1999, 

of Arthur Humes and Austin Williams in October 

1999, of Gary Brusca and Ray Manning in January 

2000, of Théodore Monod in November 2000, and 

of Denton Belk in April 2001, during the compilation 

of this classification. Their absence is keenly 

felt by all carcinologists. Deserving of special recognition 

are David K. Camp for supplying much 

needed information and literature for a wide variety 

of taxa; Anne C. Cohen for literature on ostracodes 

and maxillopods and for enlightening discussions 

of that group’s presumed monophyly; William 

Newman and Mark Grygier, both of whom provided 

literature and enlightening comments on maxillopods; 

Mark Grygier for additional comments on 

interpretation of ICZN recommendations; Trisha 

Spears and Cheryl Morrison for providing unpublished 

molecular sequence or gene rearrangement 

data for the decapods; Geoffrey Fryer for his always 

direct comments concerning the branchio- 

ACKNOWLEDGMENTS 

pods; Gary Poore for information and literature on 

several peracarid and decapod groups and for his 

detailed review of our penultimate draft; Robert 

Hessler for providing needed literature and for his 

insightful suggestions; and Lipke Holthuis for suggesting 

corrections to several taxonomic authorities 

and dates in our earlier versions. Obviously, not all 

of the suggestions we received were incorporated, 

in part because some suggested changes contradicted 

others and in part because some suggested 

changes would have involved major rearrangements 

for which we deemed the evidence insufficient 

or incomplete. Inclusion of crustacean-related 

web sites as an appendix was the idea of Keith 

Crandall. We thank Todd Zimmerman, Regina 

Wetzer, Todd Haney, and Sandra Trautwein in our 

Los Angeles crustacean laboratory for suggestions 

and assistance at various points; Regina Wetzer in 

particular was instrumental in assembling Appendix 

III. 

We thank the Natural History Museum of Los 

Angeles County, and especially John Heyning, John 

Harris, and the members of the Scientific Publications 

Committee, for support and for assistance 

with readying the manuscript for publication. We 

also thank the National Science Foundation for 

partial support via grants DEB 9020088, DEB 

9320397, and DEB 9727188 to J. W. Martin; NSF 

Biotic Surveys and Inventories grant DEB 9972100 

to T. L. Zimmerman and J. W. Martin; and NSF 

PEET grant DEB 9978193 to J. W. Martin and D. 

K. Jacobs. Finally, we sincerely thank Sue, Alex, 

and Paul Martin and Ruthe Davis for their kind 

encouragement and understanding.

CONTENTS 

Preface ......................................................................................................................... v 

Acknowledgments ......................................................................................................... vii 

General Introduction....................................................................................................... 1 

Methods ................................................................................................................... 3 

Names, Dates, and the ICZN......................................................................................... 3 

Cladistics and Classification of the Crustacea ..................................................................... 5 

Molecular Systematics and Classification of the Crustacea ..................................................... 7 

Developmental Genetics and Classification of the Crustacea ................................................... 8 

Sperm Morphology and Classification of the Crustacea......................................................... 8 

Larval Morphology and Classification of the Crustacea......................................................... 9 

The Fossil Record and Classification of the Crustacea.......................................................... 10 

A Note on the Appendices ........................................................................................... 10 

Rationale .................................................................................................................... 12 

Concluding Remarks...................................................................................................... 57 

Classification of the Recent Crustacea ................................................................................ 58 

Literature Cited ............................................................................................................ 76 

Appendix I: Comments and Opinions............................................................................... 102 

Appendix II: List of Contributors .................................................................................... 114 

Appendix III: Other Crustacean Resources......................................................................... 115

An Updated Classification of the Recent Crustacea 

By JOEL W. MARTIN 1 AND GEORGE E. DAVIS 1 

ABSTRACT. An updated classification of the Crustacea down to the level of family is 

provided. The classification is based loosely on that given by Bowman and Abele (1982) 

and includes all new families and higher level taxa described since that time. In addition, 

in several crustacean groupings, new arrangements and assignments have been incorporated, 

based usually on phylogenetic information that has accrued or that has become 

more widely accepted since 1982. Among the more salient changes, some of which are 

more controversial than others, are the recognition of the former phylum Pentastomida 

as a group of maxillopod crustaceans based on additional spermatological and molecular 

evidence, the inclusion of the parasitic Tantulocarida also among the maxillopods, the 

treatment of the Branchiopoda as the most primitive extant group of crustaceans, and 

the recognition of Guinot’s (1977, 1978) division of the higher (eubrachyuran) crabs 

into two ‘‘grades’’ based primarily on placement of the genital aperture. The revised 

classification includes 849 extant families in 42 orders and 6 classes; this is an increase 

of nearly 200 families since the Bowman and Abele classification. More than 90 specialists 

in the field were consulted and asked to contribute to the update. Some workers 

are not in agreement with our final arrangement. In particular, there are questions or 

dissenting opinions over our choice of which taxa to recognize, which authorities and 

dates to credit for various taxa, and especially over the arrangements among and/or 

within the higher taxa. As an aid to future workers in crustacean classification and 

phylogeny, comments and dissenting opinions of some of these workers are appended 

to highlight areas of uncertainty or controversy. Also appended are a list of the specialists 

who were given the opportunity to respond (Appendix II) and a list of printed and 

World Wide Web resources that contain information on crustaceans (Appendix III). The 

new classification is in part a result of one such site, the Crustacean Biodiversity Survey 

(formerly found at URL http://www.nhm.org/cbs/, now temporarily off-line). 

No group of plants or animals on the planet exhibits 

the range of morphological diversity seen among 

the extant Crustacea. This morphological diversity, 

or disparity in the paleontological jargon, is what 

makes the study of crustaceans so exciting. Yet it is 

also what makes deciphering the phylogeny of the 

group and ordering them into some sort of coherent 

classification so difficult. Because of the great age 

of the group, extending back at least as far as the 

early Cambrian and almost certainly beyond that, 

there has been ample time for endless experimentation 

with form and function. The result of these 

many millions of years of evolution is quite dazzling. 

The current estimate of the number of described 

species is approximately 52,000 (Land, 

1996; Monod and Laubier, 1996). This estimate is 

surely on the low side, as a recent estimate of the 

1 Natural History Museum of Los Angeles County, Research 

and Collections, Department of Invertebrate Zoology, 

900 Exposition Boulevard, Los Angeles, California 

90007 

Email: jmartin@nhm.org and gdavis@nhm.org 

GENERAL INTRODUCTION 

number of living species of ostracodes alone is 

10,000 to 15,000 (K. Martens, pers. comm., and 

discussions on the electronic ostracode listserver 

OSTRACON@LISTSERV.UH.EDU) and Kensley 

(1998) has estimated more than 54,000 for the reefassociated 

peracarids. Among the Metazoa, the estimate 

of 52,000 species places crustaceans fourth, 

behind insects, molluscs, and chelicerates, in terms 

of overall species diversity. But morphological diversity 

(disparity) is higher in the Crustacea than in 

any other taxon on Earth. There are probably few 

other groups of animals (squids come to mind because 

of Architeuthis) in which the difference in 

maximum size of adults can be a factor of 1,000. 

The known size of crabs now ranges from a maximum 

leg span of approximately 4minthegiant 

Japanese spider crab Macrocheira kaempferi and a 

maximum carapace width of 46 cm in the giant 

Tasmanian crab Pseudocarcinus gigas (as cited in 

Schmitt, 1965) to a minimum of 1.5 mm across the 

carapace for a mature ovigerous female pinnotherid, 

Nannotheres moorei, the smallest known spe-

cies of crab (Manning and Felder, 1996). An ovigerous 

hermit crab (probably genus Pygmaeopagurus) 

with a shield length of only 0.76 mm taken 

from dredge samples in the Seychelles (McLaughlin 

and Hogarth, 1998) might hold the record for decapods, 

and of course much smaller crustaceans exist. 

Tantulocarids, recently discovered parasites found 

on other deep-sea crustaceans, are so small that 

they are sometimes found attached to the aesthetascs 

of the antennule of copepods; the total body 

length of Stygotantulus stocki is only 94 �m ‘‘from 

tip of rostrum to end of caudal rami’’ (Boxshall and 

Huys, 1989a:127). In terms of biomass, that of the 

Antarctic krill Euphausia superba has been estimated 

at 500 million tons at any given time, probably 

surpassing the biomass of any other group of 

metazoans (reviewed by Nicol and Endo, 1999). In 

terms of sheer numbers, the crustacean nauplius 

has been called ‘‘the most abundant type of multicellular 

animal on earth’’ (Fryer, 1987d). Crustaceans 

have been found in virtually every imaginable 

habitat (see Monod and Laubier, 1996), have been 

mistaken for molluscs, worms, and other distantly 

related animals, and continue to defy our attempts 

to force them into convenient taxonomic groupings. 

Indeed, there is still considerable debate over 

whether the group is monophyletic (see below). 

Not surprisingly, the history of crustacean classification 

is a long and convoluted one. A summary 

of that history is well beyond the scope of this paper, 

and the reader is referred to the following publications 

as some of many possible starting points: 

Schram (1986); Fryer (1987a, c); Dahl and Strömberg 

(1992); Spears and Abele (1997); Rice (1980); 

Schram and Hof (1998); Monod and Forest (1996); 

and papers in the edited volumes The Biology of 

Crustacea (1982–1985; D. E. Bliss, editor-in-Chief) 

(especially volume 1); Crustacean Issues (F. R. 

Schram, general editor); Arthropod Fossils and 

Phylogeny (G. D. Edgecombe, editor); Traité de 

Zoologie (P.-P. Grassé, series editor; J. Forest, crustacean 

volumes editor); and the Treatise of Invertebrate 

Paleontology (R. C. Moore, editor) (a revision 

of this last work is currently underway). Despite 

the long history of studies on Crustacea, in 

many ways, we are just beginning our journey. New 

and significant finds continue to delight and surprise 

the student of the Crustacea. In the last two 

decades, the newly discovered taxa Remipedia, 

Tantulocarida, and Mictacea, as well as beautifully 

preserved fossils from the ‘‘Orsten’’ fauna of Sweden, 

are some of the more obvious examples. Another 

striking example of how little we know about 

crustaceans is the relatively recent discovery of an 

entirely new phylum of animal life, the Cycliophora 

(Funch and Kristensen, 1995; Winnepenninckx et 

al., 1998), found living on the mouthparts of the 

Norway lobster Nephrops norvegicus, a species of 

commercial importance that is encountered often in 

European restaurants. 

The 1982 classification of the Recent Crustacea 

by T. E. Bowman and L. G. Abele, in turn based to 

a large extent on that of Moore and McCormick 

(1969), was a benchmark compilation that has 

been of tremendous use to students of the Crustacea. 

In that classification, the extant crustaceans 

were divided among 6 classes, 13 subclasses, 38 orders, 

and 652 families. Although it was recognized 

by Bowman and Abele and other workers in the 

field, even at the time of publication, that the classification 

was intended to be little more than a stopgap 

measure, it has continued to be employed in 

many major treatments of crustaceans (e.g., Barnes 

and Harrison, 1992; Young, 1998) and has widely 

influenced the study of crustaceans since its appearance. 

Subsequent to the appearance of the 

Bowman and Abele (1982) classification, a large 

number of new families and even some higher level 

taxa have been described. Indeed, our current list 

includes 849 families, an increase of 197 families 

over the Bowman and Abele (1982) classification. 

Thus, an argument could be made that an updated 

classification is warranted on the basis of the increased 

number of new families alone. A more 

compelling reason is that several major treatises 

have appeared that offer substantially different arrangements 

of those taxa and that many exciting 

areas of phylogenetic research and improved methodology 

have contributed significantly to our understanding 

of the relationships within the Crustacea 

and of the Crustacea to other arthropod 

groups. 

While attempting to arrange the collections at the 

Natural History Museum of Los Angeles County, 

the second largest collection of crustaceans in the 

United States, we decided to update the Bowman 

and Abele (1982) classification by simply inserting 

the taxa described since that time. This proved to 

be a more difficult task than we originally envisioned. 

In part this was because the number of new 

taxa was larger than we first thought. And, in part, 

it was because there have been so many suggestions 

for new arrangements and groupings of crustacean 

assemblages, and we wanted to reflect some of the 

recent thinking in crustacean phylogeny in the arrangement 

of our museum’s collection. At about the 

same time, we announced a World Wide Web product 

(http://www.nhm.org/cbs/) called the Crustacean 

Biodiversity Survey (Martin, 1996). The Survey 

was designed to allow workers from anywhere 

in the world to add information at a variety of levels 

to a database on crustacean biodiversity. The 

currently proposed classification is one result of 

that survey. 

Lines have to be drawn at certain times in order 

to attain some level of completion. We received the 

suggestion from several workers to take the classification 

down to the level of subfamily; one worker 

even suggested we include a list of all known genera 

for each family. Others suggested that we provide 

a clear diagnosis and/or characters that distinguish 

each taxon or at least each major clade. Although 

these additions would undoubtedly be extremely 

helpful, for what we hope are obvious reasons, we 

2 � Contributions in Science, Number 39 General Introduction

did not want to attempt it. We are also aware that 

there are a number of works in progress that will 

have a bearing on our understanding of the classification 

of Crustacea (future volumes of the Traité 

de Zoologie [J. Forest, editor] and the ongoing revision 

of the Crustacea sections of the Treatise on 

Invertebrate Paleontology [edited by R. L. Kaesler, 

University of Kansas] are examples of works we 

have not yet seen). However, the field is moving 

rapidly, and we felt that there was more merit to 

publishing what we have than in waiting for additional 

analyses and publications to appear. We are 

also aware of the relatively recent suggestions to 

replace Linnaean hierarchical taxonomy and classification 

with a more phylogenetically based system. 

A brief review by Milius (1999, Science News, 

vol. 156: 268) outlines the controversy as presented 

at the International Botanical Congress meetings in 

St. Louis (see also de Queiroz and Gauthier, 1994; 

Hibbett and Donoghue, 1998; Cantino et al., 1999; 

Cantino, 2000; Nixon and Carpenter, 2000; Meier 

and Richter, 1992; and the web site for the 

PhyloCode at www.ohiou.edu/phylocode/). Some 

authors have even advocated doing away with species 

names as a supposedly logical consequence of 

using phylogenetic taxonomy (e.g., Pleijel and 

Rouse, 2000). However, we have retained a more 

classical approach for now. 

METHODS 

To arrive at the present classification, we began by 

incorporating all of the changes or rearrangements 

of which we were aware. Mostly, because of our 

own taxonomic interests and the strengths of the 

Crustacea collection of the Natural History Museum 

of Los Angeles County, this meant the changes 

or updates within the Decapoda and Branchiopoda. 

In addition, we scanned the following journals 

from 1982 until the present: Crustaceana, Journal 

of Crustacean Biology, Proceedings of the Biological 

Society of Washington, Smithsonian Contributions 

in Zoology, Contributions in Science of the 

Natural History Museum of Los Angeles County, 

Researches on Crustacea (now Crustacean Research), 

and Journal of Natural History. Knowing 

that these journals would not provide a complete 

account of the many changes and additions suggested 

since 1982, we then endeavored to solicit the 

input of a large number of crustacean systematists 

from around the world. Any measure of completeness 

is due to the considerable help and input given 

by these workers (Appendix II). At the same time, 

we accept the responsibility and inevitable criticism 

that any such undertaking generates, as final decisions 

were made by us. 

After incorporating comments received from the 

first mailing of the updated classification, we again 

sent the classification back to the same carcinologists 

and also to several other workers whose 

names had been suggested to us. Finally, in a third 

mailing, we asked those same workers (again, with 

some new names added to the list) to send us additional 

corrections and also their comments, supportive 

or otherwise, concerning the resulting classification, 

with the promise that we would try to 

publish these comments verbatim as Appendix I. In 

this way, we hope to point out areas of disagreement 

and existing controversies in the ‘‘current’’ 

classification such that future workers will know 

that what is presented here as a classification is 

merely a suggested starting point and that there is 

considerable room for improvement. 

Not all workers responded. Some responded only 

to the first mailing, others only to the second or 

third. And of course not all persons listed in Appendix 

II received all three of the mailings. It is 

important to note that the listing of a name in Appendix 

II does not necessarily imply agreement with 

the new classification, regardless of whether a dissenting 

opinion has been offered. We also received 

a large number of positive comments and letters of 

encouragement. 

The present classification will not be accepted by 

all current workers and is sure to be considered 

obsolete almost immediately. Yet we have found the 

Bowman and Abele (1982) classification to be of 

such help, in everything from organizing our museum 

collections to searching for taxa with which 

we are unfamiliar, that we hoped to provide a similar 

and updated tool that would be of at least some 

usefulness for students of the Crustacea. 

As concerns the authorship of this paper, it is 

pertinent to note that G. E. Davis has been responsible 

for the overall organization, tracking, and dissemination 

of information from the beginning of 

this project. Thus, any and all errors or oversights 

concerning the actual classification itself or concerning 

the rationale behind the choices, the literature 

reviewed and cited, and the introductory text 

are the responsibility of J. W. Martin. 

NAMES, DATES, AND THE ICZN 

The Introduction section of the fourth edition of 

the International Code of Zoological Nomenclature 

(ICZN, 1999:xix) states that the Code ‘‘does not 

fully regulate the names of taxa above the family 

group.’’ This is, as we understand it, an intentional 

move designed to allow for some flexibility in establishing 

higher order taxa. Because of this flexibility, 

there are different schools of thought for recognizing 

the names of higher taxonomic categories 

and for crediting the names and dates of these higher 

taxa. One school of thought would advocate that 

a different name (and thus a different person and 

date) should be used each time the constituency of 

the taxon is altered. Thus, for example, if the thalassinoid 

families are removed from the Anomura, 

then we should no longer use the term Anomura 

(or use it in a newly restricted sense) to describe the 

remaining (nonthalassinoid) members of that assemblage. 

Using another example, if we persist in 

keeping the taxon name Eumalacostraca and yet 

Contributions in Science, Number 39 General Introduction � 3

exclude the hoplocarids (stomatopods) from the 

group, we should not credit the name to Grobben, 

who originally coined the name but considered the 

hoplocarids to be within the Eumalacostraca. Such 

changes seem to us to detract considerably from 

stability and can result in a plethora of new names 

being proposed for major taxa that essentially have 

changed very little. An example might be the Achelata 

of Scholtz and Richter (1995), proposed for 

what is essentially the Palinura if the family Polychelidae 

is removed. 

The second school of thought maintains that stability 

is perhaps more valuable than strict accuracy 

and that there is no need to change (for example) 

the name Isopoda simply because the tanaidaceans 

were once included but have since been removed, 

or to discontinue use of Eumalacostraca because 

the stomatopods have been removed, or to change 

the Anomura to Anomala because the thalassinoids 

have been removed. The latter example was discussed 

at length by McLaughlin (1983b), who originally 

advocated using the term Anomala, rather 

than Anomura, for this reason. Later, McLaughlin 

and Holthuis (1985) argued for stability and for 

maintaining the use of the familiar name Anomura. 

For these reasons, and because the Code reminds 

us in the Introduction (ICZN, 1999) that ‘‘nomenclatural 

rules are tools that are designed to provide 

the maximum stability compatible with taxonomic 

freedom,’’ we side with the second school of 

thought. Certainly, at lower taxonomic levels, we 

would never advocate changing the name of a family 

or genus because of the transfer or synonymy of 

a single species, and similarly we are hesitant to do 

away with well-established higher names because 

their constituency has been slightly altered. Thus, 

for the most part, we have tended to retain a wellrecognized 

taxonomic name in favor of a new one 

that differs slightly in its composition. 

Another area of controversy is in the crediting of 

higher taxon names to the original author of the 

group vs. crediting them to the first person to use 

the name in its new, higher, context. For example, 

the ostracode family Darwinulidae is usually credited 

to Brady and Norman (1889). These authors 

did not use it to describe any higher taxon, and it 

was Sohn (1988) who first established the suborder 

Darwinulocopina (based on this family). Should we 

refer to the Darwinulocopina Brady and Norman 

or to the Darwinulocopina Sohn? The ICZN offers 

some guidelines for resolution of this problem at 

lower levels via article 50.3.1 (ICZN, 1999:53). 

This article states that ‘‘the authorship of the name 

of a nominal taxon within the family group, genus 

group or species group is not affected by the rank 

at which it is used.’’ This clearly applies only to 

those mentioned taxonomic levels, and so it does 

not necessarily need to be invoked for the name of 

a family that has been elevated to the rank of superfamily 

(or higher). However, in an attempt to be 

as consistent as possible, Dr. Lipke Holthuis (who 

not only is one of the most prolific writers on crus- 

tacean systematics in history but also has served on 

the International Commission of Zoological Nomenclature) 

has suggested that we extend that recommendation 

to higher levels for those cases where 

it was clear to us that the higher taxon had been 

based on a lower one. Thus, in the above example 

where the family Darwinulidae has been elevated 

to superfamily and even to suborder, we might continue 

to recognize Brady and Norman as the author 

of both of those higher taxa. Holthuis (1993a) also 

mentioned ICZN Article 36a (now 36.1), and as an 

example cited the fact that the ‘‘family name Palaemonidae, 

subfamily name Palaemoninae and the 

superfamily Palaemonoidea, all have as the author 

Rafinesque, 1815.’’ The Editorial Preface to the 

Treatise on Invertebrate Paleontology (Moore, 

1969:xi–xxxvi) stated this in a slightly different 

way, and we quote from it: 

All family-group taxa having names based on the same 

type genus are attributed to the author who first published 

the name for any of these assemblages, whether 

tribe, subfamily, or family (superfamily being almost 

inevitably a later-conceived taxon). Accordingly, if a 

family is divided into subfamilies or a subfamily into 

tribes, the name of no such subfamily or tribe can antedate 

the family name. Also, every family containing 

differentiated subfamilies must have a nominate (sensu 

stricto) subfamily, which is based on the same type genus 

as that for the family, and the author and date set 

down for the nominate subfamily invariably are identical 

with those of the family, without reference to 

whether the author of the family or some subsequent 

author introduced subdivisions. 

The negative side to following this advice (in the 

above case, using the taxon names Darwinulidae 

Brady and Norman and also Darwinulocopina Brady 

and Norman) is that some ‘‘bibliographic’’ and 

historical information is lost. The reader will know 

the original source of the name but will have a very 

difficult time discovering who first employed that 

name as a superfamily, suborder, or higher taxon 

and when this was first done. Using the name ‘‘Darwinulocopina 

Sohn, 1988’’ is therefore more informative, 

if not strictly in keeping with ICZN 50.3.1. 

Holthuis (1993a) was aware of this as well, stating: 

‘‘One could, in keeping with the rules for the family 

names, consider the authors of the family name to 

be at the same time the author of the name of these 

higher categories, but it seemed more logical to cite 

as their author the first zoologist who used such a 

name for a category above the family group level.’’ 

There are also cases in which the higher taxon was 

clearly used and described separately, by different 

authors, rather than being an ‘‘elevation’’ of a family 

name. For example, within the Peracarida, the 

family Mictocarididae is correctly credited to Bowman 

and Iliffe (1985), whereas the order Mictacea 

is credited to Bowman et al. (1985), who established 

the order in a companion paper in the same 

issue of the journal. For these reasons, the choice 

of author and date following a taxonomic name 

might at first seem arbitrary, but we have endeav- 


ored to credit the person or persons who first used 

that name in its new (higher) context when this information 

was known to us. In other instances 

where we were unsure or where we could not personally 

check the original literature, we have employed 

the oldest known name and date, more in 

keeping with the suggestion by Holthuis (pers. 

comm.) to extend ICZN 50.3.1 to higher categories. 

Thus, the present classification, like many others 

before it, is something of an unfortunate mix of 

‘‘rules’’ used to credit authors and dates with the 

establishment of taxa. M. Grygier (pers. comm.) informs 

us that the above discussion is slightly misinformed 

in that the term ‘‘family group’’ explicitly 

includes superfamilies (ICZN article 35.1), such 

that the real difficulty should be only at the level of 

suborder (or any level above that of superfamily). 

One of the specific suggestions we received from 

several workers was a plea to credit Latreille (1803) 

for a large number of higher level crustacean taxa 

(we had used the date 1802 in earlier editions of 

the classification). These taxa include Ostracoda, 

Malacostraca, Gammaridae (and thus Gammaridea), 

Oniscidea (and thus Oniscoidea), Astacidea 

(and thus Astacoidea), Palinura, Paguroidea, Brachyura, 

Squilloidea, and many more. Our choice of 

1802 instead of 1803 is based on the following information 

quoted from a letter we received from L. 

Holthuis (pers. comm., 13 July 1998) referring to 

an earlier draft of our classification: 

Some of Latreille’s names proposed in his Histoire naturelle 

générale et particulière des Crustacés et des Insectes, 

vol. 3 . . . have been cited with the year 1802 

. . . others have the year 1803. The year of publication 

of vol. 3 of Latreille’s work was studied by the best 

authority on Latreille, namely C. Dupuis, who in 1975 

(Bulletin of Zoological Nomenclature, 32: 4) stated 

that this vol. 3 was published after April 1802 and before 

6 November 1802, thus definitely in 1802. Therefore 

all the author’s names ‘Latreille, 1803’ should be 

changed to ‘Latreille, 1802.’ 

Similarly, unless we had fairly convincing evidence 

to the contrary, in those cases where we were 

faced with a choice of different dates (which usually, 

although not always, meant also different authors, 

such as White, 1850 vs. Dana, 1853 vs. Harger, 

1879, all suggested to us by different workers 

as the correct author/date of the isopod family Limnoriidae) 

for the establishment of a taxon, we went 

with the earliest date. In this particular example, at 

least, it proved the correct choice, as White (1850) 

is indeed the author of the family Limnoriidae (G. 

Poore, pers. comm.). 

Finally, we wish to caution readers that we have 

not been able to research each name to the degree 

that we would have liked, and we have depended 

instead upon the many contributors (not all of 

whom were in agreement). Consequently, we would 

advise any user of this (or any other) classification 

to take the time necessary to research carefully the 

history of each taxonomic name for his- or herself, 

which, because of the sheer number of names in- 

volved in this project, we simply were not able to 

do. 

CLADISTICS AND CLASSIFICATION OF THE 

CRUSTACEA 

Ideally, a classification should accurately reflect the 

phylogenetic history of the group. We are very 

much in favor of following rigorous cladistic analyses 

wherever possible, and some of the newly proposed 

classification reflects phylogenetic hypotheses 

based on cladistic analysis of morphological and/or 

molecular data. However, saying that we favor 

classifications based on rigorous cladistic methods 

is not the same as saying that any cladistic analysis 

is more correct than every preceding hypothesis of 

crustacean phylogeny. We wish to state this more 

clearly so that there can be no mistaking our meaning: 

A phylogeny is not correct simply because it 

was generated using cladistics. This somewhat obvious 

point is quite often overlooked. The advantage 

that cladistics imparts is the objective use of 

synapomorphies to define clades. Cladistics is a 

powerful tool, and, like all such tools, it must be 

wielded carefully. And, as with any other tool, there 

is never any guarantee that the result is ‘‘correct.’’ 

We received numerous suggestions that we employ 

a ‘‘more cladistic’’ approach to our new classification. 

For many crustacean assemblages, there have 

been no proposed phylogenies, cladistic or otherwise. 

For other groups, although cladistic methods 

may have been used, there are no published or accessible 

data for confirmation of the results, and/or 

the proposed phylogenies are in stark contrast with 

large literatures on fossil, morphological, developmental, 

or molecular studies of these taxa, making 

them, at least to us, suspect. Two taxa that demonstrate 

this problem are the Maxillopoda and the 

Decapoda, for which some of the most vocal proponents 

of cladistic approaches gave us quite different 

suggestions for the classification, all supposedly 

based on rigorous cladistic analyses of ‘‘good’’ 

data. Similar frustration concerning recent attempts 

to cladistically analyze fossil arthropods is expressed 

by Fryer (1999c). More troubling still is 

that there are other cladistic analyses of which we 

are aware, and that appear to be based on solid 

evidence, that we could not follow completely because 

to do so would have orphaned large numbers 

of families. For example, we do not doubt the revelation 

by Cunningham et al. (1992) that king crabs 

of the family Lithodidae are actually nested within 

one clade of hermit crabs (but see McLaughlin and 

Lemaitre, 1997, 2000, for a dissenting opinion). 

But there are other clades of hermits and other species 

of lithodids that were not part of this study, 

and we hesitated to make sweeping changes before 

all evidence is in. Another example concerns dromiacean 

crabs, traditionally placed among the lower 

Brachyura but whose larvae appear distinctly anomuran. 

The molecular analysis of Spears et al. 

(1992) grouped at least one dromiid with the An- 


omura rather than the Brachyura—but does this 

hold for all crabs in the former Dromiacea? Thus, 

we have in some instances knowingly presented 

groupings for which contrary evidence exists for at 

least some of the constituent taxa. We have tried to 

mention all such areas in the text of the Rationale 

section that follows. Several workers noted this 

problem and suggested that perhaps no classification 

should be attempted until such time that we 

have better supported phylogenetic analyses in 

hand for all (or at least most) crustacean groups. 

There is merit to this argument. But in keeping with 

our original goal of updating a classification of the 

entire assemblage to benefit students who wish to 

view the overall picture of crustacean diversity, we 

felt that waiting would not improve the situation. 

An additional practical problem faced by the student 

wishing to construct a cladistically based classification 

is the very real difficulty of representing 

complex relationships in a two-dimensional classification. 

To accurately depict all of the branching 

relationships and show all of the sister groupings 

would necessitate a rather large number of additional 

taxonomic categories. One proposed solution 

is to simply indent the families in the list (without 

creating additional names for groupings) to imply 

the relationships. But even this is difficult when 

dealing with the number of families in, for example, 

the gammaridean amphipods or the harpacticoid 

copepods. Another proposed solution is to completely 

abandon Linnaean hierarchical classifications 

in favor of a more phylogenetically based system 

(e.g., see Milius, 1999; Cantino et al., 1999). 

We feel that, in many cases, a ‘‘standard’’ classification—that 

is, a simple list of families—still serves 

a purpose for those taxa where the phylogeny remains 

uncertain (which is nearly every group of the 

Crustacea) in that it at least allows recognition and 

placement within well-defined higher groups for beginning 

students. Thus, while very much in favor 

of the application of cladistic methodology and of 

the construction of classifications based on these 

methods whenever possible, we have had difficulties 

in trying to arrive at a sensible or useful way 

of depicting these relationships to the beginning 

student of carcinology. Consequently, to many 

readers, our current arrangements and ‘‘lists’’ of 

families will appear old fashioned and unsatisfactory. 

The number of phylogenetic studies on the Crustacea 

has risen dramatically since Bowman and 

Abele’s (1982) classification. Christoffersen (1994: 

135) estimated that 123 cladistic analyses of crustaceans 

had appeared in print as of the end of 1992, 

and that number has increased dramatically since 

then. Reasons for the increase include improved 

methods of computation and the availability of cladistic 

programs, such as PAUP, McCLADE, and 

HENNIG 86, in addition to the growing acceptance 

of cladistics as a preferred way of thinking 

about and depicting crustacean relationships and 

relationships of all other groups as well (see papers 

cited in Nielsen, 1995, and Nielsen et al., 1996). 

Recent phylogenetic software is reviewed by Eernisse 

(1998), and a list of phylogenetic programs 

by categories is provided on J. Felsenstein’s ‘‘Phylogenetic 

Programs’’ web site at http://evolution. 

genetics.washington.edu/phylip/software.html# 

methods. The fact that cladistics is almost routinely 

employed in studies of crustacean relationships today 

can be credited largely to the efforts of F. R. 

Schram (e.g., see Schram, 1983a, and papers therein; 

Schram, 1986; Schram and Hof, 1998). Although 

it is beyond the scope of this project to review 

the many cladistic analyses of crustacean 

groups that have appeared since 1982, we list below 

a few of the more salient papers that treat crustaceans 

above the level of family, with the hope that 

this might form something of an introduction to the 

literature for students of crustacean phylogeny. The 

list is not intended to be exhaustive. Instead, we 

hope it alerts readers to the fact that very little is 

settled with regard to crustacean relationships and 

classification and to the fact that cladistic thinking 

has profoundly affected our understanding of crustacean 

relationships. 

In alphabetical order within chronological order, 

these works include: Briggs (1983, Cambrian arthropods 

and crustaceans [see also Briggs and 

Whittington, 1981]), Grygier (1983a, b, maxillopodans), 

Sieg (1983a, tanaidaceans), Takeuchi 

(1993, caprellidean amphipods), Wheeler et al. 

(1993, arthropods including crustaceans), Ho 

(1984, nereicoliform copepods), Schram (1984a, 

Eumalacostraca; 1984b, Syncarida), Martin and 

Abele (1986, anomuran decapods), Schram (1986, 

all crustacean groups), Christoffersen (1986, 1987, 

caridean shrimp), Grygier (1987a, b, maxillopodans), 

Pires (1987, peracarids), Christoffersen 

(1988a, b, caridean shrimp), Müller and Walossek 

(1988, Maxillopoda), Abele et al. (1989, pentastomids), 

Boxshall and Huys (1989a, maxillopodans), 

Briggs and Fortey (1989, Cambrian arthropods 

including crustaceans), Christoffersen (1989, 

caridean shrimp), Schmalfuss (1989, oniscidean 

isopods), Brusca and Brusca (1990, all crustacean 

groups), Christoffersen (1990, Caridea), Ho (1990, 

copepod orders), Kim and Abele (1990, decapods), 

Walossek and Müller (1990, ‘‘stem line’’ crustaceans), 

Abele (1991, decapods), Brusca and Wilson 

(1991, isopods), Abele et al. (1992, maxillopodan 

groups), Briggs et al. and Briggs and Fortey (1992, 

Cambrian arthropods including crustaceans), Høeg 

(1992a, maxillopodans), Spears et al. (1992, brachyuran 

crabs), Walossek and Müller (1992, ‘‘orsten’’ 

fossil crustaceans), Wilson (1992, most major 

extant groups), Kim and Kim (1993, gammaridean 

amphipod families and amphipod suborders), Walossek 

(1993, branchiopods and Crustacea), Poore 

(1994, thalassinideans), Spears et al. (1994, thecostracan 

maxillopodans), Wagner (1994, peracarids), 

Wilson (1994, janiroidean isopods), Glenner et al. 

(1995, cirripedes), Scholtz and Richter (1995, decapods), 

Bellwood (1996, calappid crabs), Humes 


and Boxshall (1996, lichomolgoid copepods), 

Moura and Christoffersen (1996, ‘‘mandibulate’’ 

arthropods), Wilson (1996, isopods), Ahyong 

(1997, stomatopods), Emerson and Schram (1997, 

all arthropods), Hanner and Fugate (1997, branchiopods), 

Spears and Abele (1997, several major 

groups, review), Tshudy and Babcock (1997, 

clawed lobsters), Tudge (1997b, anomurans), Walossek 

and Müller (1997, Cambrian crustaceans and 

their bearing on crustacean phylogeny), Wheeler 

(1997, arthropods including crustaceans), Wills 

(1997, all Crustacea), Jenner et al. (1998, hoplocarids), 

Olesen (1998, conchostracans and cladocerans), 

Schram and Hof (1998, all major groups, 

extant and extinct), Shen et al. (1998, spelaeogriphaceans), 

Strausfeld (1998, crustacean neurological 

features), Taylor et al. (1998, mysidaceans and 

other peracarids), Tucker (1998, raninoid crabs), 

Wheeler (1998, all arthropod groups), Wills et al. 

(1998, fossil and extant arthropod groups), Almeida 

and Christoffersen (1999, pentastomids), Cumberlidge 

and Sternberg (1999, freshwater crabs), 

Huys and Lee (1999, laophontoidean harpacticoid 

copepods), Sternberg et al. (1999, freshwater 

crabs), Olesen (1999b, leptostracans), Spears and 

Abele (1999b, crustaceans with foliaceous limbs; 

2000, branchiopods), Walossek (1999, major crustacean 

groups), Edgecomb et al. (2000, all major 

arthropod groups), Negrea et al. (1999, branchiopods), 

Shultz and Regier (2000, all major arthropod 

groups), and Richter et al. (2001, cladocerans). 

MOLECULAR SYSTEMATICS AND 

CLASSIFICATION OF THE CRUSTACEA 

Without doubt, the most exciting recent developments 

in our understanding of crustacean relationships 

have been in the realm of molecular systematics 

and phylogenetics. Indeed, many of the cladistic 

papers mentioned in the previous section are 

based on molecular sequence data, which essentially 

were not available at the time of the Bowman 

and Abele classification. Molecular systematic studies 

of arthropods have become so numerous that 

Wheeler (1998) stated ‘‘the past decade has presented 

us with nearly annual molecular analyses of 

Arthropoda.’’ For the Crustacea, most of this work 

has been championed by the laboratories of L. G. 

Abele and T. Spears at Florida State University and 

C. W. Cunningham at Duke University. This field, 

as well as the field of developmental genetics 

(which we barely touch upon here), is growing and 

changing at a phenomenal rate. Many of the early 

studies were based on relatively small sequences, so 

it is not terribly surprising that there have been 

some published results that appear unreasonable 

based on our knowledge of morphology, embryology, 

paleontology, and other sets of characters. As 

we refine our selection of which genes to target, 

improve our ability to extract and align increasingly 

larger sequences, and devise better computational 

algorithms, we might begin to see more agree- 

ment between molecular results and more traditional 

views of crustacean phylogeny, or at least 

results that are less ambiguous. Or we may not. As 

Spears and Abele (1997) state in the conclusion to 

their review paper on the use of 18S rDNA data in 

crustacean phylogeny, ‘‘Regrettably, in the crusade 

for understanding relationships among crustaceans 

and other arthropod lineages, the rDNA data represent 

but a relic, and not the Holy Grail itself.’’ 

Yet despite this sobering conclusion, Spears and 

Abele (1997) were able to make some very strong 

statements concerning at least some crustacean 

taxa. For example, Branchiopoda, Copepoda, Podocopida, 

and Myodocopida are all clearly monophyletic; 

the Malacostraca is clearly monophyletic 

and includes the Phyllocarida (Leptostraca) (supported 

also by Shultz and Regier, 2000); Maxillopoda 

does not appear monophyletic (although certain 

groups within it seem to be united); etc. 

There are, of course, known problems associated 

with some of these approaches (as one early example, 

see the responses by Nielsen and others 

(1989) to the article by Field et al. (1988) entitled 

‘‘Molecular analysis of the animal kingdom’’). Fryer 

(1997) points out several papers that question the 

results and/or validity of recent studies of arthropod 

phylogeny based on molecular data; Wägele 

and Stanjek (1995) make the point that alignment 

alone can be responsible for serious discrepancies 

in analyses of such data. And of course the history 

of a particular gene might not accurately reflect the 

phylogeny of the species containing that gene (e.g., 

see Brower et al., 1996; Doyle, 1997; Maddison, 

1997; Page and Charleston, 1998). Unfortunately, 

the branchiopod genus Artemia, which has been 

used for more molecular comparative studies than 

any other crustacean genus, is not the best choice; 

Maley and Marshall (1998) note that ‘‘brine shrimp 

[have] long been known to produce artifactual 

groupings.’’ Lake (1990) admitted that arthropod 

paraphyly as indicated in his analysis may be a result 

of long branch attraction caused by the inclusion 

of Artemia and Drosophila; this problem was 

mentioned also by Turbeville et al. (1991). It is also 

disconcerting that, after so much money and effort 

have been expended toward applying genetic data 

to resolving the evolutionary roots of modern humans, 

we still do not have a clear answer. Whether 

Homo sapiens arose from a single African source 

200,000 years ago or ‘‘multiple groups in Africa 

and elsewhere’’ at least a million years ago is still 

hotly debated (see Bower, 1999). How, then, are we 

expected to place confidence in what the molecules 

are telling us about the evolution of crustaceans 

when our efforts, in comparison, have been so limited? 

To summarize, we again quote Maley and 

Marshall (1998): ‘‘To be confident in our hypotheses 

of relationships among the animal phyla we 

need to gather more DNA sequences, especially 

from undersampled phyla; develop better methods 

of DNA analysis on the basis of more realistic models 

of DNA evolution; and develop independent 


data sets using morphological, developmental, and 

other molecular data to corroborate or falsify specific 

hypotheses or to combine in total-evidence 

analyses.’’ Thus, just as we have not accepted all 

cladistic analyses simply because they were cladistic, 

we have incorporated molecular analyses with 

caution because of perceived problems with some 

of these studies. At the same time, there is little 

question that these efforts, however preliminary 

they may be, represent the first attempts to apply 

‘‘new’’ and objective data to the resolution of crustacean 

phylogeny for the first time in some 200 

years of study, and we look forward to continued 

advances in this field. 

Papers mentioned below are merely examples of 

some of the more comprehensive or influential 

works of which we are aware. As in the previous 

section, we have included only those papers that 

deal with ‘‘higher level’’ crustacean taxa or with the 

relationships of crustaceans to other arthropods. In 

alphabetical order within chronological order, these 

papers include Abele et al. (1989, pentastomids, 

rRNA), Kim and Abele (1990, decapods, 18S 

rRNA), Abele (1991, decapods, 18s rRNA), Turbeville 

et al. (1991, arthropods including crustaceans, 

18S rRNA), Abele et al. (1992, maxillopodans, 

18S rDNA), Cunningham et al. (1992, lithodid 

and pagurid anomurans), Spears et al. (1992, 

brachyuran crabs, 18s rRNA), Wheeler et al. 

(1993, arthropods including crustaceans, 18S 

rDNA, and polyubiquitin), Raff et al. (1994, review 

of arthropod relationships [and other metazoan 

groups] based on various genes), Spears et al. 

(1994, thecostracans, 18S rDNA), Boore et al. 

(1995, arthropods including crustaceans), Friedrich 

and Tautz (1995, arthropods, 18S and 28S rDNA), 

France and Kocher (1996, DNA sequencing of formalin-fixed 

crustaceans), Wray et al. (1996, 6 mitochondrial 

and 2 nuclear genes), Eernisse (1997, 

arthropods [including crustaceans] and annelids, 

18S rRNA), Hanner and Fugate (1997, branchiopods, 

12S rDNA), Regier and Schultz (1997, major 

arthropod groups, two nuclear genes), Spears and 

Abele (1997, all crustacean groups, 18S rDNA), 

Wheeler (1997, most arthropod groups), Boore et 

al. (1998, crustaceans and insects, gene translocations), 

Colgan et al. (1998, arthropods including 

crustaceans, histone H3 and U2 snRNA), Min et 

al. (1998, arthropods, 18S rDNA), Regier and 

Schultz (1998a, b, arthropods, amino acid sequence 

of EF-1�), Schwenk et al. (1998, cladocerans, 16S 

rDNA), Wheeler (1998, arthropods [including crustaceans], 

18S and 28S rDNA), Braga et al. (1999, 

copepods, 16S and 28S rRNA), Morrison and Cunningham 

(1999, anomurans, mitochondrial gene rearrangements), 

Spears and Abele (1999b, crustaceans 

with foliaceous limbs, 18S rDNA), Crandall 

et al. (2000, Astacidea, 18S, 28S, and 16S rDNA), 

Edgecomb et al. (2000, arthropods including crustaceans, 

histone H3 and U2 snRNA sequences), Giribet 

and Ribera (2000, all arthropod groups, 18S 

and 28S rDNA), Harris et al. (2000, barnacles, 18S 

rDNA), Jarman et al. (2000, malacostracans, 28S 

rDNA), Perl-Treves et al. (2000, thecostracans, 18S 

rDNA), Remigio and Hebert (2000, anostracan 

branchiopods, 28S and 16S rDNA), Spears and 

Abele (2000, branchiopods, 18S rDNA), Schubart 

et al. (2000a, b, grapsoid crabs, 16S rDNA), Shultz 

and Regier (2000, arthropods, Ef-1� and Pol II), 

Wilson et al. (2000, Malacostraca, mitochondrial 

DNA and gene order), Mattern and Schlegel (2001, 

oniscidean isopods, ssu rDNA), and Richter et al. 

(2001, Cladocera, 12S rDNA). See also papers in 

the symposium Evolutionary Relationships of 

Metazoan Phyla organized by D. McHugh and K. 

Halanych (1998, American Zoologist 38:813–982) 

and the volume Arthropod Relationships edited by 

R. A. Fortey and R. H. Thomas (1997). 

DEVELOPMENTAL GENETICS AND 


The relatively newly emerging field of developmental 

genetics needs to be mentioned here as well, 

though we hasten to add that this field of study is 

well beyond our area of expertise and that any attempt 

at a synthesis would be premature. Recent 

discoveries concerning especially homeotic (Hox) 

genes and arthropod relationships are having a profound 

influence on our understanding of crustacean 

morphological plasticity and clearly will play an increasingly 

important role in elucidating relationships 

within Crustacea and among the various arthropod 

groups. We include this brief section only 

as a way to signal to the beginning student what is 

surely to be an active field of research for many 

years to come. Some of the recent papers in this 

field with applications to crustacean classification 

include (in alphabetical order) Akam (1998), Akam 

et al. (1994), Arhat and Kaufman (1999), Averof 

and Akam (1993, 1995a, b), Averof and Patel 

(1997), Carroll (1995), Davidson et al. (1995), Fortey 

and Thomas (1997), Grenier et al. (1997), Panganiban 

et al. (1995, 1997), Popadić et al. (1996), 

Roush (1995), Scholtz (1995), Shubin et al. (1997), 

and Williams and Nagy (1995) (some of which are 

briefly reviewed in Brusca, 2000). 

SPERM MORPHOLOGY AND 


Yet another field of research that is improving our 

understanding of crustacean relationships is the description 

and comparison of crustacean sperm, 

termed ‘‘spermiocladistics’’ by Jamieson (1987, 

1991a). While examination of crustacean sperm 

morphology for systematic purposes is not new 

(e.g., Koltzoff, 1906; Wingstrand, 1972, 1978, 

1988; Grygier, 1981, 1982), recent work has employed 

ultrastructural characters that show more 

promise for resolution of long-standing questions. 

In the words of Tudge (1997b), the ‘‘use of spermatozoal 

ultrastructure in taxonomy and phylogeny 

is now firmly established as a valid means of 

investigating phylogenetic relationships in various 


animal phyla.’’ For the Crustacea, these characters 

have been invoked mostly for resolving relationships 

within the Eumalacostraca. This work is being 

championed primarily by B. G. Jamieson and 

C. Tudge and their colleagues. Some of the many 

recent papers advocating sperm ultrastructural 

characters in phylogeny are Guinot et al. (1994, 

primitive crabs; 1997, freshwater crabs; 1998, 

dromiacean crabs), Richer de Forges et al. (1997, 

crabs), Jamieson (1989a, b, crabs; 1989c, stomatopods; 

1990, primitive crabs; 1991a, overview of 

crustacean sperm ultrastructure and phylogeny; 

1991b, 1993, 1994, crabs), Jamieson et al. (1993a– 

c, crabs; 1994a, b, 1995, 1996, 1997, crabs), Jamieson 

and Tudge (1990, crabs), Jamieson, Tudge, 

and Scheltinga (1993, primitive crabs), Jespersen 

(1979, leptostracans), Grygier (1981, 1982, maxillopodans), 

Storch and Jamieson (1992, pentastomids), 

Tudge (1991, 1992, 1995, 1997a, b, 1999a, 

b, anomuran decapods), Tudge et al. (1998a, lithodid 

crabs; 1998b, hydrothermal vent crabs), and 

Tudge et al. (2000, mud-shrimp families; 1999, hippoid 

crabs). Many of these papers and their contributions 

are discussed in the sections dealing with 

the taxa in question. 

Some of the revelations from the study of sperm 

ultrastructure are not terribly surprising and in fact 

support previous long-standing hypotheses of crustacean 

relationships (e.g., peracarid unity; Jamieson, 

1991a). Other results are more controversial 

and include the alliance of the Remipedia with the 

Maxillopoda on the basis of the shared ‘‘flagellate 

condition’’ of their spermatozoon (Jamieson, 

1991a) and placing the genus Lomis outside of, and 

thalassinids within, the Anomura (Tudge, 1997a, b) 

(in contrast with what Morrison and Cunningham, 

1999, presented based on mitochondrial gene rearrangement 

data). [As an aside, the congruence between 

the phylogenetic diagrams of Jamieson 

(1991a:111), based on sperm ultrastructure, and 

Schram (1986), based on cladistic analysis of morphological 

characters, is perhaps not so remarkable 

as Schram and Hof (1998) suggest. Schram and 

Hof (1998) refer to Jamieson’s figure and ask the 

reader to ‘‘note the general correspondence with the 

major classes as arranged in Fig. 6.1.A.’’ However, 

Jamieson’s figure was in turn based on Schram 

(1986) with a diagram of the spermatozoal ultrastructure 

simply added to Schram’s tree; it is not an 

independently derived phylogeny.] Continued use 

of sperm ultrastructure in crustacean taxonomy 

and systematics will almost certainly contribute significantly 

to our understanding of crustacean phylogeny. 

LARVAL MORPHOLOGY AND 


The study of crustacean systematics and phylogeny 

has involved larval characters from the very earliest 

times. For many groups of crustaceans, a study of 

systematic relationships is a study of the larvae, as 

these are often the only characters, or the best characters, 

that we have. For example, it could be argued 

that, until recently, the history of studies in 

barnacle phylogeny has been essentially a history 

of comparisons of barnacle larvae, and to some extent 

this is true for many groups. For some taxa, in 

particular the Facetotecta, the larvae are all that we 

know; the adult has yet to be recognized or described. 

The reverse is also true: there are still some 

important groups of crustaceans (the class Remipedia, 

for example) for which the larval forms have 

never been identified. Many of the classic treatments 

of crustacean larvae were published prior to 

the Bowman and Abele (1982) classification and 

were thus available for consideration by those authors. 

The summary of crustacean larval diversity 

published by Williamson in that same series of volumes 

(Williamson, 1982) remains a good entry 

point for the literature on crustacean larvae and 

relationships based on larval characters. 

In the years following the Bowman and Abele 

(1982) classification, there have been additional 

and significant treatments of crustacean larval characters 

and phylogeny. Indeed, nearly every modern 

publication that describes a larval stage includes at 

least some comments on the applicability of the 

findings to relationships within the group. The 

study of larval crabs, in particular, has been a rich 

source of new characters for postulating higher level 

relationships among the Brachyura (e.g., see 

Rice, 1980, 1981, 1983, 1988; Martin, 1984, 

1988; Martin et al., 1985; Felder et al., 1985, as a 

few selected examples from a huge body of literature 

on crab relationships based on larvae and postlarvae). 

Williamson (1988a, b) has proposed rather 

drastic changes in our understanding of various 

pleocyemate groups (particularly the position of the 

dromiid crabs relative to anomurans and true 

crabs, the placement of the mysidaceans within the 

Eucarida, and the separation of palinurid lobsters 

from other eucarids based on their bizarre larvae). 

Grygier (1987a–c) and others have used larval 

characters to explore maxillopod phylogeny; within 

the Maxillopoda, the work of Dahms (e.g., Dahms, 

1990) could be mentioned for advancing our understanding 

of copepod naupliar characters in phylogeny. 

Discoveries of fossilized larvae, in particular 

papers on the ‘‘Orsten’’ fauna, have added new 

characters and new insights into the evolution of 

early crustaceans and ‘‘stem-line’’ crustaceans (e.g., 

see Müller and Walossek, 1985a, 1986b; Walossek, 

1993, 1995; Walossek and Müller, 1990, 1997). 

Walossek and Müller (1997) recognize the Entomostraca, 

and exclude from the Crustacea the Pentastomida, 

in part based on larval evidence. 

We have tried to mention studies based on larval 

characters (where they have a bearing on classification 

at the family level or higher) under each 

crustacean taxon. A recent review of larval diversity 

(Harvey et al., in press) provides additional material 

geared primarily for the beginning student of 

carcinology. 


THE FOSSIL RECORD AND CLASSIFICATION 

OF THE CRUSTACEA 

No understanding of crustacean diversity and evolution 

would be complete without knowledge of the 

fascinating fossil history of the group. And many 

exciting discoveries that bear on crustacean origins, 

relationships, and classification have surfaced since 

the Bowman and Abele treatment. A recent example 

is the intriguing find of a serolid-like sphaeromatoid 

isopod from the Solnhofen of Germany 

(Brandt et al., 1999), pushing back the origin of 

sphaeromatoid isopods to at least the Late Jurassic. 

Although a thorough review of such discoveries is 

beyond the scope of this report (see papers in Edgecombe, 

1998, and reviews by Delle Cave and Simonetta, 

1991; Bergström, 1992; Schram and Hof, 

1998; Walossek and Müller, 1997, 1998; Wills, 

1998; Wills et al., 1995; Fortey et al., 1997; Fryer, 

1999c), we feel the need to mention especially the 

stem and crown group crustaceans of the ‘‘Orsten’’ 

fauna of Sweden (Orsten-type fossils have also been 

found on other continents; see review by Walossek, 

1999). These works include papers by Müller 

(1982, Hesslandona; 1983, crustaceans with soft 

parts), Müller and Walossek (1985, Skaracarida; 

1986a, Martinssonia; 1986b, various arthropod 

larvae; 1988, the maxillopod Bredocaris), Walossek 

and Müller (1990, stem line crustacean concept; 

1992, overview of the Orsten fauna; 1994, possible 

pentastomids; 1997, 1998, overviews), Walossek 

and Szaniawski (1991, Cambrocaris), Walossek et 

al. (1994, possible pentastomids), and Walossek 

(1993, 1995, the branchiopod Rehbachiella; 1999, 

overview of Cambrian crustaceans). These publications 

include detailed descriptions of several new 

taxa that have in many ways altered our view of 

primitive crustaceans and the timing of crustacean 

evolution. 

The Burgess Shale crustaceans have been reexamined 

recently by Briggs et al. (1994), and the 

remarkable fossil arthropods from the Lower Cambrian 

Chengjiang fauna of southwest China have 

been summarized by Hou and Bergström (1991, 

1997). Included in the Chengjiang fauna are no unequivocal 

crustaceans (Waptia being the only remote 

possibility), but several fossils seem to have a 

bearing on our understanding of crustacean evolution. 

Other recent studies of Chinese fossil crustaceans 

have included papers on conchostracans (e.g., 

Shen, 1984, 1990; Zhang et al., 1990; see also Orr 

and Briggs, 1999, for Carboniferous conchostracans 

from Ireland), and Lower Cambrian crustaceans 

are known from other sites around the world 

as well (e.g., see Butterfield, 1994). Studies of bradoriid 

and phosphatocopid arthropods (once 

thought to be ostracodes) (see Siveter and Williams, 

1997) have even shed light on our understanding 

of the evolution of the crustacean circulatory system 

(Vannier et al., 1997). The phosphatocopids 

are now thought to be close to the ‘‘stem-line’’ crustaceans 

(and possibly the sister taxon to Crustacea; 

see Walossek, 1999) rather than relatives of any of 

the crown-group crustaceans such as ostracodes or 

maxillopods, which had been suggested previously 

(e.g., see reviews by Walossek and Müller, 1992, 

1998). At least two major groups, and possibly 

many more unknown to us, remain enigmatic as to 

whether they belong in the Crustacea or not: Thylacocephala 

(see Pinna et al., 1982, 1985; Secretan, 

1985 [as Conchyliocarida]; Rolfe, 1985, 1992; 

Schram et al., 1999) and Cycloidea (see Schram et 

al., 1997; Schram and Hof, 1998), although cycloids 

were probably allied to the maxillopodans 

(Schram et al., 1997). Schram and Hof presented, 

as part of the Fourth International Crustacean Congress 

(ICC-4) in Amsterdam, evidence that the Thylacocephala 

are indeed crustaceans; they further 

postulate the inclusion of the Thylacocephala in the 

Thecostraca on the basis of the presence of lattice 

organs. Their paper, entitled ‘‘At last: the Thylacocephala 

are Crustacea,’’ was a late addition and 

therefore is not included among the published abstracts 

of the ICC-4 Congress, but since then, the 

information has been submitted (Lange et al., in 

press). However, Schram et al. (1999) are more 

cautious and stopped short of declaring that thylacocephalans 

were crustaceans. The Permian ‘‘pygocephalomorph’’ 

crustaceans and their relationship 

to extant mysidaceans was examined recently 

by Taylor et al. (1998). A thorough review of most 

of the above contributions is presented by Schram 

and Hof (1998). Many other papers on crustacean 

fossils continue to add to our knowledge of the history 

of the group (e.g., Brandt et al., 1999, on the 

Late Jurassic origin of sphaeromatoid isopods). 

In light of these remarkable finds, it is understandable 

that a number of colleagues have suggested, 

some rather strongly, that we incorporate 

fossil taxa into the current classification. We have 

opted not to do so, primarily because we are less 

familiar with the fossil crustacean literature (and 

with workers in that field) than we are with the 

literature on extant groups. Thus, the opportunities 

for us to inadvertently perpetuate or create errors 

would have been much greater had we attempted 

this task. Also, if the currently proposed classification 

proves to have merit, it should not be difficult 

for more paleontologically inclined carcinologists 

to, at some point, add these fossil taxa to the existing 

framework. We hope that the classification is 

of some use to paleontologists and that, at some 

point, we can incorporate fossil taxa into this 

scheme. Relatively recent lists of crustacean fossil 

taxa can be found in Whatley et al. (1993, ostracodes) 

and Briggs et al. (1993, all other crustacean 

groups) (both in M. J. Benton, editor, The Fossil 

Record 2, Chapman and Hall, 1993). However, 

since our knowledge (and time) is limited, we have 

decided to include only extant taxa for now. 

A NOTE ON THE APPENDICES 

APPENDIX I. COMMENTS AND OPINIONS 

After receiving and considering the input from various 

workers around the world, we then asked the 


same persons to comment on the resulting product. 

We did this for two reasons. First, many of the suggestions 

we received were not incorporated, and we 

wanted collaborators to have the opportunity to 

express their disagreement. Reasons for not incorporating 

a particular suggestion were many and 

ranged from simple disagreement on our part to 

conflicting suggestions or corrections from noted 

experts. Second, we wanted students of carcinology 

to know where the major areas of disagreement lie 

in our understanding of crustacean phylogeny and 

classification. By pointing out areas where other experts 

in the field disagree with the current classification, 

we hoped to avoid the impression that the 

classification is accepted or agreed upon by some 

consensus of crustacean taxonomists. 

APPENDIX II: LIST OF CONTRIBUTORS 

The list of persons to whom we sent either first, 

second, or third drafts of the classification is given 

in Appendix II. Some of those listed responded to 

only one of our mailings; some responded to all 

mailings; some workers did not respond at all. No 

person on the list should be assumed to be in agreement 

with the classification as a whole. Despite 

these caveats, we felt that we should list all of the 

workers we attempted to contact to let readers 

know the potential pool of expertise from which 

we solicited input. 

Because of the tremendous interest in the Crustacea 

worldwide, the number of qualified workers 

is much greater than this list indicates. Our decision 

on whose input to solicit was more or less arbitrary, 

based on our own knowledge of workers in the 

field and on suggestions received as a result of the 

first and second mailings. We apologize in advance 

if, by omitting someone from one or more mailings, 

we have inadvertently slighted anyone; such was 

not our intent. 

APPENDIX III: OTHER CRUSTACEAN 

RESOURCES 

Finally, a list of other crustacean resources is provided 

to give the student of Crustacea an introduction 

to the large and ever growing number of crustacean 

resources. The list includes crustacean-specific 

journals, newsletters of special interest groups 

(e.g., Zoea, Ecdysiast, Monoculus, Anostracan 

News, and Cumacean Newsletter), and URLs of 

helpful crustacean-related sites on the World Wide 

Web. 


SUBPHYLUM CRUSTACEA 

Many of the questions considered most pressing today 

have been asked for well over 100 years: Are 

crustaceans a monophyletic group? How many major 

clades, or classes, are there? Which is the most 

primitive class? What are the relationships among 

the classes? We cannot attempt to answer all of 

these questions here, but below we offer a brief explanation 

of how and why we arrived at the current 

classification. In most cases, we provide some additional 

information under the heading for each of 

the various taxa (each of which is treated later). For 

more in-depth discussions of the complex history 

of attempts to classify the Crustacea, we refer the 

reader to the following publications: Moore and 

McCormick (1969), Schram (1986), Spears and 

Abele (1997), Schram and Hof (1998), and especially 

Monod and Forest (1996). 

Are Crustaceans a Monophyletic Group? 

The question of crustacean monophyly, the place of 

the Crustacea within the Arthropoda, the question 

of arthropod monophyly, and the relationships 

among the many arthropod and crustacean groups 

have been reviewed by several recent workers (see 

especially Boore et al., 1995; Friedrich and Tautz, 

1995; Telford and Thomas, 1995; Raff et al., 1994; 

Fortey et al., 1997; Regier and Shultz, 1997, 

1998b; Wheeler, 1998; Shultz and Regier, 2000; 

Edgecomb et al., 2000). Broader questions concerning 

whether crustaceans and other arthropods belong 

in a phylum or larger clade called the Ecdysozoa 

(see Garey et al., 1996; Aguinaldo et al., 

1997) are reviewed by Schmidt-Rhaesa et al. (1998) 

and Garey (2000). We have not attempted to address 

either of these issues (that is, the relationship 

of crustaceans to other arthropods or the relationships 

within the Ecdysozoa) and instead refer the 

reader to the following publications and the papers 

cited therein. Wheeler et al. (1993) presented a 

combined analysis of morphological and molecular 

data that strongly supported arthropod monophyly, 

and this view was strengthened by Wheeler (1998). 

Lake (1990) suggested arthropod paraphyly, while 

Fryer (1997) presents several arguments in favor of 

arthropod polyphyly. Strausfeld (1998) depicts insects 

and crustaceans (both of which he feels may 

be paraphyletic) as sister groups on the basis of 

neuroanatomical data. Preliminary work on the 

neurogenesis of compound eyes supports common 

ancestry for crustaceans and insects as well (e.g., 

see Harzsch and Walossek, 2001, and references 

cited therein). Friedrich and Tautz (1995) support 

both arthropod monophyly and a crustacean–insect 

sister group arrangement with DNA sequence data, 

as do Boore et al. (1995, 1998), using mitochondrial 

gene rearrangement data, and Wilson et al. 

(2000), comparing the complete mitochondrial ge- 

RATIONALE 

nome of a malacostracan with that of Drosophila. 

Regier and Schultz (1997, 1998a, b) also questioned 

crustacean monophyly (their 1997 title suggests 

crustacean polyphyly) based on EF-1� and 

RNA polymerase II (Pol II); however, their results 

were somewhat ambiguous, as there were no 

strongly supported nodes, and support for a basal 

Malacostraca was not high (J. Regier, pers. comm.). 

Regier and Shultz also suggested (1997, 1998b), as 

had other workers, that branchiopod crustaceans 

may be more closely related to other arthropod 

groups (hexapods and myriapods) than they are to 

malacostracan crustaceans, although this too did 

not have strong node support (what was strongly 

supported was that branchiopods, and indeed all of 

our six classes of crustaceans, grouped with hexapods 

to the exclusion of myriapods, arguing against 

the concept of the ‘‘Atelocerata’’ (hexapods � myriapods); 

see also Popadić et al., 1996, and Shultz 

and Regier, 2000). Another way of stating this is 

that, if crustaceans are not monophyletic, then the 

group that breaks them up is the Hexapoda and 

not myriapods or chelicerates or groups outside Arthropoda. 

The emerging field of developmental biology 

(see references cited in the earlier section on 

developmental genetics and crustacean classification) 

also provides evidence that crustaceans and 

insects are closely linked. Brusca (2000) nicely summarizes 

the history of the controversy and the disparate 

data sets. Two recent volumes address these 

questions by way of collections of edited papers: 

Fortey and Thomas (1997, Arthropod Relationships, 

Chapman and Hall) and Edgecombe (1998, 

Arthropod Fossils and Phylogeny, Columbia University 

Press). 

In the introduction to the latter volume, Edgecombe 

notes that ‘‘the monophyly of Crustacea is 

endorsed in every chapter that investigates the issue’’ 

(see also Edgecomb et al., 2000). Yet there 

remains some doubt. We have found it advantageous, 

at least for the project at hand, to treat the 

group as monophyletic. We also note that there is 

an abundance of fossil, morphological, and molecular 

data that support this view. The ‘‘crown-’’ vs. 

‘‘stem-group’’ approach as detailed by Walossek 

and Müller (1990, 1998) is worth noting in this 

regard; those authors consider the Crustacea monophyletic 

and give several morphological characters 

that uniquely define the group, while at the same 

time they present interesting information on ‘‘stemline 

crustaceans,’’ crustacean-like arthropods that 

are not members of the crown group (their ‘‘Eucrustacea’’) 

but that share at least some features 

with true crustaceans. Other workers have argued, 

some with more data than others, that the Crustacea 

is paraphyletic (e.g., Moura and Christoffersen, 

1996; García-Machado et al., 1999; Wilson et al., 

2000) or polyphyletic (e.g., Averof and Akam, 

1995a, b) or that the question is, at best, unre- 

12 � Contributions in Science, Number 39 Rationale

solved (e.g., Regier and Schultz, 1997, 1998a, b; 

Shultz and Regier, 2000), and we would be remiss 

not to mention these dissenting opinions. Further 

arguments for or against the monophyly of the 

Crustacea (and also Arthropoda) can be found in 

the reviews by Brusca (2000) and Giribet and Ribera 

(2000). 

Our treatment of the Crustacea as a subphylum 

(of the Arthropoda) is therefore somewhat arbitrary. 

Arguments could be (and have been) made 

for recognizing the group as a distinct phylum, and 

some workers refer to the Crustacea as a superclass 

or class. Our choice of subphylum allowed us to 

use classes within the group, which to us was more 

manageable. Treating the Crustacea as a subphylum 

implies monophyly of the Arthropoda. Although 

this issue is not completely settled (see 

above references and especially Fryer, 1997, in Fortey 

and Thomas, 1997), most bodies of evidence of 

which we are aware seem to indicate that the arthropods 

are indeed a phylum (see summaries in 

Raff et al., 1994; Telford and Thomas, 1995; and 

Brusca, 2000) that includes the Crustacea. 

How Many Classes Are There? 

The history of higher level classification of the 

Crustacea is briefly discussed in Holthuis (1993a), 

Spears and Abele (1997), Schram (1986), Schram 

and Hof (1998), and especially Monod and Forest 

(1996). Some of the more notable schemes for crustacean 

classification that have appeared subsequent 

to the Bowman and Abele (1982) classification are 

those of Schram (1986), Starobogatov (1986, with 

English translation by Grygier in 1988), and Brusca 

and Brusca (1990). Other workers have presented 

phylogenies from which the reader can deduce alternative 

classifications, even if no specific classification 

is presented in the paper (e.g., Wilson, 

1992). 

Schram (1986) departed from Bowman and 

Abele’s use of six classes by recognizing four 

groups: Remipedia, Phyllopoda (which included the 

branchiopods, cephalocarids [as Brachypoda], and 

leptostracans), Maxillopoda (including tantulocarids, 

branchiurans, mystacocaridans, ostracodes, copepods, 

facetotectans, rhizocephalans, ascothoracidans, 

acrothoracicans, and thoracicans), and Malacostraca 

(containing both the hoplocarids and the 

eumalacostracans). Schram’s (1986:542–544) classification 

extends to the level of suborder and occasionally 

infraorder. It is noteworthy not only for 

attempting to derive a classification from his cladistic 

analyses but also because of his inclusion of 

a large number of fossil taxa. Unfortunately, 

Schram (1986) also introduced, or employed, some 

taxonomic names that have not been well accepted 

(e.g., ‘‘Euzygida’’ for the stenopodidean shrimps; 

‘‘Eukyphida’’ for the carideans; ‘‘Edriophthalma’’ to 

contain the isopods and amphipods as distinct from 

all other peracarids, etc.). Starobogatov (1986, 

1988) recognized four groups as well, but the com- 

position of his four groups differs appreciably from 

those of Schram and from those of all other previous 

workers. Additionally, Starobogatov employed 

some unusual names for his groupings (such 

as Carcinioides for the malacostracans and Halicynioides 

to accommodate some of the maxillopodan 

groups) that are unlikely to receive wide recognition, 

and his classification appears to be at 

odds with most of the morphological and fossil 

data (e.g., see Schram and Hof, 1998) as well as 

with the molecular data (e.g., Spears and Abele, 

1997). Brusca and Brusca (1990) recognized five 

classes (Remipedia, Branchiopoda, Cephalocarida, 

Maxillopoda, and Malacostraca), and in part because 

this usage is in a major textbook, it has received 

wide acceptance. Bousfield and Conlan 

(1990, Encyclopaedia Britannica), whose classification 

extends only to the ordinal level, followed 

Schram’s lead for some groups of the Crustacea and 

Bowman and Abele (1982) for others. Their classification 

is noteworthy because of their attempt to 

include fossil taxa as well and because of their laudable 

attempt to estimate the number of families in 

each order. Gruner (1993) treats the Crustacea as 

a class, does not recognize the Branchiopoda or 

Maxillopoda, and as a result includes 13 separate 

subclasses. Apart from the somewhat unusual treatment 

by Starobogatov, the number of proposed or 

recognized classes seems to have depended mostly 

upon whether the maxillopods are seen as a natural 

assemblage and, if they are, whether the ostracodes 

are within or outside of the Maxillopoda, and on 

whether and how the Malacostraca should be divided. 

In our classification, the subphylum Crustacea 

includes six major groups, which we are treating as 

classes: Branchiopoda, Remipedia, Cephalocarida, 

Maxillopoda, Ostracoda, and Malacostraca. However, 

this is somewhat misleading in that we are 

also positing the Branchiopoda as the sister taxon 

to all other crustacean groups. Thus, the ‘‘class’’ 

Branchiopoda should be accorded more weight 

than the remaining classes, which together constitute 

the sister group to the branchiopods in our 

arrangement. Our treatment of crustaceans as being 

comprised of six classes is quite conservative and 

follows essentially the Bowman and Abele (1982) 

classification. Perhaps the most salient problem is 

our continued recognition of the Maxillopoda as a 

valid class, when virtually all lines of evidence point 

to its being an artificial assemblage (see discussion 

under Maxillopoda). Thus, Wilson (1992) observed 

that ‘‘the concept of the Maxillopoda is not supported 

in any of the trees’’ and Spears and Abele’s 

(1997) molecular analysis ‘‘fails to provide strong 

support for a monophyletic Maxillopoda.’’ If we 

eliminated the Maxillopoda as a class, as has Gruner 

(1993) (and there are many lines of evidence 

that suggest that this is the correct course), then we 

would treat as distinct classes each of the currently 

recognized ‘‘maxillopodan’’ subclasses (the Thecostraca, 

Tantulocarida, Mystacocarida, and Copepo- 

Contributions in Science, Number 39 Rationale � 13

da). This would have the advantage of further increasing 

our perception of crustacean diversity 

(only because nine classes sounds more diverse than 

six). The number of crustacean classes that should 

be recognized is a very controversial topic, and 

opinion is sharply divided. As Spears and Abele 

(1997) noted, ‘‘surprisingly, there is as yet no consensus 

regarding even the number of constituent 

crustacean classes.’’ 

We do not recognize the taxon ‘‘Entomostraca,’’ 

which has been used historically by several workers 

in slightly different contexts (e.g., McKenzie et al., 

1983; Walossek and Müller, 1998). Walossek and 

Müller (1998:210) and Walossek (1999) recognize 

this group as one of the ‘‘two major lineages’’ of 

Crustacea (the other being the Malacostraca). Contained 

in their Entomostraca are the cephalocarids 

(depicted as the sister taxon to the Maxillopoda 

and Branchiopoda) and two extinct groups (Orstenocarida 

and Skaracarida). 

Which Is the Most Primitive Class? 

We are treating the class Branchiopoda as the most 

primitive of the extant groups of Crustacea. We arrived 

at this decision mostly because of the following 

three lines of evidence. First, the group as a 

whole is ancient and extends back into the Upper 

Cambrian and probably further (see Fryer, 1999, 

and especially Walossek, 1993). A beautifully preserved 

fossil from the Upper Cambrian of Sweden 

(Rehbachiella) appears to be a branchiopod and is 

similar in many ways to living anostracans (Walossek, 

1993; although note that Olesen (1999a) 

questions the anostracan affinities of Rehbachiella, 

while both Wills (1997) and Schram and Hof 

(1998) obtained nonbranchiopod positions for 

Rehbachiella on their cladograms). There are no 

known fossils of any cephalocarids, and the only 

fossils thought to be remipedian are from the Carboniferous 

(Mississippian and Pennsylvanian) Period 

(Schram and Hof, 1998). In fairness, we 

should state also that (1) cephalocarids, because of 

their habitat, size, and fragility, would seem unlikely 

candidates for fossilization (and yet, such could 

also be said about the minute animals in the Orsten 

fauna) and (2) there are other crustacean groups 

known from the Upper Cambrian, such that appearance 

of branchiopods in the Upper Cambrian 

is not in itself sufficient to argue for their being the 

most primitive of the extant classes. Second, there 

are developmental studies that show clear and unambiguous 

anamorphic development in at least 

some branchiopods, which is exhibited by no other 

living crustacean group (e.g., see Fryer, 1983). On 

the other hand, cephalocarids exhibit only slightly 

metamorphic development, and as of this writing, 

we still know nothing about remipede development. 

Third, some studies based on molecular sequence 

data seem to indicate that branchiopods are 

not only monophyletic but are also distinct from all 

other crustacean assemblages (e.g., Spears and 

Abele, 1997, 2000; Regier and Schultz, 1997, 

1998a, b; Shultz and Regier, 2000). As noted earlier, 

Regier and Schultz (1997) suggested that branchiopods 

may be closer to other groups of arthropods 

than to malacostracan crustaceans, although 

there was no strong support for this arrangement 

and they concluded that the EF-1� data are ambiguous 

on this question. These authors later (1998b) 

depict remipedes closer to the crustacean stem, but 

again in this analysis, node support was not strong, 

and thus the authors remain suitably cautious as to 

interpretation of these data (J. Regier, pers. comm.). 

Spears and Abele (1997) conclude that ‘‘we cannot 

identify which crustacean lineage is most basal; 

branchiopods, pentastomes, branchiurans, and ostracodes 

[but note the absence of remipedes or cephalocarids] 

all diverged from the main crustacean 

lineage in relatively rapid succession.’’ Although arguments 

on this point will surely continue for many 

years to come, we have elected to follow the 18S 

rDNA-based findings of Spears and Abele (1997), 

supported to some degree (in our estimation) by the 

EF-1� findings of Regier and Schultz (1997, 1998b; 

see also Shultz and Regier, 2000). Thus, we treat 

branchiopods first in our classification, thereby implying 

that we are in agreement with branchiopods 

being the most basal of the extant crustacean 

groups. This treatment also receives some support 

from Itô’s (1989) suggestion of a remipede � cephalocarid 

� copepod clade, an arrangement that 

was also suggested by Spears and Abele (1997) 

based on 18S rDNA data (see especially their fig. 

14.7 and accompanying discussion). We have not, 

however, created the additional taxonomic categories 

that would be required to group branchiopods 

as the sister group to all other crustaceans. In other 

words, our classification is far from being a strictly 

cladistically based arrangement. Branchiopods are 

thus accorded class status, as are the other five major 

crustacean groupings, in this classification. Additionally, 

if we are positing the branchiopods as 

the sister group to the other crustaceans, then we 

should list specific synapomorphies unique to the 

clade. Most of the morphological characters seeming 

to cast branchiopods in a primitive light (e.g., 

foliaceous limbs, anamorphic development) are indeed 

primitive features, but they may have been retained 

in this group and lost or modified in others. 

Noting simply that their morphology is ‘‘primitive’’ 

sheds no real light on phylogeny, and other groups 

of crustaceans exhibit other ‘‘primitive’’ characters. 

Possible candidates for branchiopod synapomorphies 

might include the ‘‘specialization of postnaupliar 

feeding apparatus to true filter feeding’’ (from 

Walossek, 1993:71), aspects of sperm morphology 

(Wingstrand, 1978), and the 18S rDNA sequences, 

which Spears and Abele (2000) used to conclude 

that ‘‘(1) branchiopods are monophyletic; (2) they 

are considerably divergent from other crustaceans 

(e.g., the Malacostraca), and (3) they are divided 

into two main lineages’’ (Anostraca and all others). 

The issue of which extant class is closest to the 


ancestral crustacean is of course not completely settled, 

and there are published arguments for presenting 

either the Cephalocarida or the Remipedia 

as the most primitive group of living crustaceans. 

There have also been, from time to time, hypotheses 

presented where other groups of crustaceans 

have occupied a basal position (e.g., McKenzie, 

1991, postulated a bradoriid ostracode origin for 

all other crustaceans). 

In favor of depicting remipedes as the most primitive 

class are the works of Schram (1986), Brusca 

and Brusca (1990), Briggs et al. (1993a), Schram 

and Hof (1998), Wills (1997), and Wills et al. 

(1998), all based on cladistic analyses of morphological 

characters from extant and extinct forms. 

Also supporting this view is the phylogeny presented 

by Jamieson (1991a) based on sperm ultrastructure, 

in which the Remipedia is the most basal of 

the crustacean groups. (It should be noted, however, 

that Jamieson’s study is not purely independent 

of other phylogenies in that his figure is actually 

an overlay of the various sperm types on top 

of the classification offered by Schram in 1986.) 

Thus, there are workers at several independent laboratories 

whose studies have indicated that remipedes 

occupy the most basal position among the 

crustaceans, and several textbooks have followed 

this arrangement as well (e.g., Hickman et al., 

1996: 401, figs. 20–30; Brusca and Brusca, 1990). 

Molecular evidence concerning where remipedes 

belong has been maddeningly difficult to obtain. 

Regier and Schultz (1998b) could not say with certainty 

(using EF-1�), and Spears and Abele (1997) 

were equally unsure (using 18S rDNA). Emerson 

and Schram (1990, 1997) have also suggested that 

crustacean biramous limbs arose from fusion of adjacent 

uniramous limbs, and this has a bearing on 

the placement of remipedes relative to other crustacean 

groups as well (discussed further in Schram 

and Hof, 1998, but see Spears and Abele, 1997). It 

should also be pointed out that at least one publication 

(Moura and Christoffersen, 1996) suggests 

that the Remipedia are a derived assemblage that 

may be the sister group to the Tracheata (terrestrial 

mandibulates). 

In support of cephalocarids occupying the most 

basal position among extant crustaceans are some 

surely primitive external morphological features. 

These features include the flattened and ‘‘Orstenlike’’ 

limbs, the lack of differentiation of the second 

maxilla (also shared with some of the Orsten crustaceans), 

and relatively anamorphic development. 

Hessler (1992) reviewed early considerations of the 

placement of the cephalocarids with respect to other 

crustaceans. He concluded, based on the morphology 

of some of the Upper Cambrian ‘‘Orsten’’ 

fauna of Sweden and in comparison with remipedes 

and other crustaceans, that the argument for placing 

cephalocarids at the base of the crustacean lineage 

is still strong (see also Walossek, 1993; Moura 

and Christoffersen, 1996). In Hessler’s words, 

‘‘among living crustaceans, cephalocarids still best 

personify what the ur-crustacean must have looked 

like.’’ Hessler (1992) also made the point, with 

which we agree, that remipedes are quite specialized, 

and he found it ‘‘impossible to accept the 

claim that the Remipedia better approximates the 

ur-crustacean.’’ However, cephalocarids face problems 

as primitive crustaceans as well. Schram and 

Hof (1998) point out some cephalocarid features 

they consider highly derived, and molecular studies 

(e.g., Spears and Abele, 1997; Regier and Schultz, 

1998b) and spermatological data (especially lack of 

a flagellum; see Jamieson, 1991a) do not place cephalocarids 

basal to other crustacean taxa (although 

in fairness, the EF-1� data of Regier and 

Shultz do not decisively place cephalocarids elsewhere, 

either). We have not followed the suggestion 

of Hessler (1992) to revive the taxon Thoracopoda 

to include the cephalocarids, branchiopods, and 

malacostracans (based on their shared possession of 

an epipod on the trunk limbs). 

What Are the Relationships Among the Classes? 

This question is closely related to the issues raised 

above. In fact, most of the competing phylogenetic 

hypotheses for class-level relationships have already 

been alluded to in earlier sections (e.g., in the sections 

‘‘Cladistics and Classification of the Crustacea’’ 

and ‘‘Molecular Systematics and Classification 

of the Crustacea,’’ and under the above three questions 

on crustacean monophyly, number of classes, 

and most primitive class). Rather than attempt a 

discussion of the many competing hypotheses for 

the relationships within and among the various 

classes, we have opted to treat each group individually 

below. We also refer the reader to the reviews 

by Wills et al. (1998) and Schram and Hof (1998), 

both in Edgecombe (editor, 1998, Arthropod Fossils 

and Phylogeny), and to the review of 18S rDNA 

studies by Spears and Abele (1997). 

Concerning authorship of the name Crustacea, 

although most workers credit Pennant (1777), Lipke 

Holthuis, in a detailed and well-researched footnote 

to his FAO volume on marine lobsters (Holthuis, 

1991), noted that the first usage was actually 

that of Brünnich in 1772. We have followed Holthuis’ 

(1991) suggestion and have credited Brünnich 

(1772) with authorship of this taxon. 

CLASS BRANCHIOPODA 

Virtually all evidence points to the fact that the 

branchiopods are a strongly supported monophyletic 

group, despite the staggering diversity of extant 

forms (e.g., see Martin, 1992). Lines of evidence 

indicating branchiopod monophyly include 

sperm morphology (Wingstrand, 1978), larval 

characters (e.g., Sanders, 1963), feeding apparatus 

(Walossek, 1993), adult characters (e.g., Negrea et 

al., 1999), and 18S rDNA sequence data (Spears 

and Abele, 1997, 1998, 1999a, b, 2000). However, 

the group’s tremendous morphological diversity 

and age (see Fryer, 1987a–c, 1999; Martin, 1992; 


Walossek, 1993; Negrea et al., 1999) makes it difficult 

to find characters shared by all extant members, 

and perhaps for this reason some analyses 

have hinted at para- or polyphyly (e.g., see Wilson, 

1992). Gruner (1993) does not recognize the Branchiopoda, 

instead treating the extinct Lipostraca 

and the extant Anostraca and Phyllopoda (Notostraca 

� Diplostraca) as separate subclasses within 

the class Crustacea. The fact that there appears to 

be solid support from molecular data for branchiopod 

monophyly (e.g., Spears and Abele, 1997, 

1998, 1999b, 2000) is nevertheless reassuring. 

There is also a consensus that, within the Branchiopoda, 

the Anostraca diverged early, are very 

primitive (despite a large number of apomorphic 

features in the various families), and should be depicted 

as separate from the remaining branchiopod 

groups. Beyond that, however, there is little agreement 

concerning the relationships among the constituent 

branchiopod taxa. 

Because the Anostraca are clearly a separate lineage 

from the remaining branchiopods and are an 

ancient and slowly evolving group (e.g., see Fryer, 

1992, 1999), we have elevated the group to the level 

of subclass, to be treated as the sister group of 

the other branchiopods (as was advocated also by 

Walossek, 1993, and Negrea et al., 1999). However, 

this move necessitates creating a name for the 

subclass or choosing an available name from the 

literature to contain the Anostraca (and which 

would eventually, we assume, contain also the fossil 

branchiopod order Lipostraca and possibly also the 

Cambrian Rehbachiella; see Walossek, 1993; Walossek 

and Müller, 1998). Tasch’s (1969) proposal 

to use the name Sarsostraca (to contain anostracans 

and lipostracans) is not very appealing, in part because 

Tasch originally included in his Sarsostraca a 

noncrustacean (obviously also a nonbranchiopod), 

and one of his anostracans was in fact an insect 

larva (G. Fryer, pers. comm.). Nevertheless, the 

name Sarsostraca appears to be a valid preexisting 

name by ICZN standards and would have seniority 

over any newly proposed name here, so reluctantly 

we accommodate the order Anostraca within the 

subclass Sarsostraca, as did Bowman and Abele 

(1982) and, more recently, Negrea et al. (1999). 

Finding a name suitable to contain the other 

(non-Anostraca) groups was more difficult. First of 

all, the tremendous morphological differences 

among the groups traditionally thought of as cladocerans, 

conchostracans, and notostracans has led 

several workers, most notable among them Geoffrey 

Fryer (e.g., see Fryer, 1987a, c, 1995, 1999a, 

b), to suggest that there is no reason to try to force 

such disparate groups into artificial groupings as 

‘‘cladocerans’’ and ‘‘conchostracans.’’ Fryer’s wellwritten 

articles argue convincingly for the separation 

of these ancient and diverse taxa (most of 

which he would elevate to ordinal level), and indeed 

his suggested classification (Fryer, 1987a, c) 

has been followed by several workers, such as Martin 

(1992), Alonso (1996), Amoros (1996), Frey 

(1995), Brtek and Thiéry (1995), Thiéry (1996), 

Brtek (1997), and others. However, simply recognizing 

how different these groups are from one another 

and elevating the former conchostracan or 

cladoceran taxa to higher taxonomic categories 

while doing away with the categories that once included 

them does not, in our opinion, shed light on 

their relationships. The question still remains as to 

whether these orders are more closely related to one 

another than any is to some other crustacean assemblage. 

The morphological and molecular evidence 

seems to indicate (1) that branchiopods are 

monophyletic and (2) that some of these taxa (not 

all are well represented by molecular or even morphological 

data) are indeed related more closely to 

one another than to any other crustacean group. 

The alternative is to suggest that, for example, the 

Anomopoda are more closely related to anostracans 

or to some nonbranchiopod crustacean. We 

think this is very unlikely. Thus, the value of Fryer’s 

arguments is in the recognition of the tremendous 

age and morphological differences that exist (and 

have existed for a long time) among these disparate 

taxa, a point that is well taken. Despite these arguments, 

and because we still must postulate relationships, 

we are forced to group these taxa together. 

Toward this end, several workers have suggested 

that we use the name Phyllopoda for the taxon encompassing 

the Notostraca and the bivalved branchiopods 

(see comments below about the nonmonophyly 

of the ‘‘diplostracans’’), and indeed the 

name Phyllopoda has been used often for that assemblage 

(e.g., Walossek, 1993, and later). Unfortunately, 

the name Phyllopoda has also been used 

to denote groupings that include the Anostraca or 

that include the Ostracoda or that include the Leptostraca 

and Cephalocarida and in several other 

contexts as well. In fact, the term Phyllopoda has 

been used so often in crustacean systematics, and 

with such different meanings, that Martin and 

Christiansen (1995a) argued for avoiding it completely 

to avoid further confusion. Not surprisingly, 

we agree with Martin and Christiansen (1995a) 

and would prefer to employ another available name 

for this lineage. Does one exist? Tasch (1969) employed 

the names Calmanostraca (for the notostracans) 

and Diplostraca (for the conchostracans and 

cladocerans) as subclasses, but the two groups were 

treated equally (i.e., Tasch did not depict them as 

being more closely related to each other than either 

would be to the anostracans). Because the name 

Diplostraca obviously refers to the bivalved carapace 

seen in some groups, we could have opted to 

use the name Calmanostraca suggested by Tasch 

(1969) but expanding its definition to include both 

notostracans and the bivalved groups, which seems 

to be advocated by the classification proposed by 

Spears and Abele (2000). However, the name Calmanostraca 

should probably be reserved for containing 

the extinct Kazacharthra and the extant 

Notostraca (as it was first intended) when fossil 

taxa are eventually added to the ‘‘updated’’ classi- 


fication (see also Negrea et al., 1999). Therefore, 

with trepidation and against our own recommendations 

(Martin and Christiansen, 1995a), we have 

resurrected the name Phyllopoda, using it this time 

to include the extant Notostraca and the bivalved 

branchiopod groups (i.e., all branchiopods except 

the Anostraca). We have credited the taxon name 

to Preuss (1951), who was, to our knowledge, the 

first person to use the name Phyllopoda in the sense 

that we are using it (to contain all branchiopods 

other than the anostracans). This decision will surely 

prompt arguments from many current students 

of the Branchiopoda (see especially Fryer, 1987c, 

1995, 1999b). 

There have been many significant findings in extant 

and extinct branchiopods that have altered our 

view of branchiopod relationships since the Bowman 

and Abele (1982) classification. Morphological 

treatments have included Fryer (1983, 1985, 

1987a–c, 1995, 1996a, b, 1999), Martin (1992), 

Martin and Cash-Clark (1995), Walossek (1993, 

1995), Olesen et al. (1997), Olesen (1996, 1998, 

1999), Thiéry (1996), Amoros (1996), and Negrea 

et al. (1999), to mention only a few of the recent 

papers. There have also been several attempts to 

deduce branchiopod relationships using molecular 

data, including Hanner and Fugate (1997) and 

Spears and Abele (1997, 1998, 1999b, 2000). In 

the current classification, we have attempted to reconcile 

some of the recent morphological and molecular 

findings, but earlier classifications should 

not be discarded as being out of date or invalid. 

Indeed, many of the most detailed accounts of 

branchiopods remain the older, classical treatments, 

and to ignore these is a grave mistake. Thiéry 

(1996, based in large part on Martin, 1992) reviewed 

the biology of the noncladoceran groups 

(including Cyclestheria among the conchostracans), 

and Amoros (1996) reviewed the four ‘‘former cladoceran’’ 

orders Ctenopoda, Anomopoda, Onychopoda, 

and Haplopoda. 

SUBCLASS SARSOSTRACA, ORDER 

ANOSTRACA 

Within the Anostraca, Brtek (1995) elevated the 

former chirocephalid subfamily Artemiopsinae to 

family level and thus recognized the Artemiopsidae. 

Earlier, Brtek (1964) established the family Linderiellidae. 

However, Denton Belk (pers. comm.) believed 

these moves are unwarranted. Concerning 

the Artemiopsidae, Belk stated, ‘‘placing this single 

genus in a separate family obscures the many features 

it shares with other genera in the Chirocephalidae, 

and is thus a hindrance to having a meaningful 

taxonomic classification of the Anostraca.’’ 

Concerning the Linderiellidae, he noted that ‘‘these 

genera have antennal appendages and some penal 

features that suggest they are related to other genera 

of the Chirocephalidae; separate familial status 

obscures these seemingly significant similarities.’’ In 

light of Belk’s expertise with anostracans, we have 

followed his suggestion and have not recognized 

these two families, although they are recognized in 

the latest key to families and genera (Brtek and 

Mura, 2000). Our classification of the Anostraca 

therefore follows Belk (1996), with the exception 

of the Linderiellidae (which was included by Belk, 

1996, but is not included here). A recent molecular 

analysis (Remigio and Hebert, 2000) of the relationships 

among extant anostracan families suggested 

two clades, one containing Artemiidae and 

Branchipodidae and the other containing the other 

five families. 

SUBCLASS PHYLLOPODA 

By placing anostracans in a subclass separate from 

all other branchiopods, we are assuming also that 

the other branchiopods form a monophyletic 

grouping. In other words, we believe that the notostracans, 

conchostracans, and cladocerans are 

more closely related to one another than any of 

those groups is to the anostracans. There are some 

morphological features (e.g., Negrea et al., 1999) 

and molecular data (e.g., Spears and Abele 1997, 

1999b, 2000) that suggest this might be true. This 

arrangement has been proposed by many other 

workers as well (some of whom, such as Walossek, 

1993, 1995; Walossek and Müller, 1998, have also 

employed the name Phyllopoda in the same sense 

that we are using it). 

ORDER NOTOSTRACA 

It may be necessary, once fossil taxa are included 

in this classification, to someday resurrect Tasch’s 

(1969) name Calmanostraca to accommodate the 

extant notostracans and the extinct and obviously 

closely related Kazacharthra. The sole family of extant 

Notostraca, Triopsidae, is credited to Keilhack 

(‘‘Kielhack’’ was a misspelling in Bowman and 

Abele, 1982), and that date has been changed from 

1910 to 1909 (L. Holthuis, pers. comm.). Although 

the original spelling was Triopidae, as listed in 

Bowman and Abele (1982), the spelling Triopsidae 

(based on the genus Triops) was entered in the Official 

List of Family-Group Names in Zoology by 

the ICZN, Opinion 502 (M. Grygier, pers. comm.). 

ORDER DIPLOSTRACA 

As noted above, the Phyllopoda as used here includes 

the orders Notostraca and Diplostraca (a 

name that predates Onychura used by some authors, 

such as Walossek, 1993, and Negrea et al., 

1999). Whether these are indeed sister taxa is unclear; 

there is some morphological and molecular 

evidence to suggest that this might not be the case. 

Recognition of the taxon Diplostraca indicates our 

feeling that the former conchostracan and cladoceran 

groups are indeed related. There appears to be 

some morphological (e.g., see Walossek, 1993; Olesen, 

1998; Negrea et al., 1999) and molecular 

(Spears and Abele, 2000) evidence supporting this 


elationship, although the view is certainly not universally 

shared (e.g., see the exchange between Olesen, 

1998, 2000, and Fryer, 1999, 2001), and there 

is a large body of evidence suggesting that Diplostraca 

is nonmonophyletic. Additionally, there is 

considerable doubt concerning the monophyly of 

some of the groups we have included within it, such 

as the Cladocera. Fryer (1987a, 1995, 1999a, b) 

discusses the great morphological differences 

among the four groups traditionally placed in the 

‘‘so-called Cladocera’’ and highlights the trenchant 

differences among these taxa and the difficulty in 

reconciling these forms within one taxonomic category. 

We should also point out that the ‘‘secondary 

shield’’ mentioned as unifying these taxa (e.g., by 

Walossek, 1993; Olesen et al., 1997; Olesen, 1998) 

is, according to Fryer (1996b, 1999b), simply nonexistant, 

a misunderstanding of the nature of the 

crustacean carapace. Other characters that supposedly 

unite the ‘‘diplostracan’’ groups are similarly 

called into question by Fryer in a series of papers 

(1987a–c, 1995, 1996a, b, 1999b). In particular, 

after considerable work in attempting to reconstruct 

a primitive anomopod from which extant anomopods 

could have been derived and by so doing 

highlighting the great difficulties of any such exercise, 

Fryer (1995) argued against attempting to 

force such disparate taxa as Leptodora, Bythotrephes, 

and the superficially similar ctenopods into a 

taxon with the Anomopoda, stating (pers. comm.) 

that ‘‘when those who make these proposals can 

support them by evolutionary series that involve 

animals that would work, I’ll pay more attention 

to them.’’ 

Within the Diplostraca, we have removed the 

‘‘Conchostraca’’ (following to some extent the suggestions 

of Fryer, 1987c, and Olesen, 1998) in recognition 

of (1) the distinct nature of the Laevicaudata 

(Lynceidae), (2) the stark differences that separate 

Cyclestheria hislopi (sole member of the Cyclestheriidae) 

from all other conchostracans, and 

(3) Cyclestheria’s possible affinities to the cladocerans 

on morphological and molecular grounds (see 

Martin and Cash-Clark, 1995; Olesen et al., 1997; 

Olesen, 1998; Spears and Abele, 1998, 2000). The 

fact that Cyclestheria differs significantly from other 

spinicaudate conchostracans, and probably to 

the extent that it should not be placed among them, 

has also been highlighted (Martin and Cash-Clark, 

1995; Olesen et al., 1997; Olesen, 1999; Negrea et 

al., 1999). Thus, our resulting classification within 

the Diplostraca differs slightly from, and is in some 

ways a compromise between, the classification suggested 

by Olesen (1998) based on morphological 

characters and that suggested by Spears and Abele 

(2000) based on molecular data and is easily reconciled 

with the phylogeny proposed by Negrea et 

al. (1999). Our arrangement does not agree with 

the somewhat preliminary findings of Hanner and 

Fugate (1997) based on a relatively small segment 

of the genome. 

Removal of Cyclestheria from the Spinicaudata 

and placing it on an equal footing with the remaining 

Spinicaudata and with the Cladocera necessitated 

the creation of a separate suborder, the Cyclestherida, 

which we are crediting to Sars (1899) 

in keeping with ICZN article 50.3.1. Negrea et al. 

(1999) used the same spelling to refer to an order 

(Cyclestherida) within their superorder Conchostraca, 

thus indicating a closer affinity of Cyclestheria 

to the conchostracans rather than the cladocerans. 

We have not taken the bolder step of actually 

including the Cyclestheriidae among the Cladocera, 

although there is apparently evidence for 

this as well. Spears and Abele (1999a, b, 2000) note 

that, not only do 18S rDNA sequence data support 

the close relationships of Cyclestheria and the cladocerans, 

the two groups also share certain hypervariable 

regions of the gene that are not found in 

other branchiopods, and these are potential synapomorphies. 

Ax (1999) first suggested the term 

‘‘Cladoceromorpha’’ for the clade containing Cyclestheria 

plus Cladocera. Papers by Crease and 

Taylor (1998) and Taylor et al. (1999) appear to 

offer additional molecular support, and the phylogeny 

suggested by Negrea et al. (1999:196) supports 

such a clade as well, although their resulting classification 

of the Branchiopoda into five superorders 

does not. 

Sassaman (1995) presented fascinating insights 

into possible phylogenetic models for the conchostracan 

families based on the evolution of unisexuality 

in the group; he views lynceids as the sister 

group to all other families, while noting at the same 

time the unusual nature of the cyclestheriids, which 

he posits as the sister group to the remaining ‘‘spinicaudatan’’ 

families. Thus, in many ways, Sassaman’s 

(1995) phylogeny is consistent with our classification. 

Within the former ‘‘conchostracan’’ groups, the 

spelling of the Lynceidae has been corrected (from 

Lyncaeidae, a typographical error in Bowman and 

Abele, 1982), and authorship for the family is now 

credited to Baird, 1845 (L. Holthuis, pers. comm.). 

Mark Grygier points out (pers. comm.) that ICZN 

Opinion 532 attributes the family name to Sayce, 

1902; however, there are clearly earlier uses of the 

family name Lynceidae (e.g., see review by Martin 

and Belk, 1988), and we are crediting the family 

name to Baird as noted above. 

Although the genera Imnadia and Metalimnadia 

at times have been suggested to represent distinct 

families (the Imnadiidae Botnariuc and Orghidan 

and the Metalimnadiidae Straskraba; see Marincek 

and Petrov, 1991; Roessler, 1991, 1995a, b; Orr 

and Briggs, 1999:8), most workers (e.g., Martin, 

1992; Sassaman, 1995) consider them members of 

the family Limnadiidae, as do we. Roessler’s (1991) 

erection of the family Paraimnadiidae was based on 

a species he described as Paraimnadia guayanensis, 

a junior synonym of Metalimnadia serratura (see 

Orr and Briggs, 1999). We also include among the 

limnadiids the genus Limnadopsis and agree with 


Bowman and Abele in not recognizing Tasch’s 

(1969) family Limnadopsidae. 

The superfamilies Cyzicoidea (which contained 

only Cyzicidae) and Limnadioidea have been removed, 

as there is no longer any need for them in 

light of the above reassignments. Indeed, the families 

Cyzicidae and Leptestheriidae are probably 

more closely related to each other than either is to 

the Limnadiidae (Martin, 1992; Sassaman, 1995). 

Within the Cladocera, the spelling of the Holopediidae 

has been corrected (from Holopedidae in 

Bowman and Abele, 1982) in light of the spelling 

of the type genus Holopedium (M. Grygier, pers. 

comm.). The correct spelling of Macrotrichidae 

(rather than Macrothricidae) was also pointed out 

to us by M. Grygier (pers. comm), referring us to 

Appendix D of the ICZN, third edition, example 

24, page 223 (ICZN, 1985a), for examples of family 

names formed from genus names ending in - 

thrix. However, the fourth edition of the Code 

(ICZN, 1999) now allows such misspellings to 

stand if they are in ‘‘prevailing use,’’ which the family 

name Macrothricidae certainly is. Thus, we retain 

the spelling Macrothricidae. (This same logic 

(i.e., retention of a misspelling because of prevailing 

use) applies also to the family Rhizothricidae in the 

harpacticoid copepods.) 

Within the Anomopoda, we have removed the 

family Moinidae, following the suggestion of G. 

Fryer (1995, and pers. comm.). Comparisons of the 

trunk limbs of species of Moina and Daphnia indicate 

great similarity between these groups; certainly 

they are much more similar than are many 

macrothricid and chydorid genera to each other. If 

a separate family were recognized for Moina and 

Moinodaphnia, then we would have to erect a series 

of families for various chydorids and macrothricids, 

which we see as only adding to the confusion. 

Thus, the Moinidae is not recognized here. 

For the same reason, we have decided not to recognize 

the family Ilyocryptidae as treated by Smirnov 

(1992) based on the genus Ilyocryptus (see also 

Young, 1998:23). However, it is possible that the 

correct course of action would be to acknowledge 

anomopodan diversity by recognizing both the 

Moinidae and Ilyocryptidae as valid families and 

establishing the additional families for other genera 

as needed. 

The four main cladoceran groupings have been 

treated as infraorders. Although we are in full 

agreement with Fryer’s (1987a–c, 1995) assessment 

of the distinct nature of, and tremendous differences 

among, these taxa (Fryer argued for removal of 

the terms ‘‘cladocera’’ and ‘‘conchostraca’’ as formal 

taxonomic entities), we nevertheless felt that 

the four groups are more closely related to one another 

than any one of them is to any other crustacean 

assemblage, the same conclusion reached by 

Richter et al. (2001) and several earlier workers. 

This may prove to be a mistake. Certainly, treatment 

of the cladocerans as a single order containing 

four infraorders and a handful of families has the 

unfortunate appearance of minimizing the staggering 

morphological and ecological diversity of this 

group, and we very much regret that. Schwenk et 

al. (1998) provided a preliminary estimate of the 

relationships of the Ctenopoda, Haplopoda, Onychopoda, 

and Anomopoda based on 16S rDNA sequence 

data. See Fryer (1995) for suggested relationships 

among the families of the Anomopoda 

and Richter et al. (2001) for 12S rDNA-based relationships 

among onychopods and between the 

‘‘gymnomerans’’ (� onychopods � Leptodora) and 

other cladoceran groups. 

The taxon ‘‘Eucladocera’’ has been removed, as 

we saw no evidence for grouping together all other 

cladocerans as the sister taxon to the monotypic 

Haplopoda (Leptodora), as proposed by several 

workers (most recently by Negrea et al., 1999). Our 

classification is more in keeping with the study by 

Richter et al. (2001), who supported the monophyly 

of the Onychopoda � Haplopoda (the former 

Gymnomera) and argued for cladoceran monophyly. 

The superfamilies Sidoidea, Daphnioidea, and 

Polyphemoidea have also been removed. 

CLASS REMIPEDIA 

It is a little discouraging that we still know so little 

about the phylogenetic relationships of this fascinating 

group. The initial establishment of a separate 

class (Yager, 1981) met with criticism early on, 

and similarities between the limbs of remipedes and 

those of certain maxillopods have been pointed out 

(Itô, 1989). Felgenhauer et al. (1992) hinted at molecular 

data that suggested maxillopodan affinities 

as well, although, to our knowledge, these data 

have not been published. Spears and Abele (1997) 

also suggested possible maxillopodan affinities. In 

an early draft of this classification, we had the remipede 

families included among the Maxillopoda, 

but this was criticized, and rightly so, by several 

persons who pointed out that some of the similarities 

between Remipedia and Maxillopoda are symplesiomorphies 

(although others, such as the loss of 

the maxillary endopod, defined precoxa of the 

maxillule, and three-segmented endopod of the 

trunk limbs, may be synapomorphies) and are insufficient 

to warrant the inclusion of the former 

among the latter. More detailed morphological 

studies (e.g., Schram et al., 1986; Itô and Schram, 

1988; Schram and Lewis, 1989; Yager, 1989a, b, 

1991; Yager and Schram, 1986; Emerson and 

Schram, 1991; Felgenhauer et al., 1992) seem to 

confirm the unique nature of the group. Their status 

as a distinct class is therefore maintained in this 

classification. See also our introductory comments 

concerning which class of extant Crustacea appears 

most plesiomorphic. 

As noted above in the general discussion of the 

primitive groups of Crustacea, several workers 

(e.g., see Schram, 1986; Brusca and Brusca, 1990; 

Briggs et al., 1993a; Schram and Hof, 1998; Wills, 

1997; Wills et al., 1998) have suggested that re- 


mipedes occupy the most basal position among the 

extant crustaceans. These arguments are perhaps 

best summarized in Schram and Hof (1998) and in 

Wills (1997), where remipedes come out at the base 

of all other Crustacea groups following cladistic 

analyses of large datasets. Moura and Christoffersen 

(1996) take an opposing stance, suggesting that 

remipedes are an apical group of crustaceans that 

are possibly the sister group to terrestrial mandibulates. 

To us, the evidence (morphological, molecular, 

and developmental) for branchiopods being 

basal appears stronger (see earlier comments on 

primitive crustaceans). Emerson and Schram (1990; 

see also Emerson and Schram, 1991) have suggested 

that crustacean biramous limbs may have arisen 

from fusion of adjacent uniramous limbs, and this 

has a bearing on the placement of remipedes relative 

to other crustacean groups (discussed further 

in Schram and Hof, 1998). Spears and Abele 

(1997) also discussed possible affinities between remipedes 

and cephalocarids, some of which may be 

artifactual as a result of long branch attractions. 

Within the Remipedia, the order Nectiopoda was 

erected by Schram (1986) to separate extant remipede 

families from some fossils that appear remipedian 

(and that are treated as the fossil order Enantiopoda). 

One additional family, the Godzilliidae, 

was added by Schram et al. (1986). Yager and 

Humphreys (1996) reported the first species from 

Australia and the Indian Ocean and presented a key 

to the world species known at that time. Cals 

(1996) reviewed the biology of the group and presented 

a table comparing the characteristics of the 

two currently accepted families, Speleonectidae and 

Godzilliidae; more recently, Yager and Carpenter 

(1999) and Carpenter (1999) have added to what 

is known of the natural history of speleonectids. 

CLASS CEPHALOCARIDA 

Our classification differs from that of Bowman and 

Abele (1982) only in recognizing a single family, 

Hutchinsoniellidae, rather than two families. The 

family Lightiellidae proposed by Jones (1961) is 

thought to differ only slightly and insignificantly 

from the characters established for the former family 

(R. Hessler, pers. comm.). Our placement of the 

cephalocarids here, between the remipedes and 

maxillopods, to some degree reflects the summary 

finding of Spears and Abele (1997) that remipedes 

and cephalocarids may constitute a clade that is the 

sister group to one of the maxillopodan groups (the 

Copepoda) (e.g., Spears and Abele, 1997, figs. 14.4, 

14.7, and accompanying text), although Spears and 

Abele (1997) also note that this arrangement is not 

well supported by their bootstrap analysis. The 

placement of cephalocarids and remipedes together, 

and adjacent to the maxillopods, in some ways also 

supports Itô’s (1989) morphology-based suggestion 

of a remipede � cephalocarid � copepod clade. 

Hessler and Elofsson (1996) recently reviewed 

what is known of cephalocarid biology and phylogeny. 

CLASS MAXILLOPODA 

The Maxillopoda continues to be a terribly controversial 

assemblage concerning both the number of 

constituent groups and the monophyly of the entire 

taxon. We were tempted to abandon, once and for 

all, the concept of a monophyletic Maxillopoda, as 

there seems very little in the way of morphological 

or molecular evidence uniting the disparate groups 

(Wilson, 1992; Spears and Abele, 1997; Shultz and 

Regier, 2000). Ostracodes in particular have been 

placed sometimes within the Maxillopoda (e.g., see 

Boxshall and Huys, 1989a) and sometimes in their 

own class, and the issue remains unresolved despite 

much debate (e.g., see Boxshall et al., editors, Acta 

Zoologica, vol. 73(5), 1992). It is certainly no secret 

that the characters used in defining the group 

do not hold for many of the taxa traditionally 

thought of as being ‘‘maxillopodan.’’ Abandoning 

the Maxillopoda seems to have been implied in 

tome VII fascicule II of the Traité de Zoologie 

(1996), as only the constituent groups are treated 

with no mention of maxillopod affinities or relationships 

(e.g., see Grygier 1996a, b), and Gruner 

(1993) similarly did not recognize the Maxillopoda. 

Boxshall (1983) and others have argued against 

recognition of the Maxillopoda on morphological 

grounds, although Boxshall has also continued to 

employ it from time to time (e.g., in Huys et al., 

1994). Yet other workers (e.g., see Newman, 1983; 

Grygier, 1983a; Walossek, 1993; Wills, 1997; Walossek 

and Müller, 1998) have argued, some quite 

forcefully, that there is merit to recognition of the 

Maxillopoda as a natural (monophyletic) assemblage, 

despite the fact that there seem to be exceptions 

to every synapomorphy proposed. In fairness, 

so many maxillopodan taxa are so small and/or 

modified as parasites that it should come as no surprise 

to find exceptions to groundplans. Removal 

of the Maxillopoda as a class would raise the number 

of crustacean classes from six to nine once the 

maxillopodan subclasses were elevated (each to the 

level of class). 

The somewhat controversial history of the concept 

of the Maxillopoda (whether it is monophyletic, 

and if so, which groups should be included, 

and what the relationships are within the group and 

of the group to other crustaceans) is reviewed on 

morphological grounds by Grygier (1983a, b, 

1985, 1987a–c), Müller and Walossek (1988), 

Boxshall and Huys (1989a), Huys (1991), Newman 

(1992), Schram et al. (1997), Schram and Hof 

(1998), and papers cited therein, and on molecular 

grounds by Abele et al. (1992), Spears et al. (1994), 

and Spears and Abele (1997). Some of the fossil 

discoveries since the Bowman and Abele classification 

have a bearing on our understanding of the 

monophyly and definitions of the Maxillopoda as 

well, such as the description of the Skaracarida 


(Müller and Walossek, 1985), the Orstenocarida 

(Müller and Walossek, 1988), and the Mazon 

Creek Cycloidea (Schram et al., 1997). A relatively 

recent and widely used text on invertebrates (Brusca 

and Brusca, 1990) recognizes the Maxillopoda 

(including the Ostracoda), and that text is often cited 

in other listings of crustaceans (e.g., the Tree of 

Life web project; see URL http://ag.arizona.edu/ 

tree/eukaryotes/animals/arthropoda/crustacea/ 

maxillopoda.html), whereas another recent text 

(Gruner, 1993) treats the various maxillopod 

groups separately. 

While it is clear that there is not a single ‘‘good’’ 

character shared by the various maxillopod groups 

(see especially Boxshall, 1992), it is also true that 

some of them seem closely related on morphological 

and molecular grounds. Furthermore, even 

some of the more vocal opponents to the Maxillopoda 

will argue from time to time that there 

seems to be a core group of taxa that ‘‘hang together 

well’’ (although the members of this core 

group change depending on the speaker). The question 

as to which groups are and which are not 

‘‘true’’ maxillopods and whether any of the constituent 

groups should remain allied in a classification 

has not been, in our opinion, satisfactorily 

answered. 

Although the issue is still unresolved, we have 

found it useful to continue to recognize the Maxillopoda, 

and refer the reader to discussions of morphological 

characters seeming to unite the maxillopodan 

groups (see above). At the same time, we 

caution readers that acceptance of the Maxillopoda 

as monophyletic and acceptance of the constituent 

groups are not universal and nowhere near as finalized 

as envisioned by Walossek (1993; see review 

of this work by Martin, 1995) or by Walossek 

and Müller (1994). In the latter paper, Walossek 

and Müller state that the ‘‘interrelationships of the 

majority of maxillopod taxa, particularly of the 

thecostracan lineage, are well-founded on morphological, 

ontogenetic, and fossil data.’’ This could 

hardly be further from the truth. We have followed, 

for the most part, the treatment by Newman (1992) 

for higher classification of the Maxillopoda and his 

subsequent work (especially Newman, 1996) for 

lower taxonomic divisions. We differ from Newman’s 

treatment in not using the ‘‘superclass’’ rank, 

in an attempt to be consistent with our other uses 

and categories. This necessitated the creation of 

some lower level taxonomic names (superorders, 

infraorders, etc.) that unfortunately add to the clutter 

of this already confusing assemblage. We also 

differ from Newman’s treatment in that we have 

treated the Rhizocephala as members of the cirripedian 

line (see below), as suggested by J. Høeg 

(pers. comm.) and others (see below). 

Published and unpublished hypotheses of relationships 

within the Maxillopoda are numerous. As 

one example, Walossek and Müller (1998) feel that 

there are two rather clear lines and presented character 

states for each. The first is the ‘‘copepod line,’’ 

including the copepods, mystacocarids, and the extinct 

Skaracarida (which is in keeping with the 

analysis of maxillopod orders by Boxshall and 

Huys, 1989a). The second is the ‘‘thecostracan line’’ 

that includes the tantulocarids, ascothoracidans, facetotectans, 

acrothoracicans, and cirripeds. However, 

this division does not appear to have much 

neontological (e.g., Høeg, 1992a) or molecular 

(Spears et al., 1994; Spears and Abele, 1997) support. 

Some of the major areas of disagreement in 

the various maxillopod hypotheses include whether 

the ostracodes should be included vs. excluded, 

where the Facetotecta belong, where the Tantulocarida 

belong, and the placement (and subdivision) 

of the cirripedes. We have attempted to list the 

more salient of these efforts in the individual sections 

that follow. For an overview of maxillopod 

classification and phylogenetic studies, we refer 

readers to Grygier (1987a, b), Newman (1987), 

Boxshall and Huys (1989a), Boxshall (1992), Huys 

et al. (1993), Spears et al. (1994), and Spears and 

Abele (1997). 

SUBCLASS THECOSTRACA 

Spears et al. (1994) concluded, based on 18S rDNA 

sequence data, that the Thecostraca, as recognized 

by Grygier (1987a; see also Grygier, 1987b) and 

Newman (1987, 1992) on morphological grounds, 

is a monophyletic assemblage. Furthermore, within 

the Thecostraca, Spears et al. (1994) recognized 

two major subdivisions, one containing the Ascothoracida 

and a second (a modified ‘‘Cirripedia’’) 

containing the Acrothoracica, Rhizocephala, and 

Thoracica. Although we have maintained the Thecostraca, 

we have not divided the group as suggested 

by Spears et al., treating instead the Facetotecta 

(which was not treated by Spears et al.), 

Ascothoracida, and Cirripedia (now including the 

Acrothoracica, Rhizocephala, and Thoracica) as 

taxa of equivalent rank (infraclasses in the current 

scheme) within the Thecostraca. Huys et al. (1993) 

recognized the Thecostraca (without the tantulocarids) 

and postulated a sister-group relationship 

between the Tantulocarida and Thecostraca, noting 

that ‘‘inclusion of the Tantulocarida in the Thecostraca, 

as proposed by Newman (1992), would significantly 

dilute the otherwise robust concept of the 

Thecostraca.’’ Jensen et al. (1994b) described cuticular 

autapomorphies (details of the lattice organs; 

see also Høeg et al., 1998) that also support the 

Thecostraca as a monophyletic assemblage. 

INFRACLASS FACETOTECTA 

Surely one of the biggest remaining mysteries of 

crustacean classification is the taxon Facetotecta. 

Credited to Grygier (1985, corrected from 1984 in 

Bowman and Abele by M. Grygier, pers. comm.; 

see also Grygier, 1987a, b, 1996a), the taxon currently 

contains no further taxonomic divisions other 

than a single genus, Hansenocaris Itô, to accommodate 

the curious ‘‘y-larvae.’’ The group consists 


of small (250–620 micrometers) nauplii with a 

vaulted and ornamented cephalic shield, sometimes 

with complex honeycomb patterns, followed by a 

relatively long and ornamented trunk region. The 

intriguing possibility that these planktonic forms 

may be larval tantulocaridans (which would result 

in tantulocaridans being classified under the Facetotecta) 

has also been suggested (M. Grygier and 

W. Newman, pers. comm.), based in part on the 

fact that there are still gaps in the known life cycle 

of tantulocarids following the work of Boxshall 

and Lincoln (1987) and Huys et al. (1993). As Grygier 

(pers. comm.) points out, ‘‘there is a hole in the 

tantulocaridan life cycle where y-larvae might fit 

(i.e., the progeny of the supposedly sexual males 

and females), but it would be a very tough fit.’’ 

Newman (pers. comm.) succinctly describes the 

current state of our knowledge: ‘‘They [facetotectans] 

are the larvae of some very small, parasitic 

maxillopodan, and if not tantulocarids, they are the 

last survivors of some other great free-living radiation 

close to them.’’ A recent review of the Facetotecta 

was provided by Grygier (1996a). 

INFRACLASS ASCOTHORACIDA 

The Ascothoracida have been treated in the past 

sometimes as an order (e.g., by Newman, 1992), 

but that rank is changed to infraclass here to accommodate 

the constituent taxa that have been elevated 

to (or treated as) orders by Grygier (1987a, 

b) and Newman (1987, 1996), whose classifications 

we follow (see also Grygier, 1983a, b, 1987c, 

1996b). Our classification thus includes two families, 

Ascothoracidae Grygier, 1987, and Ctenosculidae 

Thiele, 1925, that were not included in the 

Bowman and Abele (1982) listing. Thus, the infraclass 

currently consists of two orders, Laurida and 

Dendrogastrida, each with three families. 

INFRACLASS CIRRIPEDIA 

Whether the Cirripedia should include the Rhizocephala 

(e.g., Høeg, 1992a) or whether the Rhizocephala 

are early offshoots of the cirripedian line 

and not members of the crown group (as in Newman, 

1982, 1987; Grygier, 1983a; Schram, 1986) 

is not settled. However, there appears to be a growing 

consensus that the Rhizocephala and the Cirripedia 

form a monophyletic group. Høeg (1992a) 

provides strong evidence based on larval morphology, 

and Spears et al. (1994) support this with molecular 

data. There is some evidence (both morphological 

and molecular) that Cirripedia, with or 

without the Rhizocephala, may be paraphyletic 

(Newman, 1987; Spears et al., 1994). Our classification 

treats the Cirripedia as one of three infraclasses 

of the subclass Thecostraca. Included in our 

Cirripedia are the Rhizocephala. This is more in 

line with Høeg’s (1992a) view, where he suggested 

that Cirripedia be defined as containing the Rhizocephala, 

Thoracica, and Acrothoracica, than 

with Newman’s (1992) view, although Newman 

(pers. comm.) has indicated to us more recently that 

he now agrees with placing the rhizocephalans 

within the Cirripedia. Characters of the naupliar 

and cypris larval stages argue for inclusion of the 

rhizocephalans within the Cirripedia (Høeg, 

1992a), and molecular evidence (in the form of 

rRNA sequences) supports this (Spears et al., 

1994). A close relationship between Rhizocephala 

and Thoracica is supported by 18S rDNA data as 

well (Abele and Spears, 1997). 

Although earlier molecular studies (Spears et al., 

1994) seemed to indicate that the Ascothoracida 

might be the sister taxon to the Acrothoracica 

(which we have included in the Cirripedia), further 

analyses have not supported this arrangement 

(Spears and Abele, 1997). Thus, our current arrangement 

maintains the inclusion of the Acrothoracica 

within the Cirripedia. 

Treatment of the Iblomorpha as one of four thoracican 

suborders (with no phylogenetic order implied) 

is at least in keeping with the finding of Mizrahi 

et al. (1998) that Ibla is not as different from 

other thoracicans as some earlier workers had supposed 

and should not be treated as near the base 

of the stem of the Thoracica. 

An extensive morphology-based cladistic analysis 

of the Cirripedia Thoracica by Glenner et al. 

(1995), reanalyzed with some characters rescored 

by Høeg et al. (1999), supported the monophyly of 

the Balanomorpha and Verrucomorpha and suggested 

that several groups, among them the Pedunculata, 

Scalpellomorpha, and Chthamaloidea, were 

demonstrably paraphyletic. Yet other major questions 

remained unresolved, and Glenner et al. 

(1995) suggested that the fields of larval ultrastructure, 

early ontogeny, and molecular sequencing 

might be promising areas for future research. Anderson 

(1994:326) presented a slightly different 

classification, where the Cirripedia (which he treats 

as a subclass within the class Thecostraca) comprises 

five superorders (two of which, the Archithoracica 

and Prothoracica, would be new taxa 

coined by him), but this has not been followed by 

many other workers. Naupliar evidence seems to 

support, in general, the classification we have depicted 

within the cirripedes based on adult morphology 

(Korn, 1995). Høeg (1995) presents some 

interesting alternatives based on evolution of the 

sexual system of cirripedes and related groups, 

where again thecostracans and tantulocaridans are 

depicted as sister taxa. 

A study of the brachylepadomorphs (Newman, 

1987) led Newman to abandon thoughts of polyphyly 

in favor of monophyly of the sessile barnacles 

(Newman, 1991, 1993, 1996, and pers. comm.). 

Thus, the Sessilia was resurrected to contain the 

brachylepadomorphs, verrucomorphs, and balanomorphs, 

as was the Penduculata for the pedunculate 

barnacles. This has been challenged by Glenner 

et al. (1995) (see above and see also the reanalysis 

of the Glenner at al. data by Høeg et al., 1999). 

A review of various bodies of information con- 


cerning barnacle evolution (Schram and Høeg, 

1995) reveals mostly that we still have much to 

learn about the relationships of the various groups 

of maxillopods. 

SUPERORDER ACROTHORACICA 

For this group, we have followed the classification 

of Newman (1996), where acrothoracicans are divided 

among two orders, Pygophora (with two 

families) and Apygophora (with a single family). 

SUPERORDER RHIZOCEPHALA 

Our classification of this group follows Høeg 

(1992), Høeg and Rybakov (1992), Høeg and Lützen 

(1993, 1996), Huys (1991), and Lützen and 

Takahashi (1996). Thus, we treat the Rhizocephala 

as an infraclass that contains two orders, Kentrogonida 

(with three families) and Akentrogonida 

(with six families), although there is concern that 

one or both of these orders may be paraphyletic 

(see Høeg and Lützen, 1993). Jensen et al. (1994a, 

b) supported monophyly of the Akentrogonida on 

the basis of details of the lattice organs. 

Within the Kentrogonida, concerning the issue of 

authorship of the families Peltogastridae and Sacculinidae 

(which we had earlier credited to Boschma), 

W. Vervoort writes (pers. comm.): ‘‘. . . both 

the families Peltogastridae and Sacculinidae must 

be ascribed to Lilljeborg, 1860. This has been duly 

checked. Boschma lived [from] 1893–1976 and 

cannot possibly be the author of these two families. 

Holthuis and I consulted Lilljeborg’s 1860 publication, 

a copy of which is in our library; there is 

not a shadow of a doubt concerning his authorship.’’ 

The family Sylonidae (Sylidae in Bowman 

and Abele) has been subsumed within the Clistosaccidae 

Boschma, which is now included in the 

Akentrogonida (J. Høeg, pers. comm.). 

Within the Akentrogonida, three new families 

(Duplorbidae, Mycetomorphidae, and Thompsoniidae) 

were described by Høeg and Rybakov (1992) 

and one new family (Polysaccidae) was added by 

Lützen and Takahashi (1996). The Chthamalophilidae 

is recognized as a valid family (also following 

Høeg and Rybakov, 1992), and, as noted above, 

the Clistosaccidae was transferred into the Akentrogonida 

from the Kentrogonida. 

SUPERORDER THORACICA 

Although few new extant families have been suggested 

since 1982, there have been significant rearrangements 

of the cirripedes (or attempts to rearrange 

them) by workers using morphological and 

molecular data. Perhaps the most comprehensive is 

the cladistic study by Glenner et al. (1995), who 

concluded that many currently recognized groups 

appear to be paraphyletic, including the groups that 

appear in our classification under the headings ‘‘Lepadomorpha’’ 

and ‘‘Pedunculata.’’ However, Glenner 

et al. (1995) also noted that ‘‘we have far to go 

before a new taxonomy can emerge’’ and suggested 

the continued use of such commonly used terms as 

‘‘lepadomorphs’’ or ‘‘pedunculates’’ as long as 

workers understand that these are groupings more 

of convenience than of common descent. We are 

not in agreement with this philosophy and would 

prefer to recognize taxa that reflect common descent, 

but in this group, it is apparent that we are 

not yet at the point where we know which clades 

are valid. 

For the most part, we have followed the classification 

of the Thoracica given by Newman (1996). 

Thus, we are recognizing the order Pedunculata (an 

old name that was previously thought to lack validity 

but that Newman (1996) feels is a natural 

assemblage and thus has resurrected) as containing 

four suborders. Some of the names in this order 

(e.g., Heteralepadomorpha, Iblomorpha, Scalpellomorpha) 

are credited to Newman (1987), although 

it is clear that these higher taxon names are 

based on older works, which perhaps should be 

credited as the taxon author and date if we were 

to closely adhere to ICZN article 50.3.1 as extended 

to higher taxa. Many of the families now treated 

in these four suborders were elevated from subfamily 

status by Newman (1987). For example, within 

the Scalpellomorpha, only the family Scalpellidae 

Pilsbry is also found in the Bowman and Abele 

(1982) classification. Within the resurrected order 

Sessilia (see Newman, 1987; Buckeridge, 1995), the 

brachylepadomorph family Neobrachylepadidae 

was described by Newman and Yamaguchi (1995) 

and the verrucomorph family Neoverrucidae was 

described by Newman (1989, in Newman and Hessler, 

1989:268; see also Newman, 1989). Within 

the Balanomorpha, Buckeridge (1983) added the 

superfamily Chionelasmatoidea, containing the single 

family Chionelasmatidae. Suggestions for evolutionary 

radiations within the Balanomorpha were 

presented by Yamaguchi and Newman (1990). A 

recent molecular analysis of several thoracican taxa 

(Harris et al., 2000) suggests that the sessile barnacles 

are monophyletic but that the pedunculate 

forms (our Pedunculata) may not be. 

SUBCLASS TANTULOCARIDA 

The Tantulocarida, bizarre parasites of other deepsea 

crustaceans, were known as early as the beginning 

of the 20th century (reviewed by Huys, 1990e, 

1991; Boxshall, 1991, 1996) but were recognized 

as a distinct class of Crustacea only in 1983 

(Boxshall and Lincoln, 1983), just too late for inclusion 

by Bowman and Abele (1982). They have 

since been relegated to a subclass or infraclass within 

the Thecostraca or have been proposed as the 

sister group to the Thecostraca within the Maxillopoda 

(e.g., Boxshall and Huys, 1989a; Boxshall, 

1991; Huys et al., 1993). Our classification follows 

that of Huys (1990e) (see also Huys, 1991, where 

two families are also described). Discussions of the 

relationships of tantulocaridans (all of which lack 


ecognizable cephalic limbs, other than paired antennules 

in one known stage, which makes elucidation 

of their affinities very difficult) to other 

Crustacea can be found in the above works as well 

as in Boxshall and Lincoln (1987) and Huys et al. 

(1993). Newman (pers. comm.) feels that, based on 

the placement of the male and female genital apertures 

and based also on the fact that the male 

genital aperture empties at the end of a median intromittant 

organ, tantulocarids are so closely related 

to the Thecostraca that placement within the 

subclass Thecostraca may be warranted. Certainly 

they appear more closely related to the Thecostraca 

than to any other maxillopodan group (W. Newman, 

pers. comm.; J. Høeg, pers. comm.; and some 

of the above references). However, for the present 

classification, we have retained them as a separate 

group within the Maxillopoda but not within the 

Thecostraca. Separate status of the Thecostraca and 

Tantulocarida was also suggested on morphological 

grounds by Boxshall and Huys (1989a), although 

we have not closely followed their proposed arrangement 

(their fig. 6) for the organization of the 

Maxillopoda. 

The unusual and confusing life cycle of the tantulocarids 

is now more completely known, thanks 

to the work of Boxshall and Lincoln (1987) and 

Huys et al. (1993). Based on these works and because 

of the gap still remaining in the known tantulocarid 

life cycle, the possibility that y-larvae (the 

Facetotecta) might belong to this taxon has at least 

been considered (M. Grygier, pers. comm.; see earlier 

discussion under Facetotecta). 

SUBCLASS BRANCHIURA 

To our knowledge, this subclass, containing a single 

order and family, has not changed since Bowman 

and Abele (1982) (see also Gruner, 1996). Bill Poly 

(pers. comm.) alerted us to the fact that, although 

the order Arguloida is often credited to Rafinesque 

(1815), Rafinesque employed only the term ‘‘Argulia’’ 

without treating it as a family or order. The 

first person to use the name as an order was apparently 

S. Yamaguti (1963, as Argulidea) (B. Poly, 

pers. comm.). Bowman and Abele (1982) credited 

the family name to Leach (1819) (as did Yamaguti, 

1963, and Gruner, 1996). Although Leach’s usage 

appeared after Rafinesque’s work, we have credited 

Leach with recognition of the family and Yamaguti 

(1963) for the order, despite Rafinesque’s original 

(1815) use of ‘‘Argulia,’’ which of course became 

the basis of both family and order names. Yamaguti 

(1963) also established the family Dipteropeltidae, 

and some subsequent workers (e.g., Overstreet et 

al., 1992; Young, 1998) have continued to recognize 

it, although we do not. 

SUBCLASS PENTASTOMIDA 

One of the most contentious changes in the new 

classification is the inclusion within the Crustacea 

Maxillopoda of the former phylum Pentastomida, 

all members of which are, as adults, parasites in the 

respiratory passages of vertebrates (see reviews by 

Riley, 1986, and Palmer et al., 1993). An alliance 

between pentastomes and branchiuran crustaceans 

was first suggested on the basis of sperm morphology 

some 29 years ago (Wingstrand, 1972; see also 

Wingstrand, 1978; Riley et al., 1978; Grygier, 

1983). Inclusion of pentastomes among the Crustacea 

was actually considered but rejected by Bowman 

and Abele (1982), who at the time felt that 

insufficient evidence was available on that issue. 

Ironically, it was Abele et al. (1989) (see also Abele 

et al., 1992) who finally confirmed this relationship 

(although some would debate whether this was 

confirmed or not) by comparison of 18S rRNA sequences. 

Additional supporting spermatological evidence 

has accumulated since that publication (e.g., 

Storch, 1984; Storch and Jamieson, 1992). Storch 

and Jamieson (1992) concluded that ‘‘a sister-group 

relationship of pentastomids and Branchiura . . . is 

confirmed’’ and that ‘‘the sperm of the pentastomebranchiuran 

assemblage appear to be the most 

highly evolved of the flagellate crustacean sperm.’’ 

Some modern invertebrate texts now treat the pentastomids 

as crustaceans (e.g., Brusca and Brusca, 

1990; Ruppert and Barnes, 1994). Brusca and Brusca 

(1990) mention additional evidence such as similarities 

in the type of embryogenesis, cuticular fine 

structure, and arrangement of the nervous system. 

Nevertheless, the amazing discovery of fossils 

from Middle Cambrian limestones that are extremely 

similar to extant pentastomes (Walossek 

and Müller, 1994; Walossek et al., 1994) would 

seem to cast doubt on placing them within the 

Crustacea (see discussions in Walossek and Müller, 

1994, 1998; also Almeida and Christoffersen, 

1999) and certainly would argue against their being 

maxillopods. If these fossils are indeed related to 

modern-day pentastomids (an issue we feel is not 

yet settled, but see Almeida and Christoffersen, 

1999, for a dissenting opinion), then this finding 

would dispel any notion that the pentastomes are 

a recently derived group. Walossek and Müller 

(1994) make the point that, if pentastomids are related 

to branchiurans, then the morphology of the 

two groups as well as their modes of development 

have differed markedly for more than 500 million 

years, such that present day similarities of their 

sperm morphology might seem to carry less weight. 

If the Cambrian fossils are indeed pentastomids— 

appearing hundreds of millions of years before 

most of their present day hosts were on the scene— 

we must rethink whether we can accept such a major 

divergence in body plan so soon after the Crustacea 

itself appears in the fossil record. Thus, our 

inclusion of them here represents an acceptance of 

the available molecular and sperm morphology 

data (for additional molecular support, see Garey 

et al., 1996, and Eernisse, 1997) over apparently 

sound fossil evidence to the contrary; this may 

prove to be an error. Brusca (2000) suggests a way 

to reconcile the issues if early pentastomids were 


parasites of early fish-like vertebrates as represented 

by the conodonts, many of which were present in 

the Cambrian. 

The classification we follow for the pentastomids 

is from Riley (1986; see also Riley et al., 1978). 

This classification has been questioned recently by 

Almeida and Christoffersen (1999), who do not 

consider pentastomes to be crustaceans. Almeida 

and Christoffersen suggest, based on a cladistic 

analysis of available genera, the recognition of the 

Raillietiellida as a new order to contain their new 

family Raillietiellidae (for the genus Raillietiella), 

the recognition of the Reighardiida as a new order 

to contain the family Reighardiidae, and the dissolution 

of the family Sambonidae. Additionally, 

the Porocephalida was partitioned by them into 

two superfamilies. We have not followed the Almeida 

and Christoffersen (1999:702) classification 

here. 

Authority for the taxon name Pentastomida was 

somewhat difficult to decipher. Riley (pers. comm.) 

informs us that the name ‘‘Pentastomum’’ was first 

employed by Rudolphi (1819) to refer to a single 

species, and several workers (e.g., Almeida and 

Christoffersen, 1999) credit the taxon name Pentastomida 

to Rudolphi. We have been unable to locate 

a work by Rudolphi in 1819 and suspect that 

Rudolphi, 1809, was the intended reference, as Rudolphi 

described the genus Pentastoma and used 

the group name Pentastomata in this 1809 work 

(L. Holthuis, pers. comm.). Diesing (1836) first 

used it (as Pentastoma) for the entire group, although 

the rank was not given. Elevation to phylum 

status was not suggested until 1969 (Self, 

1969), although his evidence and reasoning were 

flawed (Riley, pers. comm.). Prior to that, there 

were various spellings and ranks assigned (e.g., by 

Heymons, 1935; Fain, 1961; and others; see Riley, 

1986). Thus, because Diesing was the first to use 

the name Pentastoma for the entire assemblage, we 

have attributed the authorship of the Pentastomida 

to him. 

Riley (1986) also was of the opinion that pentastomids 

were allied with arthropods and probably 

with crustaceans, noting that ‘‘the available evidence 

overwhelmingly indicates that pentastomids 

are euarthropods and, more specifically, that their 

affinities are closer to crustaceans than uniramians.’’ 

More recently, however, he has indicated that 

the return to the status of separate phylum is probably 

warranted (pers. comm., 1998). Riley’s (1986) 

classification (his table 1), which we have followed, 

recognized nine families in two orders. Two suborders 

of the Porocephalida are mentioned in Riley’s 

text, but he chose not to recognize them in his 

table, and we have followed his lead. 

The inclusion of pentastomids among the Crustacea 

takes the known morphological diversity and 

lifestyle extremes of the Crustacea—already far 

greater than for any other taxon on earth—to new 

heights. How many other predominantly marine invertebrate 

taxa can claim to have representatives 

living in the respiratory passages of crocodilians, 

reindeer, and lions? 

SUBCLASS MYSTACOCARIDA, ORDER 

MYSTACOCARIDIDA 

To our knowledge, there have been no suggested 

changes in the classification of, or in our understanding 

of the phylogeny of, the mystacocarids 

since Bowman and Abele (1982). The subclass continues 

to be represented by a single extant order 

(Mystacocaridida) and family (Derocheilocarididae). 

In their review of crustacean relationships 

based on 18S rDNA, Spears and Abele (1997) noted, 

within the maxillopodan groups, that ‘‘the long 

branch leading to the first lineage, the Mystacocarida, 

indicates extensive divergence relative to other 

crustaceans.’’ Schram et al. (1997) suggest a mystacocarid 

� copepod lineage; a relationship with 

copepods has also been suggested by Boxshall and 

Huys (1989) and Walossek and Muller (1998). The 

group was most recently reviewed by Boxshall and 

Defaye (1996) and Olesen (2001). 

SUBCLASS COPEPODA 

What could have been a truly daunting task for us 

has been made considerably easier by the relatively 

recent publication of Copepod Evolution by Huys 

and Boxshall (1991), by Damkaer’s (1996) list of 

families of copepods (along with their type genus), 

and by three recent treatments of copepods by Razouls 

(1996, free-living copepods), Raibaut (1996, 

parasitic copepods), and Razouls and Raibaut 

(1996, phylogeny and classification). Our acceptance 

of the Huys and Boxshall classification resulted 

in 26 families that have been added, while 

18 families recognized by Bowman and Abele have 

been replaced, resulting in a net gain of 8 families. 

Additional families have been described or recognized 

since then (listed below). Huys and Boxshall 

(1991) proposed some rather sweeping changes in 

some of the higher taxonomic levels as well. Indeed, 

most of the suborders and superfamilies appearing 

in the Bowman and Abele (1982) list have been 

suppressed. This tack was taken also by Damkaer 

(1996), although he does not cite Huys and Boxshall. 

Where the two classifications differ, we tended 

to follow Huys and Boxshall (1991), and readers 

are referred to that tome for arguments underlying 

these changes. However, we must also point out 

that not everyone has accepted the changes suggested 

by Huys and Boxshall (1991) (see especially 

the critique by Ho, 1994a). Indeed, Huys continues 

to use the superfamily concept in some instances 

(see Huys and Lee, 1999, for the Laophontoidea) 

even though it was not used in Huys and Boxshall 

(1991). W. Vervoort (pers. comm.) reminds us that 

‘‘a subdivision of a subclass the size as that of the 

Copepoda will always remain a matter of personal 

choice,’’ and indeed some of the changes advocated 

by Huys and Boxshall have been corrected by these 

same authors in subsequent personal communica- 


tions, as noted below. These changes represent not 

a capricious nature but our constantly changing understanding 

of a tremendously diverse group of organisms. 

Just prior to the publication of Huys and Boxshall’s 

book, Ho (1990) presented a cladistic analysis 

of the orders of the copepods. The results of 

that analysis differ in several significant ways from 

the classification of Huys and Boxshall (and thus 

from our classification). For example, Ho (1990) 

recognized a gymnoplean clade that included the 

Platycopioidea and Calanoidea, and this clade was 

the sister group to the remaining copepod orders. 

In contrast, Huys and Boxshall (1991) treated the 

Platycopioidea as being outside of the Gymnoplea. 

There are other differences as well, such as the 

placement of the monstrilloids and cyclopoids. Ho 

(1990) consistently placed these taxa near each other, 

whereas Huys and Boxshall (1991) separate 

them in their classification, at least implying that 

they are not closely related. In his subsequent critique 

of the Huys and Boxshall (1991) phylogeny, 

Ho (1994a) pointed out an alternative phylogeny 

where the Misophrioida was depicted as the sister 

group to the remaining seven orders of the Podoplea. 

For an in-depth review of recent attempts at 

producing copepod phylogenies, interested readers 

should consult Huys and Boxshall (1991) and the 

critique by Ho (1994a). A more recent molecular 

study (Braga et al., 1999) of relationships among 

the Poecilostomatoida, Calanoida, and Harpacticoida 

yielded somewhat different results, with the 

Poecilostomatoida depicted as basal to the calanoids 

and harpacticoids, in contrast with the abovementioned 

morphology-based hypotheses. 

The review of copepod phylogeny and classification 

presented by Razouls and Raibaut (1996) 

(based in part on Boxshall, 1983, 1986; Boxshall 

et al., 1984; Por, 1984) recognizes 10 orders of copepods, 

as did Huys and Boxshall (1991). However, 

Razouls and Raibaut (1996) did not list the 

orders under superorders or subclasses, preferring 

instead to treat each order separately and refrain 

from phylogenetic hypotheses (although they reproduce 

the ‘‘tree’’ of important events in the evolution 

of copepods from Boxshall, 1986). Also, the list of 

accepted families within each order is not always 

the same in the two treatments. The included families 

are not always given by Razouls and Raibaut 

(1996), and there are differences in the names and 

dates assigned to some of the families. It is also 

apparent that some phylogenetic information may 

be forthcoming from detailed studies of copepod 

developmental (naupliar and copepodid) stages 

(e.g., see Dahms, 1990, 1993), but the data to date 

are preliminary and incomplete (Dahms, 1990). 

M. Grygier informs us (pers. comm.) that the 

correct date for the many copepod taxa named by 

Giesbrecht should perhaps be 1893 rather than 

1892; he refers to Scott’s (1909) note in the Siboga 

Expedition (a note added to the entry for Giesbrecht’s 

Naples volume in the reference list of Scott, 

1909). We have not seen Scott’s 1909 reference list, 

but the date 1893 has been confirmed by W. Vervoort 

(pers. comm.), who additionally notes that 

Scott was a contemporary of Giesbrecht and that 

there is therefore ‘‘no reason at all to doubt [his] 

accuracy.’’ We have thus used this date (1893) instead 

of the often-used 1892. 

Publications describing or recognizing additional 

families subsequent to Bowman and Abele (1982), 

some of which appeared too late for inclusion in 

(or subsequent to) Huys and Boxshall (1991), are 

listed in the following sections on copepod orders. 

ORDER PLATYCOPIOIDA 

This newly recognized order (established by Fosshagen, 

1985, in Fosshagen and Iliffe, 1985) is 

based on the family Platycopiidae Sars, 1911, and 

currently contains only that family and its four genera 

(Platycopia, Nanocopia, Sarsicopia, and Antrisocopia). 

Because Sars established the family Platycopiidae, 

an argument could be made that Sars 

should be the name associated with the higher taxon 

as well, although most workers credit Fosshagen 

(correctly) and/or Fosshagen and Iliffe (1985). 

ORDER CALANOIDA 

Publications with newly described calanoid taxa include 

Fosshagen and Iliffe (1985; Boholinidae), 

Suarez-Morales and Iliffe (1996; Fosshageniidae), 

Ohtsuka, Roe, and Boxshall (1993; Hyperbionychidae), 

and Ferrari and Markhaseva (1996; Parkiidae). 

Suarez-Morales and Iliffe (1996) also erected 

a superfamily, the Fosshagenioidea, to accommodate 

their new family Fosshageniidae, but in 

keeping with our decision to follow the Huys and 

Boxshall (1991) classification, which avoids superfamilies, 

we have not included that taxon, instead 

listing the Fosshageniidae alphabetically among the 

other calanoid families. The family name Phyllopodidae 

has been replaced (because an older use of 

the name Phyllopus was suppressed only for purposes 

of synonymy and not homonymy; G. Boxshall, 

pers. comm.), and the family name erected to 

replace it is Nullosetigeridae (Soh et al., 1999). The 

very similar spelling of the families Pseudocyclopidae 

and Pseudocyclopiidae, pointed out earlier by 

some readers as a possible error, is in fact correct 

and results from the former being based on the genus 

Pseudocyclops Brady while the latter is based 

on the genus Pseudocyclopia Scott (G. Boxshall, 

pers. comm.). Park (1986) presented a brief discussion 

of calanoid phylogeny (based largely on that 

of Andronov, 1974); more recently, Braga et al. 

(1999) examined relationships among calanoid superfamilies 

using 28s rRNA data. 

ORDER MISOPHRIOIDA 

Two new families of misophrioidans, the Palpophriidae 

and Speleophriidae, both comprising genera 

found in anchialine habitats, were described by 


Boxshall and Jaume (2000, see also 1999). The palpophriids 

and misophriids constitute a clade that is 

the sister group to the Speleophriidae (Boxshall and 

Jaume, 1999). 

ORDER CYCLOPOIDA 

Papers with new cyclopoid taxa include Boxshall 

(1988; Chordeumiidae), Ho and Thatcher (1989; 

Ozmanidae [of interest because this family is based 

on a new genus and species from a freshwater snail, 

making it, according to the authors, the ‘‘first parasitic 

copepod ever recorded from a freshwater invertebrate’’]), 

da Rocha and Iliffe (1991; Speleoithonidae), 

and Ho et al. (1998; Fratiidae). The 

family Thespesiopsyllidae has been removed, as it 

is an objective synonym of Thaumatopsyllidae (see 

McKinnon, 1994). The family Mantridae, originally 

placed in the Poecilostomatoida, was transferred 

to the Cyclopoida by Huys (1990d). 

We initially removed from the cyclopoids the Botrylophyllidae 

and Buproridae, following Huys and 

Boxshall (1991). Illg and Dudley (1980) recognized 

these as subfamilies of the Ascidicolidae (along 

with five other subfamilies), and Huys and Boxshall 

(1991) followed that arrangement. However, Huys 

(pers. comm.) has suggested that the Buproridae 

(and also the Botrylophyllidae; see below) should 

be reinstated. G. Boxshall (pers. comm.) also feels 

that the Ascidicolidae, as constituted, ‘‘is too heterogeneous 

and the Buproridae at least should be 

accorded separate family status.’’ However, the situation 

with the Botrylophyllidae is more problematic, 

one problem being that it is a junior synonym 

of the Schizoproctidae (Illg and Dudley, 1980; G. 

Boxshall, pers. comm.); Boxshall (pers. comm.) 

feels that most, but not all, of the seven ascidicolid 

subfamilies recognized by Illg and Dudley (1980) 

‘‘will eventually be given full family status.’’ Thus, 

we have reinstated the Buproridae but not the Botrylophyllidae. 

The former families Enterocolidae, 

Enteropsidae, and Schizoproctidae were also reduced 

to subfamilies of the Ascidicolidae by Illg 

and Dudley (1980), according to J.-S. Ho (pers. 

comm.). The family Cucumaricolidae was transferred 

here from the Poecilostomatoidea following 

Huys and Boxshall (1991), among other such 

changes (see their book). Other changes to the 

Bowman and Abele (1982) list include the removal 

of the Doropygidae (long known to be a synonym 

of the Notodelphyidae) and the Namakosiramiidae 

(a synonym of the harpacticoid family Laophontidae) 

(J.-S. Ho, pers. comm.; G. Boxshall, pers. 

comm.). Ho (1994b) discussed cyclopoid phylogeny 

(based on cladistic analysis of the 10 families 

known at that time) and concluded that parasitism 

had arisen twice in the group. 

ORDERS GELYELLOIDA AND 

MORMONILLOIDA 

The order Gelyelloida was established by Huys 

(1988) for the family Gelyellidae, treated in the past 

as a harpacticoid family and listed as ‘‘infraorder 

incertae cedis’’ by Bowman and Abele (1982:11). 

The Mormonilloida is unchanged, consisting still of 

the single family Mormonillidae. 

ORDER HARPACTICOIDA 

Papers describing new harpacticoid taxa (or elevating 

former subfamilies) include Huys (1990a, Adenopleurellidae; 

1990b, Hamondiidae, Ambunguipedidae; 

1990c, Cristacoxidae, Orthopsyllidae), 

Por (1986, Argestidae, Huntemanniidae, Paranannopidae 

[revised by Huys and Gee, 1996], Rhizothricidae 

[splitting the polyphyletic Cletodidae]), 

Fiers (1990, Cancrincolidae), Huys and Willems 

(1989, Laophontopsidae, Normanellidae; see also 

Huys and Lee, 1999), Huys and Iliffe (1998, Novocriniidae), 

Huys (1988, Rotundiclipeidae), Huys 

(1993, Styracothoracidae), and Huys (1997, Superornatiremidae). 

Huys and Lee (1999) elevated to 

family level the Cletopsyllinae, formerly a subfamily 

of the Normanellidae (following Huys and Willems, 

1989). The Paranannopidae established by 

Por (1986) was relegated to a subfamily of the 

Pseudotachidiidae by Willen (1999); the Pseudotachidiidae 

was formerly a subfamily of the Thalestriidae. 

Huys et al. (1996) referred to this assemblage 

(the Paranannopidae) as the Danielsseniidae 

Huys and Gee because Paranannopidae was based 

on an unavailable genus name. Thus, the family 

Paranannopidae (� the Danielsseniidae of Huys et 

al., 1996) does not appear in our list, as it is considered 

a subfamily of the Pseudotachiidae following 

Willen’s (1999) preliminary study. The subfamily 

Leptastacinae Lang was upgraded to a family by 

Huys (1992). The family Gelyellidae, treated by 

Bowman and Abele (1982) as a harpacticoid family, 

was transferred to its own order, Gelyelloida, 

by Huys (1988). Relationships among the laophontoidean 

families were addressed by Huys (1990b) 

and Huys and Lee (1999). 

Arbizu and Moura (1994) found the family Cylindropsyllidae 

polyphyletic and elevated the former 

subfamily Leptopontiinae to family level (family 

Leptopontiidae). Although they also suggested 

that the family Cylindropsyllidae should be relegated 

to a subfamily of the Canthocamptidae, we 

have retained the family Cylindropsyllidae for now 

(and on the advice of R. Huys, pers. comm.). 

ORDER POECILOSTOMATOIDA 

Papers describing new poecilostomatoid taxa include 

Humes (1986, Anthessiidae), Humes and 

Boxshall (1996, Anchimolgidae, Kelleriidae, Macrochironidae, 

Octopicolidae, Synapticolidae, 

Thamnomolgidae), Avdeev and Sirenko (1991, Chitonophilidae 

[incomplete description; tentative 

placement in the Poecilostomatoida is based on 

pers. comm. from W. Vervoort, A. Humes, and G. 

Boxshall]), Ho (1984, Entobiidae, Spiophanicolidae), 

Humes (1987, Erebonasteridae), Marchenkov 

and Boxshall (1995, Intramolgidae), Huys and 


Böttger-Schnack (1997, Lubbockiidae), Lamb et al. 

(1996, Nucellicolidae), Boxshall and Huys (1989b, 

Paralubbockiidae), Ho and Kim (1997, Polyankylidae). 

The family Phyllodicolidae was transferred 

here from the cyclopoids by Huys and Boxshall 

(1991) (it still appears as a cyclopoid in Damkaer, 

1996). 

Also within the poecilostomatoids, the Lernaeosoleidae 

was elevated (from the Lernaeosoleinae 

Yamaguti, 1963) by Hogans and Benz (1990). The 

family Amazonicopeidae proposed by Thatcher 

(1986) has not been recognized; it is thought to be 

a synonym of the Ergasilidae by G. Boxshall (pers. 

comm.) and J. Ho (pers. comm., and citing Amado 

et al., 1995). The family Anomopsyllidae (included 

in Bowman and Abele, 1982, and in Huys and 

Boxshall, 1991) is not listed here. According to G. 

Boxshall (pers. comm.), ‘‘the genus Anomopsyllus 

was included in the Nereicolidae by Stock (1968), 

the family Anomopsyllidae thus becoming a junior 

synonym of the Nereicolidae.’’ Laubier (1988) (unfortunately 

overlooked by Huys and Boxshall, 

1991) described both sexes of the genus and confirmed 

that it is a nereicolid. The family Vaigamidae 

proposed by Thatcher and Robertson (1984) also 

is not included here, as it was shown to be a synonym 

of the Ergasilidae by Amado et al. (1995). 

The Nucellicolidae, although retained for now, may 

prove to be a junior synonym of the Chitonophilidae 

(R. Huys, pers. comm.). Finally, the family Micrallectidae 

has been established recently (Huys, 

2001) to accommodate poecilostomatoid genera associated 

with pteropods. 

Ho (1984) suggested phylogenetic relationships 

among the nereicoliform families, indicating three 

main lines of evolution. Later, Ho (1991) conducted 

a more thorough analysis of the 47 known poecilostomatoid 

families, which remains the most indepth 

study of poecilostomatoid relationships while 

at the same time being somewhat preliminary in 

nature. Relationships of 10 poecilostomatoid families 

(in the lichomolgoid complex) are presented by 

Humes and Boxshall (1996). Unfortunately, we 

could not follow their suggestions here because of 

the absence of knowledge concerning the other 

(nonlichomolgoid) poecilostomatoid families. 

ORDER SIPHONOSTOMATOIDA 

Papers describing new siphonostomatoid taxa include 

Izawa (1996, Archidactylinidae [questionable, 

as this is an incomplete description]), Humes 

and Stock (1991, Coralliomyzontidae), and Humes 

(1987, Ecbathyriontidae). The Herpyllobiidae 

(treated as siphonostomes by Huys and Boxshall, 

1991) have been removed to the Poecilostomatoidea 

(R. Huys, pers. comm.). Two new families, Dichelinidae 

and Codobidae, have been proposed recently 

for siphonostomatoid genera parasitic on 

echinoderms (Boxshall and Ohtsuka, 2001), and 

the family Scottomyzontidae (erected for Scottomyzon 

gibberum, a symbiont of the asteroid Aste- 

rias rubens) was established by Ivanenko et al. 

(2001). 

ORDER MONSTRILLOIDA 

With the transfer of the Thaumatopsyllidae to the 

Cyclopoida (Huys and Boxshall, 1991; Grygier, 

pers. comm.), the order Monstrilloida has been reduced 

to a single family, Monstrillidae, which now 

is credited to Dana rather than to Giesbrecht following 

ICZN Opinion 1869 (M. Grygier, pers. 

comm.). 

There have also been many additional changes to 

the list of copepod families that are not detailed 

here—including additions, deletions, reinstatements 

of older families, corrected spellings and authors, 

etc.—suggested by various workers, mostly Ju-Shey 

Ho, Arthur Humes, H.-E. Dahms, G. Boxshall, and 

Rony Huys. In some cases, we did not ask for a 

published reference, instead taking these workers at 

their word (and also because in some cases the suggestion 

has not been published). 

CLASS OSTRACODA 

This section received extensive input from Dr. Anne 

Cohen, and our treatment of this group is in many 

ways based on her impressive knowledge of this 

taxon. Major references included Morin and Cohen 

(1991), Martens (1992), Whatley et al. (1993), 

Hartmann and Guillaume (1996), Martens et al. 

(1998), and Cohen et al. (1998). 

Many previous workers have considered ostracodes 

to be a subclass of the Maxillopoda. The 

strongest reason for including ostracodes among 

maxillopods is, apparently, the presence in ostracodes 

of a naupliar eye with three cups and tapetal 

cells between the sensory and pigment cells (e.g., 

see Elofsson, 1992; Huvard, 1990; and earlier papers 

cited in these works). This feature is found also 

in the Thecostraca, Branchiura, and Copepoda, and 

for this reason, Schram (1986), Brusca and Brusca 

(1990), and others have placed ostracodes within 

the Maxillopoda (see also discussions in Grygier, 

1983a; Boxshall, 1992; Elofsson, 1992; Cohen et 

al., 1998). Schram (pers. comm., and citing K. 

Schultz, Das Chitinskelett der Podocopida und der 

Frage der Metamerie dieser Gruppe, doctoral dissertation, 

University of Hamburg, which we have 

not seen) informs us that an additional apomorphy 

that argues for inclusion of ostracodes within the 

Maxillopoda is the location of the gonopods. 

Swanson’s (1989a, b, 1990, 1991) discovery of living 

specimens of the primitive ostracode genus 

Manawa (family Punciidae) caused him to suggest 

the inclusion of ostracodes within the Maxillopoda 

as well. Cohen et al. (1998) note the following 

‘‘perhaps homologous morphological characters’’: a 

medial naupliar eye that has three cups and a tapetal 

layer (present in most Myodocopida and in 

many Podocopida), and overall reduction in body 

size and limb number. 

However, other workers are quick to point out 


that reduction in body segmentation has occurred 

independently as a functional adaptation in many 

different and unrelated crustacean taxa and that the 

unique features of the Ostracoda argue for their 

recognition as a separate class (see especially discussions 

in Newman, 1992; Boxshall, 1992; Wilson, 

1992). Treatment of ostracodes as a subclass 

of the Maxillopoda has additional problems as 

well. Wilson (1992) could not find support for placing 

the former within the latter based on morphological 

grounds (although Schram and Hof, 1998, 

point out errors in Wilson’s analysis that, if corrected, 

would indeed group ostracodes with one 

cluster of Maxillopoda). Abele et al. (1992) rejected 

the inclusion of ostracodes in the Maxillopoda on 

molecular grounds. Spears and Abele (1997) suggest 

the possibility that, based on molecular data, 

both Ostracoda and Maxillopoda might be paraphyletic. 

There is also some evidence, both morphological 

and molecular, that the two major groupings of the 

Ostracoda (Myodocopa and Podocopa) may not 

constitute a monophyletic assemblage (e.g., see 

Vannier and Abe, 1995; Spears and Abele, 1997). 

On the other hand, Cohen et al. (1998), based on 

the many similarities between these two groups, 

‘‘regard it more parsimonious and useful to assume 

that they do.’’ This older view—that ostracodes are 

monophyletic—has been adopted here and is in fact 

held by a majority of current workers in the field. 

The assignment of ostracodes to a group ‘‘Entomostraca’’ 

(which included, in addition to ostracodes, 

the Branchiopoda, Cirripedia, Branchiura, 

and Phyllocarida) by McKenzie et al. (1983) was 

clearly an unsupported departure (see also discussions 

on Branchiopoda and Phyllocarida and notes 

on Entomostraca under the general heading Crustacea). 

A modified version of the classification of the Ostracoda 

used by Whatley et al. (1993), which will 

be the basis for the classification used in the upcoming 

revision of the Treatise on Invertebrate Paleontology 

(‘‘more or less,’’ according to Whatley, 

pers. comm.; R. Kaesler, pers. comm.), was sent to 

us by R. Whatley. This classification, which differs 

considerably from what was proposed by Mc- 

Kenzie et al. (1983) and also from the classification 

used by Hartmann and Guillaume (1996), has been 

followed fairly closely. Differences include the spelling 

of the endings of superfamilies. We use the 

ICZN-recommended ending ‘‘–oidea’’ (which in the 

latest (fourth) edition of the International Code of 

Zoological Nomenclature is mandatory rather than 

a recommendation; ICZN, 1999, article 29.2). 

Whatley, in one of the more interesting responses 

we received, has indicated that the ‘‘-oidea’’ spelling 

is an ‘‘attempted imposition’’ by the ICZN. Kaesler 

(pers. comm.) and Whatley (pers. comm.) note that 

ostracodologists prefer to think of the higher 

groups as superfamilies rather than as suborders 

and are also more accustomed to the use of the 

ending ‘‘-acea’’ for superfamilies and thus are more 

familiar with, and prefer, the concept of a superfamily 

Bairdiacea as opposed to a superfamily Bairdioidea 

or suborder Bairdiocopina. On the spelling 

of superfamily names, however, the ICZN recommendation 

(ICZN, 1999, fourth edition, article 

29.2) is rather clear: ‘‘The suffix -OIDEA is used 

for a superfamily name, -IDAE for a family name, 

-INAE for a subfamily name . . .’’ etc. And it appears 

to us that it is primarily the paleontologists 

(who are, we admit, the majority of the ostracodologists) 

rather than neontologists who prefer 

(and use) the ‘‘-acea’’ ending for superfamilies (e.g., 

see Martens, 1992, and Martens et al., 1998, for 

living freshwater ostracode superfamilies, all of 

which are spelled according to ICZN recommendation 

29.A [now 29.2]). As Martens et al. (1998: 

41) explain in a note to accompany their classification, 

‘‘. . . as ostracods are animals, we will follow 

the ICZN throughout this book.’’ 

Thus, we have followed the ICZN recommendation 

(as did Bowman and Abele, 1982, and 

Schram, 1986) for spellings of superfamilies (e.g., 

Bairdioidea, not Bairdiacea). Whatley (pers. 

comm.) also feels that, relative to the Podocopida, 

the Myodocopa is probably ‘‘one hierarchical level 

too high.’’ Whatley (pers. comm.) considers his 

own arrangement (Whatley et al., 1993) ‘‘old fashioned 

but acceptable to people who actually work 

on the group,’’ a justification that we feel is baseless 

but that, at the moment, faces nothing in the way 

of a serious alternative classification. Martens 

(1992) and Martens et al. (1998) appear to base 

their decisions more on shared derived characters 

and more often than not employ characters of the 

entire animal (as opposed to those of the shell 

only). Consequently, we have followed their lead 

for the names, spellings, and arrangement of the 

superfamilies and families of the freshwater families 

as far as was possible (not all families are treated 

in those works). Thus, although Whatley would remove 

the superfamilies Macrocypridoidea and Pontocypridoidea 

(placing their families among the Cypridoidea), 

we have maintained these groupings 

following Martens (1992) and Martens et al. 

(1998). Whatley (pers. comm.) also feels that the 

family Saipanettidae (� Sigilliidae; see later) is no 

more than a subfamily of the Bairdiidae, whereas 

Martens (1992) recognized a separate superfamily, 

the Sigillioidea Mandelstam, to accommodate this 

unusual group, and here again we have followed 

Martens (1992). 

Whatley (pers. comm.) and Whatley et al. (1993) 

also place the unusual and primitive family Punciidae 

in the Platycopida (he considers Manawa to be 

a member of the Cytherellidae), indicating that 

there are still no living members of the Palaeocopidae. 

Martens et al. (1998) also feel that there are 

no living palaeocopids, which also supports transfer 

of the punciids. We have followed Whatley’s advice 

in moving the punciids to the Platycopida (although 

they appear to share no unique characters 

with platycopids and differ in many respects), but 


we have retained them in their own family, the Punciidae, 

as we are not aware of any publications that 

demonstrate that they belong among the cytherellids. 

Possibly a better solution would have been to 

list them as incertae sedis for now. 

Although there have been rearrangements of the 

Ostracoda, there have been surprisingly few higher 

taxa described or recognized since Bowman and 

Abele (1982) and Cohen (1982). The fossil bradoriids 

and the ‘‘phosphatocopines’’ of Sweden’s Upper 

Cambrian ‘‘Orsten’’ fauna are no longer considered 

true ostracodes. Walossek and Müller 

(1998, in Edgecombe) hypothesize that, although 

phosphatocopines are not crown group crustaceans 

(their ‘‘Eucrustacea’’), they may be the sister taxon 

to this group. 

SUBCLASS MYODOCOPA 

Arrangement of families in the Myodocopa follows 

Kornicker (1986:178), which in turn was based 

largely on McKenzie et al. (1983), although some 

of the higher taxon spellings have been changed for 

consistency. The suborder Cladocopina may be deserving 

of status as a separate order (A. Cohen, 

pers. comm.), although this step has not been taken 

here (see also Kornicker and Sohn, 1976, who first 

suggested the inclusion of the Cladocopina and 

Halocypridina within the Halocyprida). 

SUBCLASS PODOCOPA 

The superfamilies Bairdioidea and Cytheroidea 

have been elevated to suborders, with spelling 

changed to Bairdiocopina and Cytherocopina (respectively) 

(following Martens, 1992, and A. Cohen, 

pers. comm.). Alexander Liebau (pers. comm.) 

informs us that the Cytherocopina has been divided 

by him (Liebau, 1991; not seen by us) into two 

infraorders, the Nomocytherinina (which includes 

species showing epidermal cell constancy reflected 

by mesh constancy of reticulate sculptures) and the 

Archaeocytherinina, containing the paraphyletic remaining 

cytherocopines. We have not used this division 

here. Within the Cytheroidea, we have used 

the list of families supplied by R. Whatley (pers. 

comm.), based in part on Whatley et al. (1993) and 

on his anticipation of the Ostracoda section of the 

next edition of the Treatise on Invertebrate Paleontology 

(Whatley, pers. comm.; R. Kaesler, pers. 

comm.). The family Bonaducecytheridae McKenzie 

has been removed (R. Maddocks, pers. comm.). 

The superfamily Terrestricytherioidea and its sole 

family, the Terrestricytheridae, have been removed; 

Martens et al. (1998), citing Danielopol and Betsch 

(1980), note that Terrestricypris is a modified member 

of the Candonidae (the spelling of which has 

been corrected from Candoniidae; R. Maddocks, 

pers. comm.). 

The suborder Metacopina now contains only fossils 

and thus has been removed from our classification, 

as the Darwinulocopina has now been established 

by Sohn (1988) to accommodate the fam- 

ily Darwinulidae (A. Cohen, pers. comm.). The former 

superfamily Cypridoidea is now treated as a 

suborder, Cypridocopina Jones (Martens et al., 

1998). The family Paracyprididae has been removed; 

this group also is now thought to be a subfamily 

of the Candonidae (Martens et al., 1998). 

The Cypridopsidae has been removed (Martens et 

al., 1998). The family Saipanettidae, formerly in 

the superfamily Healdioidea (which has been removed), 

also has been removed. The Saipanettidae 

was found to be a junior synonym of the Sigilliidae, 

an extant family reviewed recently by Tabuki and 

Hanai (1999). Spelling of the Sigilliidae was initially 

given as Sigillidae by Tabuki and Hanai 

(1999); we have corrected it based on the spelling 

of the genus Sigillium. The Sigilliidae is now treated 

as a member of the superfamily Sigillioidea (see Tabuki 

and Hanai, 1999; spelling emended from Sigilloidea; 

R. Maddocks, pers. comm.), which in turn 

has been placed in its own suborder, the Sigilliocopina 

(see Martens, 1992). Martens (1992) originally 

suggested recognition at the infraorder level, 

as ‘‘infraorder 3, ‘Sigillioidea.’ ’’ The spelling we use 

for the suborder was first employed by Cohen et al. 

(1998). 

CLASS MALACOSTRACA 

Because of their size and numbers, malacostracans 

have been the subject of a huge number of classificatory 

and phylogenetic studies employing morphological 

characters, molecular characters, or 

both. For the most part, there seems to be agreement 

that the Malacostraca itself is a monophyletic 

grouping (e.g., see Hessler, 1983; Dahl, 1983a, b, 

1991; Mayrat and Saint Laurent, 1996; Shultz and 

Regier, 2000; Watling et al., 2000; Richter and 

Scholtz, in press), although differing opinions can 

certainly be found. There is considerably less agreement 

concerning the constituencies and relationships 

of the various groupings of the Malacostraca, 

and these topics are the subject of a vast body of 

literature (much of which was reviewed recently by 

Richter and Scholtz, in press). Attempts to place 

phyllocarids outside the Malacostraca have largely 

been shown to be misguided (see below). We have 

tried to refer readers to the salient papers that offer 

arrangements that differ from our own in the individual 

sections that follow. 

SUBCLASS PHYLLOCARIDA, ORDER 

LEPTOSTRACA 

The status of the subclass Phyllocarida (which includes 

only one extant order, the Leptostraca) as 

true malacostracans is now fairly well accepted. Arguments 

can be found in Dahl (1987), in rebuttal 

to Schram (1986), who had been in favor of resurrecting 

the older term Phyllopoda to include 

branchiopods, cephalocarids, and leptostracans 

(see also Rolfe, 1981; Dahl, 1992; Martin and 

Christiansen, 1995a; Spears and Abele, 1999; Richter 

and Scholtz, in press; but see also Ferrari, 1988, 


for a rebuttal of Dahl’s criticism). Inclusion of leptostracans 

within the Malacostraca has been further 

supported by molecular evidence (rDNA data 

summarized in Spears and Abele, 1997, 1999; see 

also Shultz and Regier, 2000, for EF-1� and Pol II 

data). Hessler (1984) established the family Nebaliopsidae 

in recognition of the great differences setting 

the genus Nebaliopsis apart from other leptostracans, 

thereby doubling the number of recognized 

families of the extant phyllocarids. However, J. 

Olesen (1999b, and pers. comm.) finds that, depending 

upon the choice of outgroups (and characters) 

used in cladistic analyses of the group (based 

on descriptions in the literature), there is still some 

room for doubt as to whether Nebaliidae is monophyletic 

or paraphyletic (with Nebaliopsis nested 

within the other nebaliacean genera). Most recently, 

Walker-Smith and Poore (2001) have erected a 

third family, Paranebaliidae, to contain the genera 

Paranebalia and Levinebalia (the latter of which 

was described by Walker-Smith, 2000). 

Our treatment of the Phyllocarida follows Hessler 

(1984), Martin et al. (1996), Dahl and Wägele 

(1996), and our PEET web page for Leptostraca 

(URL http://www.nhm.org/�peet/) in recognizing 

two extant families (see Rolfe, 1981, for extinct 

phyllocarids) plus the recently established family 

Paranebaliidae following Walker-Smith and Poore 

(2001). Most authors in the past have credited the 

family Nebaliidae to Baird (1850). However, according 

to L. Holthuis (pers. comm.), Samouelle 

(1819:100) mentioned ‘‘Fam. VI. Nebaliadae’’ [sic] 

in his ‘‘Entomologist’s Useful Compendium,’’ which 

of course predates Baird’s (1850) work. Thus, we 

have attributed the family Nebaliidae to Samouelle, 

1819. 

SUBCLASS HOPLOCARIDA, ORDER 

STOMATOPODA 

Several workers, today and in the past (examples 

include Hessler, 1983; Scholtz, 1995; Richter and 

Scholtz, in press), have considered the hoplocarids 

to be members of the Eumalacostraca, a placement 

that has been used often and in some textbooks as 

well (e.g., Brusca and Brusca, 1990). However, we 

have retained their placement as a separate subclass 

within the Malacostraca pending further exploration 

of this question (see review by Watling et al., 

2000). Our treatment of the hoplocarids as separate 

from the other Eumalacostraca also is consistent 

with some (admittedly weak) molecular evidence 

(see Spears and Abele, 1997, 1999b) and 

with cladistic analyses based mostly on fossil taxa 

(e.g., Hof, 1998a, b; Hof and Schram, 1999). 

Schram (1971, 1986) had argued earlier for separate 

status of the hoplocarids as well. Spears and 

Abele (1997) could state only that the position of 

the ‘‘Hoplocarida relative to the Eumalacostraca is 

equivocal’’ (low bootstrap value) based on rDNA 

sequence data, and thus they were ‘‘unable to determine 

whether hoplocarids represent a separate, 

independent malacostracan lineage with taxonomic 

rank (subclass) equivalent to that of phyllocarids 

and eumalacostracans.’’ Their subsequent paper 

(Spears and Abele, 1999b) seems (to us) to indicate 

somewhat stronger evidence that hoplocarids are 

not eumalacostracans, but the authors are suitably 

cautious in not saying so. Without firm indications 

that we should do otherwise, we have maintained 

separate status for the Hoplocarida and Eumalacostraca. 

Although a thorough cladistic analysis of 

fossil and extant crustacean taxa by Schram and 

Hof (1998) resulted in a tree that showed hoplocarids 

arising from somewhere within the Eumalacostraca, 

these authors also noted that forcing the 

hoplocarids into a ‘‘sister group’’ position to the 

Eumalacostraca increased tree length by only 1%. 

Other workers (e.g., Watling, 1999a), recognizing 

how very derived the stomatopods are, place them 

in the Eumalacostraca as the sister taxon to the Eucarida. 

Most recently, Richter and Scholtz (in press) 

suggested that hoplocarids occupy a basal position 

within the Eumalacostraca. Thus, placement of the 

hoplocarids continues to be an unresolved issue, 

but we felt that the weight of the evidence placed 

them outside, rather than within, the Eumalacostraca. 

Scholtz (pers. comm.) additionally suggests 

that our crediting the name Eumalacostraca to 

Grobben is therefore incorrect, as Grobben included 

the hoplocarids among the Eumalacostraca (but 

see earlier notes on names, dates, and the ICZN). 

Within the Hoplocarida, most of our changes are 

based on the catalog provided by H.-G. Müller 

(1994) and on Manning (1995), and our final arrangement 

of families and superfamilies follows the 

recent cladistic analysis by Ahyong and Harling 

(2000). Publications that describe or recognize families 

or higher taxa of stomatopods subsequent to 

Bowman and Abele (1982) include Manning (1995, 

Indosquillidae, Parasquillidae, Heterosquillidae), 

Manning and Bruce (1984, Erythrosquillidae [for 

which the superfamily Erythrosquilloidea was later 

created by Manning and Camp, 1993]), Manning 

and Camp (1993, Tetrasquillidae), Moosa (1991, 

Alainosquillidae), and Ahyong and Harling (2000, 

superfamilies Eurysquilloidea and Parasquilloidea). 

Concerning phylogeny within the Hoplocarida, 

there is recent evidence from several laboratories 

that the superfamily Gonodactyloidea as presented 

in Bowman and Abele (1982) is not a monophyletic 

grouping (Hof, 1998b; Ahyong, 1997; Barber and 

Erdmann, 2000; Ahyong and Harling, 2000; Cappola 

and Manning, 1998; Cappola, 1999) and that 

within the gonodactyloids the eurysquillids may be 

paraphyletic. These same authors disagree over 

whether the Bathysquilloidea are monophyletic 

(Cappola and Manning, 1998) or not (Ahyong, 

1997). A comparative study of eye design in stomatopods 

(Harling, 2000) also supports a nonmonophyletic 

Gonodactyloidea and questions the fivesuperfamily 

scheme of Müller (1994). The nonmonophyly 

of the Gonodactyloidea necessitates the 

creation of additional families and superfamilies to 


accommodate some of the former gonodactyloid 

taxa (Ahyong, 1997; Ahyong and Harling, 2000; 

Cappola and Manning, 1998). Cappola and Manning 

(1998) also suggested that a new superfamily 

and family (Eurysquilloidoidea, Eurysquilloididae) 

should be established to accommodate the former 

eurysquillid genus Eurysquilloides. We have followed 

the classification suggested by Ahyong and 

Harling (2000). According to their scheme, the 

families Eurysquillidae and Parasquillidae, formerly 

treated as members of the Gonodactyloidea, are 

each deserving of superfamily status, and thus they 

established the superfamilies Eurysquilloidea and 

Parasquilloidea to accommodate them. The Gonodactyloidea 

has been reconfigured and now contains 

the Alainosquillidae, Hemisquillidae, Gonodactylidae, 

Odontodactylidae, Protosquillidae, 

Pseudosquillidae, and Takuidae. The family Heterosquillidae 

established by Manning (1995) has 

been removed, as it was suggested to be a synonym 

of Tetrasquillidae (see Ahyong and Harling, 2000). 

In the most recent treatment, Ahyong (2001) synonymized 

the Harpiosquillidae Manning with the 

Squillidae; thus the Harpiosquillidae is not in our 

list. 

Hof (1998b) recognized two main clades of extant 

stomatopods. One clade included most of the 

gonodactyloid families but excluded the alainosquillids 

and the eurysquillids. The second clade 

contained the remaining extant families and indicated 

possible affinities between the squilloids and 

lysiosquilloids and also between the bathysquilloids 

and erythrosquilloids. Hof (1998b) points out that, 

although his results are preliminary, the fact that 

fossils should be included when at all possible in 

any cladistic analysis is clear and obvious from his 

work. A cladistic analysis of the hoplocarids that 

incorporated Paleozoic taxa was presented by Jenner 

et al. (1998), but it did not resolve relationships 

within the sole extant order (their Unipeltata). In 

the most recent treatment, Ahyong and Harling 

(2000) have also suggested that the recent stomatopods 

have evolved ‘‘in two broad directions from 

the outset,’’ corresponding roughly to the smashing 

and spearing types. 

SUBCLASS EUMALACOSTRACA 

The concept of the Eumalacostraca as a monophyletic 

assemblage has not been seriously challenged, 

with the exception of the question of whether hoplocarids 

belong (see above discussion under Hoplocarida 

for arguments as to their inclusion or exclusion). 

Our classification is roughly similar to 

that of Bowman and Abele (1982) in recognizing 

the Eumalacostraca and its constituent groups, although 

there have been several significant rearrangements 

within and among those groups, as noted 

below (see also Richter and Scholtz, in press). 

Schram (1984a) reviewed characters that defined 

the various eumalacostracan groups recognized at 

that time and presented alternatives to more traditional 

classifications. 

SUPERORDER SYNCARIDA, ORDERS 

BATHYNELLACEA AND ANASPIDACEA 

Monophyly of the Syncarida appears to be fairly 

well accepted (e.g., Schram, 1984b; Richter and 

Scholtz, in press). Within the Bathynellacea, we 

have removed the family Leptobathynellidae, as 

this was synonymized with the Parabathynellidae 

by Schminke (1973:56). Schram (1984b) credits 

both names (Bathynellidae, Bathynellacea) to 

Chappuis (1915), whereas Lopretto and Morrone 

(1998) credit the Bathynellidae to Grobben (as did 

Bowman and Abele, 1982) and the Bathynellacea 

to Chappuis. We have not been able to locate a 

paper by Grobben describing bathynellids and so 

have followed Schram’s (1984b, 1986) lead, crediting 

both taxa to Chappuis (1915). The Anaspidacea 

remains unchanged, with four extant families. 

Thus, our classification of the Syncarida and its 

two orders (Anaspidacea and Bathynellacea) is the 

same as that presented by Lopretto and Morrone 

(1998), where all known syncarid genera are also 

listed, and is essentially the same as the classification 

suggested earlier by Schram (1984a:196) based 

on a phylogenetic analysis of fossil syncarids (excluding 

the entirely fossil order Paleocaridacea). 

SUPERORDER PERACARIDA 

We continue to recognize the Peracarida, treating it 

as a superorder that contains nine orders. This is 

mostly in keeping with Bowman and Abele (1982) 

and most major treatments since that time (see especially 

Hessler and Watling, 1999; Richter and 

Scholtz, in press). However, there have been suggestions 

made to abandon the Peracarida or at least 

significantly revise it (e.g., Dahl, 1983a), and the 

relationships among the various peracarid groups 

(and of peracarids to other groups of crustaceans) 

are very controversial. Schram (1986) advocated 

eliminating the Peracarida of earlier workers, feeling 

that it united groups that were only superficially 

similar. Other workers (e.g., Pires, 1987; Brusca 

and Brusca, 1990; Wagner, 1994; Hessler and Watling, 

1999; Richter and Scholtz, in press) recognize 

the group, but the treatments occasionally differ as 

to which orders are included. Hessler and Watling 

(1999) review major attempts to phyletically order 

the peracarids, including Schram (1986), Watling 

(1983), Wills (1997), and Wheeler (1997), all of 

which have appeared subsequent to the Bowman 

and Abele (1982) classification. There is little agreement 

among these various schemes. Mysidaceans in 

particular are sometimes treated as one order, 

sometimes as the separate orders Lophogastrida 

and Mysida within the Peracarida, and sometimes 

suggested to fall outside of the Peracarida altogether. 

As examples, Watling (1998, 1999b) argues that 

mysids should fall outside the Peracarida and that 


the Amphipoda are deserving of status separate 

from all other peracarids and should constitute 

their own superorder as a sister group to the remaining 

taxa, which would then constitute a reduced 

Peracarida sensu stricto. (Interestingly, if the 

Mysidacea and Thermosbaenacea are removed 

from Watling’s (1981) fig. 1, then the Amphipoda 

would indeed appear as the sister group to all other 

‘‘true’’ peracaridans in that diagram.) But this is not 

in agreement with Wagner (1994), who depicted 

amphipods and isopods as closely related and depicted 

amphipods, isopods, cumaceans, and tanaidaceans 

as a monophyletic clade. Wagner (1994) 

also suggested affinities between the Thermosbaenacea 

and Mictacea and between those two groups 

and the Spelaeogriphacea, whereas Pires (1987) 

treated amphipods and mysidaceans as related taxa 

that were in turn the sister group to all other peracarids. 

In Wagner’s phylogenies, the mysids (both 

Mysida and Lophogastrida) are shown as the sister 

group to the other Peracarida. Depending on where 

the line is drawn, Wagner’s phylogeny could be 

used as an argument for inclusion or exclusion of 

the mysids within the Peracarida. 

Spears and Abele (1997, 1998) have suggested, 

on the basis of molecular data, that the two groups 

of mysidaceans are not monophyletic (suggested 

earlier by Dahl, 1983a, and others based on morphological 

features), with the Lophogastrida grouping 

with other peracarids but with the Mysida falling 

outside that clade (see below). Jarman et al. 

(2000) also concluded (on the basis of 28S rDNA 

sequence data) that the Mysida and Lophogastrida 

are not closely related but posited the Mysida closer 

to the Euphausiacea. Thermosbaenaceans, treated 

as true peracarids by us (see arguments below and 

also Richter and Scholtz, in press), have in the past 

been treated by some workers (e.g., Bowman and 

Abele, 1982; Pires, 1987) as the separate order Pancarida, 

which we have abandoned. A more radical 

departure is suggested by Mayrat and Saint Laurent 

(1996), who suggested a phylogeny (their fig. 342) 

of the Malacostraca in which the peracarids are 

polyphyletic, with amphipods depicted as the sister 

taxon to all other malacostracans (except the leptostracans) 

and with cumaceans and mysids associated 

with the higher eumalacostracans. This, to 

us, seems unlikely. Richter (1999), after a thorough 

analysis of characters of the compound eyes of malacostracans, 

felt that ‘‘Lophogastrida and Mysida 

are clearly members of the Peracarida.’’ These are 

only a few of the suggestions to be found in the 

rather confusing literature on the diverse peracarid 

crustaceans. The most recent coverage is a wonderful 

in-depth treatment of the entire Peracarida 

in Tome VII, fascicule IIIA of the Traité de Zoologie 

edited by J. Forest (see especially the review by Hessler 

and Watling, 1999). 

The suggestion that the orders Spelaeogriphacea, 

Cumacea, Tanaidacea, and Thermosbaenacea constitute 

a grouping termed the ‘‘Brachycarida’’ that 

is the sister group to the Isopoda, first suggested by 

Schram (1981) and supported by Watling (1983, 

1999b) [although note that the suggested placement 

of isopods and amphipods differs in these two papers], 

is not followed here. However, removal of the 

thermosbaenaceans from the ‘‘Pancarida’’ and 

grouping them with the other peracarids, which we 

have done, could be seen as supportive of that 

move (see below under order Thermosbaenacea). 

Gutu (1998) and Gutu and Iliffe (1998) have suggested 

a novel reorganization of the peracarids, 

where both the spelaeogriphaceans and mictaceans 

would be treated as suborders of a new peracarid 

order, the Cosinzeneacea (Gutu, 1998). The mictacean 

family Hirsutiidae would be removed to the 

new order Bochusacea (Gutu and Iliffe, 1998). We 

have not followed this suggestion. 

Thus, our Peracarida contains the two orders of 

former ‘‘mysids’’ treated as the separate orders Lophogastrida 

and Mysida (as in many earlier treatments 

as well; see below), plus the Thermosbaenacea, 

in addition to the Spelaeogriphacea, Mictacea, 

Amphipoda, Isopoda, Tanaidacea, and Cumacea. 

Additional comments on each group are 

given below. 

ORDER SPELAEOGRIPHACEA 

To date, there are only three known extant species 

of this group, from South America (Brazil), South 

Africa, and Australia (Pires, 1987; Poore and Humphreys, 

1998; see also Shen et al., 1998). Pires 

(1987) suggested that spelaeogriphaceans and mictaceans 

might be sister taxa. A recent cladistic analysis 

stemming from the discovery of a new genus 

and species from the Upper Jurassic of China (Shen 

et al., 1998) indicates that the Spelaeogriphacea 

may be paraphyletic. Although Shen et al. treat the 

Spelaeogriphacea as a suborder under the order 

Hemicaridea Schram, we have not followed that 

suggestion. This may change if fossil taxa are incorporated 

into the next edition of this classification. 

All species are currently considered members 

of a single extant family, the Spelaeogriphidae, and 

the group has been reviewed recently by Boxshall 

(1999). Gutu (1998) has suggested recently that 

spelaeogriphaceans and some former mictaceans 

(the family Mictocarididae, not the Hirsutiidae) 

should be treated as suborders within the newly 

created order Cosinzeneacea. We have not followed 

this suggestion, as most other workers seem to be 

in agreement that the two groups are deserving of 

separate status within the Peracarida. 

ORDER THERMOSBAENACEA 

The former order Pancarida (as used in Bowman 

and Abele, 1982), erected to accommodate the order 

Thermosbaenacea, has been eliminated in light 

of suggestions that thermosbaenaceans are members 

of a redefined Peracarida clade (see discussion 

in Wagner, 1994; see also Monod and Cals, 1988; 

Cals and Monod, 1988; Spears and Abele, 1998; 

Richter and Scholtz, in press; and above under Per- 


acarida). Our treatment of the Thermosbaenacea as 

true peracarids is in agreement with morphological 

interpretations (e.g., Monod, 1984; Cals and Monod, 

1988; Monod and Cals, 1988, 1999) and recent 

molecular evidence (Spears and Abele, 1998). 

Other workers (e.g., Newman, 1983; Sieg, 1983a, 

b; Pires, 1987; A. Brandt, pers. comm.) have argued 

for maintaining separate status from the other peracarid 

groups (reviewed by Wagner, 1994). Wagner 

(1994), whose extensive review we followed in the 

current classification, also was of the opinion that 

there is no real justification for excluding the Thermosbaenacea 

from the Peracarida. 

Within the Thermosbaenacea, two new families 

have been described since 1982: Halosbaenidae 

(Monod and Cals, 1988) and Tulumellidae (Wagner, 

1994). The family Monodellidae was also recognized 

by Wagner (1994), bringing the total to 

four recognized extant families (up from one in 

Bowman and Abele, 1982). Wagner’s (1994) thorough 

treatment also suggests some phylogenetic relationships 

among the thermosbaenaceans (as did 

Monod and Cals, 1988). The Thermosbaenidae 

and Monodellidae appear to be sister taxa, but the 

position of the Tulumellidae was undetermined, 

sometimes appearing as the sister group to the Halosbaenidae 

and sometimes as part of the thermosbaenid 

� monodellid clade (as in his ‘‘final proposed 

phylogenetic tree’’; Wagner, 1994, fig. 498). 

Thus, we have not attempted to phyletically order 

the four recognized families at this time. See also 

the recent review by Monod and Cals (1999), 

where previous systematic arrangements (Cals and 

Monod, 1988; Wagner, 1994) are briefly discussed. 

ORDERS LOPHOGASTRIDA AND MYSIDA 

Abele and Spears (1997) concluded, based on 

rDNA studies, that the Peracarida (including the 

Thermosbaenacea) is indeed a monophyletic assemblage, 

but only if the Mysida are excluded. Jarman 

et al. (2000) also would separate the Mysida, which 

they felt are closer to the Decapoda, from the Lophogastrida. 

Supporting evidence is also found in 

the fact that all peracarids (again including thermosbaenaceans 

but excluding Mysida) contain similar 

hypervariable regions of 18S rDNA (Spears and 

Abele, 1998). However, these distinctly peracarid 

features appear to be present in the other mysidacean 

group, the Lophogastrida. The inclusion of the 

mysids (both Mysida and Lophogastrida) in the 

Peracarida (e.g., as suggested most recently by 

Richter and Scholtz, in press) has also been questioned 

on morphological grounds. For example, as 

noted above, Watling (1998, 1999a, b) feels that 

the mysidaceans (i.e., both the Mysida and Lophogastrida 

as the taxon Mysidacea) do not belong to 

the Peracarida and are instead more closely allied 

to the eucarids. Yet both groups of the Mysidacea 

(Mysida and Lophogastrida) share some unique 

and possibly synapomorphic morphological features 

of the walking limbs (Hessler, 1982; see also 

Hessler, 1985) and foregut (De Jong-Moreau and 

Casanova, 2001) that suggest monophyly. Additionally, 

Richter (1999; see also Richter and 

Scholtz, in press) has shown that lophogastridans 

and mysidans share unique morphological components 

to the design of their ommatidia (although 

these features also are shared with Anaspidacea and 

Euphausiacea). The recent treatment by Nouvel et 

al. (1999) treats the Mysidacea as monophyletic 

(see also Richter, 1994, for further arguments in 

favor of monophyly of the Mysidacea). 

Are mysidaceans paraphyletic? Is it possible that 

the Mysida fall outside the Peracarida sensu stricta 

but that the Lophogastrida are true peracarids (ignoring, 

for the moment, the larger question of 

whether the Peracarida itself is monophyletic)? This 

seems unlikely based on limb morphology (e.g., 

Hessler, 1982), and foregut morphology (De Jong- 

Moreau and Casanova, 2001), and yet other workers 

have noted significant differences between the 

Mysida and Lophogastrida on morphological (and 

now, it appears, on molecular) grounds. Several 

other workers (e.g., G. Scholtz and S. Richter, pers. 

comm.) commented on the distinct morphological 

differences between the Lophogastrida and Mysida 

and suggested that these taxa be elevated to ordinal 

status and that the former Mysidacea that contained 

the two be abandoned (but see also Richter, 

1994, De Jong-Moreau and Casanova, 2001, and 

Richter and Scholtz, in press, for arguments in favor 

of monophyly). We have split the former order 

Mysidacea, elevating each of the former mysid suborders 

to order level, as have several other workers 

before us, such as Schram (1984, 1986), and Brusca 

and Brusca (1990:624, who note that an increasing 

number of specialists have begun to treat the two 

groups separately). This could be seen as a preliminary 

for removing one or both of these groups 

from the Peracarida, if the suggestions of Watling 

(1998, 1999a, b) and Spears and Abele (1998) find 

additional support in the future. However, we have 

kept the two groups within the Peracarida for now. 

Taylor et al. (1998) analyzed the relationships of 

a group of fossil malacostracans (the Pygocephalomorpha) 

that are possibly allied with mysids; one 

of their conclusions was that the recent mysids and 

lophogastrids do form a clade (albeit a somewhat 

‘‘confused’’ one). Thus, our classification is most 

similar to that of Brusca and Brusca (1990) in recognizing 

both former ‘‘mysidacean’’ groups as orders 

within the superorder Peracarida rather than 

as suborders within the Mysidacea (as presented by 

Nouvel et al., 1999). Casanova et al. (1998) examined 

relationships of the two lophogastrid families 

(Eucopiidae and Lophogastridae) based on 

morphological and limited molecular data. Among 

their conclusions was that the monogeneric eucopiids 

(Eucopia) originated from within the Lophogastridae. 

Authorities and dates for some taxa in the Mysida 

have been changed to earlier workers and dates 

(e.g., Mysida Haworth and Mysidae Haworth rath- 


er than Mysida Boas or Mysida Dana) following 

the recommendation of L. Holthuis (pers. comm.) 

citing ICZN article 50(c)(i) (now 50.3.1, ICZN 

fourth edition, 1999). Tchindonova (1981) suggested 

the erection within the Mysida of the suborders 

Petalopthalmina and Stygiomysina as well as 

the tribe Amblyopsini and the family Boreomysidae 

(in addition to several new subfamilies, tribes, and 

genera; P. Chevaldonne, pers. comm.). We have not 

followed this suggestion. 

ORDER MICTACEA 

In 1985, two groups of workers simultaneously described 

two new families of an entirely new order 

of peracarid crustaceans and then jointly described 

the new order (Bowman et al., 1985). The new 

families were the Hirsutiidae (Sanders et al., 1985) 

and the Mictocarididae (Bowman and Iliffe, 1985), 

the latter of which formed the basis of the name of 

the new order Mictacea. A second species of the 

Hirsutiidae was described from Australia by Just 

and Poore (1988). Although discovery of the Mictacea 

has prompted speculation about its phylogenetic 

affinities, most workers are in agreement that 

the group fits comfortably within the Peracarida. 

Thus, we include the order and its two families 

among the Peracarida, as does the most recent 

treatment (Hessler, 1999) of the order. Gutu and 

Iliffe (1998) described a new (third) species of hirsutiid 

from anchialine and submarine caves in the 

Bahamas and suggested that the family be removed 

to a new order, the Bochusacea (separate order status 

for the hirsutiids had been suggested also by 

Sanders et al., 1985). The other family of Mictacea 

(Mictocarididae) was then proposed by Gutu 

(1998) to belong to a new order, Cosinzeneacea, 

which would include as suborders the Spelaeogriphacea 

and Mictacea. We have not followed the 

suggestions of Gutu and Iliffe (1998) and Gutu 

(1998). 

ORDER AMPHIPODA 

The Amphipoda, despite a large number of dedicated 

workers and numerous proposed phylogenies 

and classificatory schemes, remain to a large extent 

an unresolved mess. Families proposed by one 

worker often are not recognized by another, and 

disparate classifications based on poorly defined 

features seem to be the rule. The Gammaridea, containing 

the vast majority of amphipod families, is 

the most confusing suborder, although several 

workers (e.g., Kim and Kim, 1993) have proposed 

cladistically based rearrangements of the taxa. We 

should comment especially on the ‘‘semi-phyletic 

classification’’ put forth by Bousfield and Shih 

(1994) in the journal Amphipacifica. This classification 

apparently is being used as the basis for amphipod 

classification in an upcoming publication 

on common names of North American invertebrates 

overseen by the American Fisheries Society 

(although ‘‘minor changes may yet be made’’; E. 

Bousfield, pers. comm., March, 1999). Consequently, 

the Bousfield and Shih (1994) classification or 

its successor in the AFS publication (see Bousfield, 

2001) is likely to be cited often in the years to 

come. Although the Bousfield and Shih (1994) 

work is of value in reviewing previous classificatory 

attempts in recent years, we have not adopted it 

here. The classification divides the group into the 

Amphipoda ‘‘Natantia’’ and Amphipoda ‘‘Reptantia,’’ 

without assigning taxonomic rank to these divisions, 

and then lists the amphipod families under 

superfamily headings. Unfortunately, no authors or 

dates are provided for any of the higher taxa. A 

further point of frustration is that the authors include 

in that paper several different phylogenetic 

hypotheses based on different morphological features; 

however, the phylogenies are not concordant, 

so it is difficult to determine the characters on 

which they base their resulting ‘‘semi-phyletic’’ classification. 

These disparaging comments should not 

be taken as reflecting adversely on other papers 

from these authors. And indeed, a large number of 

papers in which various gammaridean amphipod 

superfamilies and families are revised have been authored 

by Bousfield and his colleagues in recent 

years and should be consulted by workers interested 

in those families. These works include Jarett and 

Bousfield (1994a, b, superfamily Phoxocephaloidea: 

Phoxocephalidae), Bousfield and Hendrycks 

(1994, superfamily Leucothoidea: Pleustidae; 1997, 

superfamily Eusiroidea: Calliopidae), Bousfield and 

Kendall (1994, superfamily Dexaminoidea: Atylidae, 

Dexaminidae), Bousfield and Hoover (1995, 

superfamily Pontoporeioidea: Haustoriidae), Bousfield 

and Hendrycks (1997, superfamily Eusiroidea: 

Calliopiidae), and Bousfield and Hoover (1997, superfamily 

Corophioidea: Corophiidae), and other 

papers in the journal Amphipacifica. 

Following the Fourth International Crustacean 

Congress in Amsterdam, there was a meeting of 

amphipod specialists in Kronenburg, Germany (the 

IXth International Meeting on Amphipoda, July, 

1998). One topic discussed in Kronenburg was 

‘‘Whither amphipod family-level taxonomy?’’ The 

report stemming from that discussion (Vader et al., 

1998) is interesting and informative, and we quote 

from it here: 

Currently the classification of the Amphipoda is still in 

a state of flux; the schedules of Jerry Barnard and Ed 

Bousfield, often not very compatible and neither of 

them based on cladistic analyses, are still prevalent. 

Discussions revolved around the bush-like evolution of 

the Amphipoda and envious comparisons to the Isopoda 

where the general classification appears clearer. 

Not unexpectedly, the classification problems of the 

Amphipoda were not solved! However, it was suggested 

that a cladistic analysis of the amphipod families should 

have high priority, simply to give a general idea of the 

overall relationships, and to generate topics for further 

studies. 

To summarize, in the words of Les Watling (pers. 

comm.), ‘‘most of us working in the amphipod 


world would rather that the [gammaridean] families 

be listed alphabetically rather than by superfamilies.’’ 

Thus, somewhat to our disappointment, we have 

followed that group’s suggestion and also the work 

of Barnard and Karaman (1991) (which has been 

followed by several other workers such as De Broyer 

and Jazdzewski, 1993) in listing alphabetically 

the many families of gammaridean amphipods in 

the current classification. This was done in the 

Bowman and Abele classification as well. The most 

recent treatment, an indispensable review by Bellan-Santini 

(1999), also lists the families of gammaridean 

amphipods (67 of them) alphabetically 

(in addition to listing another 24 families of questionable 

standing) without using superfamilies. 

This work (Bellan-Santini, 1999) differs from our 

compilation slightly and should be consulted by 

any serious student of gammaridean amphipods. 

The alphabetical list of families presented here 

has the advantage of not espousing one worker’s 

view over another (although because Barnard and 

Karaman, 1991, also listed families alphabetically, 

it could be argued that we are preferring their approach; 

E. Bousfield, pers. comm.). It has the additional 

advantage of signaling to future workers 

that the gammarideans are in serious need of further 

attention. However, our alphabetical listing 

has the clear disadvantage of discarding some 

groupings (e.g., corophioids, talitroids, lysianassoids) 

that seem to be fairly well accepted. An additional 

problem that should be noted is that, while 

we are avoiding superfamilies because they are controversial 

and/or not widely used, the same could 

be said for a large percentage of the families that 

we have chosen to recognize. 

Works appearing subsequent to the Bowman and 

Abele (1982) classification that employ these superfamily 

groupings (although not all in perfect 

agreement as to the constituent families) of the 

gammarideans include Schram (1986), Ishimaru 

(1994), Bousfield (1983), and Bousfield and Shih 

(1994). These papers should be consulted for further 

information on gammaridean superfamily hypotheses. 

Further advances in our understanding of 

amphipod phylogeny were presented as part of the 

10th Colloquium on Amphipoda (Heraklion, Crete, 

April, 2000) and include Berge et al. (2000), 

Bousfield (2000a, b), Serejo (2000), and Lowry and 

Myers (2000), abstracts of all of which are available 

via the Amphipod Homepage hosted by Old 

Dominion University in Norfolk, Virginia (URL 

http://www.odu.edu/%7Ejrh100f/amphome). 

SUBORDER GAMMARIDEA 

Gammaridean amphipod families that have been 

described or recognized since the Bowman and 

Abele (1982) list include, in alphabetical order of 

the families, Acanthonotozomellidae (by Coleman 

and Barnard, 1991), Amathillopsidae (recognized 

by Coleman and Barnard, 1991, credited to Pirlot, 

1934, but considered only a subfamily of the Epimeriidae 

by Lowry and Myers, 2000), Allocrangonyctidae 

(by Holsinger, 1989), Aristiidae (by 

Lowry and Stoddart, 1997), Bolttsiidae, Cardenioidae, 

Clarenciidae (all by Barnard and Karaman, 

1987), Cheidae (by Thurston, 1982), Condukiidae 

(by Barnard and Drummond, 1982), Cyphocarididae 

(by Lowry and Stoddart, 1997), Dikwidae (by 

Coleman and Barnard, 1991, suggested to be only 

a tribe within the subfamily Amathillopsinae by 

Lowry and Myers, 2000), Didymocheliidae (by Bellan-Santini 

and Ledoyer, 1986), Endevouridae (by 

Lowry and Stoddart, 1997), Ipanemidae and Megaluropidae 

(by Barnard and Thomas, 1988), Metacrangonyctidae 

(by Boutin and Missouli, 1988), 

Micruropidae (by Kamaltynov, 1999), Odiidae (by 

Coleman and Barnard, 1991, but see Berge et al., 

1998, 1999, who believe that the Odiidae is paraphyletic 

and that its genera belong instead within 

the Ochlesidae), Opisidae (by Lowry and Stoddart, 

1995), Pachyschesidae (by Kamaltynov, 1999), Paracalliopiidae 

(by Barnard and Karaman, 1982), 

Paracrangonyctidae (by Bousfield, 1982), Paraleptamphopidae 

(by Bousfield, 1983), Perthiidae (by 

Williams and Barnard, 1988), Phoxocephalopsidae 

(by Barnard and Clark, 1984, who credit Barnard 

and Drummond, 1982), Phreatogammaridae (by 

Bousfield, 1982), Pseudamphilochidae Schellenberg 

(revised and reinserted by Barnard and Karaman, 

1982), Podoprionidae (by Lowry and Stoddart, 

1996), Pseudocrangonyctidae (by Holsinger, 1989), 

Scopelocheiridae (by Lowry and Stoddart, 1997), 

Sinurothoidae (by Ren, 1999), Sternophysingidae 

(by Holsinger, 1992), Urohaustoriidae (by Barnard 

and Drummond, 1982), Valettidae (by Thurston, 

1989), Wandinidae (by Lowry and Stoddart, 1990), 

and Zobrachoidae (by Barnard and Drummond, 

1982). Additionally, we include the Podoceridae 

Leach, as this appears to be a widely recognized 

and relatively uncontroversial family (e.g., in Barnard 

and Karaman, 1991, and Bellan-Santini, 

1999), although it was not listed by Bowman and 

Abele (1982). Iphimedioid amphipods, like many 

other groupings, are currently being revised, and as 

a result, some of the names and ranks above will 

undoubtedly change (see Lowry and Myers, 2000). 

The family Lepechinellidae Schelenberg, listed in 

Bowman and Abele (1982), has been removed. Barnard 

and Karaman (1991) listed the genus Lepichenella 

in the Dexaminidae and considered the 

Lepichenellidae a synonym of the Dexaminidae 

(but note that Bousfield and Kendall, 1994, treated 

the Lepichinellidae as a subfamily of the Atylidae). 

The family Conicostomatidae is listed in the Zoological 

Record (1983, vol. 20, section 10), where it 

is attributed to Lowry and Stoddart (1983). However, 

although those authors recognized it as a 

grouping of related taxa, they did not establish it 

as a family in their 1983 paper, and they have not 

done so subsequently (J. Lowry, pers. comm.). 

Thus, the listing of the family in the Zoological Record 

is in error. The family Anamixidae is main- 


tained in our classification, although there is reason 

to believe that this family was erected to accommodate 

what are turning out to be highly derived 

males of some species of the Leucothoidae (J. Lowry, 

pers. comm.). If true, the Anamixidae will have 

to be synonymized at some point. A few workers 

asked us to ‘‘correct’’ the spelling of the family 

name Liljeborgiidae to Lilljeborgiidae to reflect the 

fact that the family name honors William Lilljeborg 

(1816–1908). The confusion stems from the fact 

that Vilhelm Liljeborg changed the spelling of his 

name to William Lilljeborg sometime in the early 

1860s. When Bate (1862) established the genus Liljeborgia, 

he used the then-correct spelling honoring 

Vilhelm Liljeborg. Thus, when Stebbing in 1899 established 

the family Lilejborgiidae based on the genus 

Liljeborgia, he was obliged to use this spelling 

as well even though, by that time, the man was 

known as William Lilljeborg (J. Lowry, pers. 

comm., and see Vader, 1972). (As an aside, the 

spelling of the genus Lilljeborgiella, erected by 

Schellenberg in 1931, is therefore also correct, as 

by that time the name was William Lilljeborg). 

All of the 67 families that Bellan-Santini (1999) 

lists as those that ‘‘ne présent pas actuellement de 

problème majeur d’interprétation’’ are included in 

our list. Bellan-Santini (1999) also lists another 24 

families that do present problems, and some of 

those are in our list as well. Some of the names and 

dates attributed to some families differ between our 

list and hers as well. 

SUBORDER CAPRELLIDEA 

Takeuchi (1993) indicated that the Caprellidea may 

not be monophyletic but stopped short of proposing 

a new classification of the group. His results 

(Takeuchi, 1993, figs. 1, 5) indicated that the phtisicids 

are the sister group to all other caprellideans 

and that the paracercopids are more closely related 

to the caprellid-caprogammarid line (he did not 

deal with the parasitic family Cyamidae). Thus, we 

have removed the family Paracercopidae from the 

superfamily Phtisicoidea and have placed it instead 

in the superfamily Caprelloidea, leaving the Phtisicidae 

the sole family of the Phtisicoidea. We saw 

this move as preferable to creating yet another superfamily 

(to contain the paracercopids) in an already 

taxon-dense suborder. In the same year and 

in the same volume, Laubitz (1993) described two 

new caprellidean families (Caprellinoididae and 

Pariambidae). She also recognized as valid the Protellidae 

McCain and tentatively suggested some 

evolutionary lines or trends within and leading up 

to the Caprellidea. Some of these ideas differ from 

those proposed by Takeuchi (1993), although both 

workers recognize the same eight families (as does 

Bellan-Santini, 1999). Also in that same volume, 

Kim and Kim (1993) suggested affinities between 

caprellideans and corophioids. Margolis et al. 

(2000) have suggested that the Cyamidae may be 

closer to the Caprogammaridae-Caprellidae lineage 

rather than to the Caprellinoididae-Phtiscidae line, 

as suggested by Laubitz (1993). Several names and 

dates have reverted to earlier workers (suggestions 

of L. Holthuis, pers. comm.). The families Aeginellidae 

and Dodecadidae have been deleted, as they 

are now considered subfamilies of the Caprellidae 

and Phtisicidae (K. Larsen, pers. comm.; Laubitz, 

1993). See also Bellan-Santini (1999). 

SUBORDER HYPERIIDEA 

Workers familiar with hyperiideans may wonder 

why we did not follow the revision of the Hyperiidea 

by Vinogradov et al. (1982, with English translation 

edited by D. Siegel-Causey appearing in 

1996). While that work contains much updated information 

concerning the biology of hyperiideans 

and nomenclatural changes below the level of family, 

the authors followed, for the higher classification, 

the earlier work by Bowman and Gruner 

(1973). Thus, the Bowman and Abele (1982) classification 

is the more current of the two for higher 

level taxa, although workers will want to consult 

the Vinogradov et al. volume for information within 

families and genera (D. Causey, pers. comm.). 

Our classification is also consistent with the classifications 

of Schram (1986, which in turn was 

based largely on Bousfield, 1983) and Bellan-Santini 

(1999). Kim and Kim (1993) suggested that hyperiids 

may be related to certain leucothoid members 

(Amphilochidae and Stenothoidae) of the 

Gammaridea. 

SUBORDER INGOLFIELLIDEA 

Several workers (e.g., J. Holsinger, pers. comm.) 

have pointed out that the ingolfiellids and metaingolfiellids 

may not justify their own suborder and 

could probably be accommodated within the Gammaridea. 

Indeed, Bowman and Abele (1982) listed 

them alphabetically among the other gammaridean 

families. However, Holsinger notes at the same time 

that this view is not universally shared by other 

amphipod workers, and most workers (e.g., Bellan- 

Santini, 1999) continue to treat these two families 

as the sole members of the suborder Ingolfiellidea. 

Vonk and Schram (1998) argue for maintaining 

separate status for the group. We have retained 

their separate status pending further investigations 

into the group’s affinities. 

ORDER ISOPODA 

The diversity of and fascination with isopods are 

reflected in the relatively large number of carcinologists 

currently working on isopod systematics and 

phylogeny. Although it is encouraging to see so 

many skilled workers dedicated to resolving questions 

of isopod systematics, there are negative aspects, 

one of which is the relatively large number 

of responses we received that contained conflicting 

ideas or information. For the most part, we have 

relied on the rather straightforward list of the ma- 


ine isopods that has been posted on the World 

Wide Web by B. Kensley and M. Schotte (http:// 

www.nmnh.si.edu/iz/isopod). However, in that 

compilation, the various suborders and their constituent 

superfamilies and families are arranged alphabetically. 

Brusca and Wilson (1991), while proposing 

some phylogenetic changes that would seriously 

alter the arrangement of groups as presented 

here (and at the same time countering several of 

the hypotheses forwarded earlier by Wägele, 1989), 

stopped short of proposing a new classification 

based on their hypothesis. Their feeling was that 

insufficient evidence had been amassed for proposing 

classifications based on the phylogenetic hypotheses 

they were presenting as testable ideas. The 

Brusca and Wilson (1991) analysis was criticized 

by Wägele (1994), who in fact used their paper to 

point out potential pitfalls in any attempt at computer-generated 

cladistic analyses. Wägele (1994) 

was in turn rebutted by Wilson (1996), who was 

answered by Wägele (1996), and it would seem 

that we have a long way to go before any consensus 

concerning isopod phylogeny (not to mention phylogenetic 

method) is reached. Thus, our classification 

is in some ways a step backward in that we 

continue to recognize some groups, such as the Flabellifera, 

that appear clearly paraphyletic (following 

the analyses of both Brusca and Wilson, 1991, 

and Wägele, 1989) but for which no alternative 

classifications have been proposed. In the most recent 

overall treatment of isopods, Roman and Dalens 

(1999) continue to recognize the Flabellifera as 

well while acknowledging that it is a heterogeneous 

assemblage. 

Additionally, many changes, especially those concerning 

names and dates of the authorities credited 

with establishing families but also concerning 

whether or not to recognize a particular family, 

have been incorporated at the request of some of 

the major workers (e.g., L. Holthuis, B. Kensley, R. 

Brusca, G. Poore, W. Wägele, and G. Wilson) via 

personal communications. It has not always been 

possible for us to verify these suggestions. Often, 

despite a rather large library on crustacean systematics 

at our disposal, we have been unable to see 

the original references. In cases where we received 

conflicting information (such as whether the family 

Arcturidae should be credited to White, 1850 vs. 

Bate and Westwood, 1868 vs. Sars, 1899) and/or 

we could not verify by checking on all of the suggested 

references ourselves, we have chosen the first 

known usage (in this case, using Arcturidae White, 

1850, which turns out to be correct according to 

G. Poore, who owns the book) in accordance with 

ICZN article 50.3.1. One such change involves the 

establishment of a large number of families and superfamilies 

credited to Latreille. L. Holthuis (pers. 

comm.) assures us that 1802 is the correct date for 

the many taxa that have been, in the past, credited 

to Latrielle (1803) (see earlier section on names, 

dates, and the ICZN). 

Major papers suggesting changes in how we or- 

ganize the Isopoda that have appeared subsequent 

to Bowman and Abele (1982) include Wägele 

(1989) and Brusca and Wilson (1991). Poore 

(2001a) presented a phylogeny of the Anthuridea 

suggesting relationships among the six families 

(two new), but to our knowledge, there have not 

as yet been names proposed for the divisions suggested 

by him. The most recent review, by Roman 

and Dalens (1999), recognizes eight suborders. 

Their arrangement differs from ours in that (1) they 

recognize the suborder Gnathiidea, which we do 

not, and (2) they do not recognize the suborders 

Microcerberidea and Calabozoidea, which we do, 

for reasons discussed below. 

Concerning the former suborder Gnathiidea, 

Brusca and Wilson (1991) suggested that the gnathiids 

were derived from among the families traditionally 

thought of as ‘‘flabelliferan’’ isopods (a 

group that they demonstrate is not monophyletic). 

Wägele (1989, pers. comm.) also would remove the 

gnathiids from their own suborder, but his preference 

was to place them among the Cymothoida, a 

group he recognizes as containing a large number 

of former Flabellifera families. We have, for the 

current classification, removed the gnathiids from 

their own superfamily and have placed them within 

the Flabellifera, knowing that the Flabellifera itself 

is not monophyletic and must some day be extensively 

revised. L. Holthuis (pers. comm.) has suggested 

that we credit the family name Gnathiidae 

to Leach (1814) rather than to Harger (1880), as 

was used by Bowman and Abele (1982) and Roman 

and Dalens (1999). 

SUBORDER PHREATOICIDEA 

Wilson (pers. comm.) suggests that many of the 

subfamilies of the Amphisopodidae recognized by 

Nicholls (1943, 1944) will need to be elevated to 

family level (e.g., as Hypsimetopodidae, Mesamphisopodidae, 

Phreatoicopsididae) once this suborder 

is revised (see also Wilson and Johnson, 

1999; Wilson and Keable, 1999, 2001). Our classification 

follows Roman and Dalens (1999) in recognizing 

three families (the same three that appear 

in Bowman and Abele, 1982). By listing the phreatoicids 

first among all isopod suborders, we are acknowledging 

the primitive nature of these isopods. 

Brusca and Wilson (1991) and Wilson and Johnson 

(1999) have indicated that the phreatoicideans, all 

of which are restricted to Gondwanan fresh waters, 

may be ‘‘the earliest derived isopod Crustaca’’ (Wilson 

and Johnson, 1999:264). The phreatoicidean 

fossil record extends back to the Carboniferous 

(Wilson and Johnson, 1999). 

SUBORDER ANTHURIDEA 

Within this suborder, the family Antheluridae was 

described by Poore and Lew Ton (1988) and the 

families Expanathuridae and Leptanthuridae were 

described recently by Poore (2001a; see also Poore, 

1998). Our treatment differs from that of Roman 


and Dalens (1999) in that we include six families. 

Roman and Dalens do not recognize the family Antheluridae 

and of course could not have known 

about the Expanathuridae and Leptanthuridae. 

SUBORDER MICROCERBERIDEA 

Wägele (1983) placed the family Microcerberidae 

within the Aselloidea; Brusca and Wilson (1991) 

considered the Microcerberoidea the sister group to 

the Asellota and consequently suggested they not 

be included among the Asellota. Our treatment of 

the family as belonging to its own suborder and 

superfamily follows Bowman and Abele (1982) but 

is also in keeping with the suggestion of Brusca and 

Wilson (1991). Additionally, we now treat the 

monotypic family Atlantasellidae in this suborder 

on the recommendation of G. D. F. Wilson (pers. 

comm.). 

SUBORDER FLABELLIFERA 

Brusca and Wilson (1991) showed that the Flabellifera 

was a paraphyletic grouping, a finding that 

has been suggested also by other workers. Wägele 

(1989) (rebutted to some degree by Wilson, 1996) 

argued for dividing the flabelliferan families into 

two somewhat smaller groups, the Cymothoida 

and Sphaeromatidea (see Wägele, 1989). Wägele 

would remove from the Flabellifera the family Atlantasellidae 

(which he considers an Aselloidea). 

The families Aegidae, Anuropidae, Argathonidae, 

Cirolanidae, Corallanidae, Cymothoidae, and Tridentellidae 

would belong to his grouping Cymothoida 

Leach, 1814. The remaining families (Bathynataliidae, 

Hadromastacidae, Keuyphyliidae, Limnoriidae, 

Phoratopodidae, Plakarthriidae, Serolidae, 

Sphaeromatidae, and Tecticepitidae) he would 

place in the Sphaeromatoidea. Thus, the two most 

current and most ambitious schemes of isopod phylogeny, 

although agreeing in some respects, do not 

agree even closely on how to treat the former flabelliferan 

families (see also Brandt et al., 1999, for 

a comparison of phylogenetic hypotheses of sphaeromatoid 

families in light of the fossil family 

Schweglerellidae). Roman and Dalens (1999) recognize 

the Flabellifera, and divide it into three superfamilies: 

Cirolanoidea (seven families), Sphaeromatoidea 

(two families), and Seroloidea (two 

families). We have retained the Flabellifera for the 

current classification, knowing that this assemblage 

cannot be considered monophyletic, and for now, 

we have avoided the use of superfamilies. Recent 

fossil finds (see Brandt et al., 1999) have pushed 

back the origin of some former flabelliferan isopods, 

indicating that the sphaeromatoid isopods, at 

least, are of Late Jurassic ancestry or older. 

Within the Flabellifera, the following changes 

have been incorporated (listed alphabetically by 

family): Ancinidae (elevated to family status by N. 

L. Bruce, 1993), Argathonidae (removed per R. 

Brusca, pers. comm.), Bathynomidae (removed per 

B. Kensley, pers. comm.), Excorallanidae (removed 

per B. Kensley, pers. comm.), Hadromastacidae (described 

by Bruce and Müller, 1991), Lynseiidae (described 

by Poore, 1987; removed per Cookson and 

Poore, 1994; see also Bruce, 1988), Protognathiidae 

(described by Wägele and Brandt, 1988; moved 

from Gnathiidea per R. Brusca and also G. Wilson, 

pers. comm.), Tecticepitidae (originally described as 

a subfamily by Iverson, 1982; elevated to family 

status by N. L. Bruce, 1993), and Tridentellidae 

(described by Bruce, 1984). 

N. L. Bruce (1993) presented a key to the known 

flabelliferan families, reappraised the family Sphaeromatidae 

Latreille (a family in rather dire need of 

internal revision; see Harrison and Ellis, 1991), and 

recognized as families the Ancinidae Dana and Tecticipitidae 

Iverson. 

G. Poore (pers. comm.) informs us that the Aegidae 

is correctly attributed to White (1850) rather 

than to Leach (there are no families mentioned in 

the only paper that Leach published in 1815, the 

date given in Bowman and Abele for this family). 

He also informs us that the families Ancinidae, Cirolanidae, 

and Serolidae are correctly attributed to 

Dana (1852) instead of 1853 (as in Bowman and 

Abele, 1982). 

Bowman and Abele (1982) used the spelling Anuropodidae 

for this isopod family, while noting 

(1982: 21) that the tanaid family Anuropodidae Băcescu 

was a homonym of the isopod family Anuropodidae 

Stebbing. ICZN Opinion 1357 (ICZN, 

1985b) dictated that the spelling of the isopod family 

should be Anuropidae to remove the homonymy, 

and thus we use Anuropidae as the correct 

spelling of this isopod family. 

The Plakarthriidae Hansen is, according to G. 

Poore (pers. comm.), ‘‘an effective replacement 

name for Chelonidiidae Pfeffer, 1887, but is conserved 

under ICZN article 40’’; Dr. Poore suggests 

that the date 1887 should follow Hansen, 1905, in 

parentheses, as Plakarthriidae Hansen, 1905 

(1887). 

SUBORDER ASELLOTA 

According to G. Wilson and G. Poore (pers. 

comm.), the currently recognized superfamilies of 

the Asellota are either poly- or paraphyletic (see 

also Wilson, 1987) and will not stand the test of 

time. Roman and Dalens (1999) treat the Asellota 

as being comprised of four superfamilies (down one 

from Bowman and Abele, 1982; the Protallocoxoidea 

and its single family, Protallocoxidae, have 

been removed). We have followed this arrangement 

here, recognizing the superfamilies Aselloidea, Stenetrioidea, 

Janiroidea, and Gnathostenetroidea. 

The superfamily Pseudojaniroidea, proposed by 

Wilson (1986), has been removed at his suggestion 

(G. Wilson, pers. comm.; see also Serov and Wilson, 

1999). Its former family, the Pseudojaniridae, 

has been transferred to the Stenetrioidea following 

the revision of the Pseudojaniridae by Serov and 

Wilson (1999). 


In the superfamily Aselloidea, the family Atlantasellidae 

has been removed. Brusca and Wilson 

(1991) suggested its removal to the Microcerberoidea, 

where we have placed it. Although Roman and 

Dalens (1999) treat the family Microcerberidae as 

a member of the Aselloidea, we are keeping it in its 

own suborder (Microcerberidea) and superfamily 

(Microcerberoidea) as per Bowman and Abele 

(1982) (as noted earlier). Thus, the Aselloidea presently 

contains only the Asellidae and Stenasellidae. 

The superfamily Stenetrioidea now contains the 

Pseudojaniridae (as noted above), although Roman 

and Dalens (1999) have kept it at one family, the 

Stenetriidae. 

Within the enormous superfamily Janiroidea, the 

Abyssianiridae was removed (incorporated into the 

Paramunnidae) following Just (1990). Species formerly 

within that family are now considered to belong 

to the Paramunnidae. The former families Eurycopidae, 

Ilyarachnidae, and Munnopsididae are 

now considered subfamilies of the Munnopsididae 

(Wilson, 1989). The Microparasellidae is apparently 

polyphyletic; ‘‘some taxa may be moved to the 

Vermectiadidae or put in a new family; Microparasellus 

will stay in the Janiroidea’’ (Wilson, pers. 

comm.). The Janiridae was shown to be nonmonophyletic 

by Wilson (1994) but remains a valid 

family; some of its genera will eventually be reassigned 

to other families. The Katianiridae was described 

by Svavarsson (1987). Although the family 

Pleurogoniidae is recognized by some workers (e.g., 

Roman and Dalens, 1999), we have removed it at 

the suggestion that it is a junior synonym of the 

Paramunnidae (G. Poore, pers. comm.; G. Wilson, 

pers. comm.). The family Pseudomesidae was sunk 

into the Desmosomatidae by Svavarsson (1984). 

Although the family Santiidae is credited to Kussakin 

(1988) by many workers (e.g., Wolff, 1989), 

it was first used (in a figure) by Wilson (1987). In 

Wilson’s (1987) paper, he acknowledges Fresi et al. 

(1980) as the source for one of the phylogenetic 

trees in that paper (Wilson’s fig. 5B). However, Fresi 

et al. (1980) did not include the Santiidae in their 

figure; it was apparently added (and therefore first 

used) by Wilson (1987). Thus, we have credited the 

family Santiidae to Wilson. Cohen (1998), in his 

review of the family Dendrotiidae, explains why 

this spelling of the family name is preferred over 

Dendrotionidae (used by Lincoln and Boxshall, 

1983). Interested workers should also consult Roman 

and Dalens (1999), whose list of families differs 

from ours in several respects. 

The superfamily Protallocoxoidea and family 

Protallocoxidae were removed per G. Wilson (pers. 

comm.). 

The superfamily Gnathostenetroidoidea contains 

the families Gnathostenetroididae and Protojaniridae 

(following Roman and Dalens, 1999). Additionally, 

the interesting family Vermectiadidae was 

described by Just and Poore (1992), and our tentative 

inclusion of the vermectiadids in the super- 

family Gnathostenetroidoidea is based mostly on 

the recommendation of R. Brusca (pers. comm.). 

SUBORDER CALABAZOIDA 

This family (Calabozoidae) and its suborder were 

erected by Van Lieshout (1983). Brusca and Wilson 

(1991) suggest that the calabazoids are oniscideans 

and so they should probably be moved, but we 

have not done so in this classification. Wägele 

(pers. comm.) points out that the ending -oidea 

should be reserved for superfamilies and suggested 

that we change the spelling of the suborder to Calabazoida, 

which we have done. 

SUBORDER VALVIFERA 

Within the Valvifera, several families have been 

added since the Bowman and Abele (1982) classification. 

The family Austrarcturellidae was described 

by Poore and Bardsley (1992), and the families 

Antarcturidae, Arcturididae, and Rectarcturidae 

were added by Poore (2001b). Poore (2001b) 

also recognized the Holidoteidae, crediting it to 

Wägele (1989), who first suggested it as a subfamily. 

Current research shows that the family Amesopodidae 

is probably a junior synonym of the Arcturidae 

(G. Poore, G. Wilson, pers. comm.), and so 

we have removed it, although the family was listed 

by Roman and Dalens (1999), who did not list the 

Austrarcturellidae. Thus, we recognize 11 families, 

4 more than did Bowman and Abele (1982). The 

family Arcturidae, credited by Bowman and Abele 

(1982) to Sars, is correctly credited to Dana (1849), 

and the family Idoteidae is correctly attributed to 

Samouelle (G. Poore, pers. comm.). 

SUBORDER EPICARIDEA 

Wägele (1989, pers. comm.) suggested that all of 

the epicaridean families we have listed should be 

treated as families or subfamilies of the Cymothoida 

Leach (see above). We have not made this rather 

radical change and instead have followed the more 

conservative classification given by Trilles (1999). 

Trilles (1999) divides the epicaridean families into 

two sections, Bopyrina and Cryptoniscina, which 

we have treated as superfamilies (Bopyroidea and 

Cryptoniscoidea) to allow a more consistent spelling 

and in keeping with our treatments of other 

peracarid groups. In the Bopyroidea are the three 

families Bopyridae, Dajidae, and Entoniscidae (all 

of which were listed by Bowman and Abele, 1982). 

In the section (now superfamily) Cryptoniscoidea, 

Trilles (1999) treats an additional eight families not 

listed by Bowman and Abele (1982); the family Liriopsidae 

has been deleted (see arguments in Grygier 

and Bowman, 1990, 1991; Trilles, 1999). Thus, 11 

epicaridean families are recognized. The families 

added since Bowman and Abele (1982) are not 

newly described families but instead represent recognition 

of formerly described families that were 

treated in the past, at least by some authors, as 


subfamilies of the Cryptoniscidae, for which Bowman 

and Abele (1982), followed by Schram (1986), 

used the name Liriopsidae (see Grygier and Bowman, 

1990). Crediting authorship of the family 

Cryptoniscidae (and thus Cryptoniscoidea) to Kossman 

rather than to Gerstaecker is based on the correction 

published by Grygier and Bowman (1991). 

Following Trilles (1999), we also do not recognize 

the family Microniscidae Müller for the genus Microniscus, 

although this family is still listed in some 

compendia (e.g., by Brasil-Lima, 1998:641, in 

Young, 1998). The spelling Cabiropsidae used by 

Trilles (1999) and some earlier workers is corrected 

to Cabiropidae based on the explanation given by 

Sassaman (1992). 

SUBORDER ONISCIDEA 

The relationships of the terrestrial isopod groups to 

one another and to marine relatives are still poorly 

understood. Although Schmalfuss (1989, in Ferrara, 

1989) proposed some relationships among oniscideans 

and compared the classification of oniscideans 

presented by Holdich et al. (1984) with a 

new one based on his analysis, Schmalfuss’ work 

was based on relatively few characters and was criticized 

by Brusca (1990). Wägele (pers. comm.) informs 

us that there are ‘‘enormous advances that 

will be published next year’’ concerning the phylogeny 

of the Oniscidea and that several groups 

presented here are not monophyletic; further, he informs 

us that the ‘‘section’’ Diplochaeta is currently 

being revised. Until these advances become known 

to us, we are unsure as to what relationships our 

classification should suggest. Holdich et al. (1984) 

used two infraorders (the Tylidae were placed in a 

separate infraorder, Tylomorpha), and within the 

infraorder Ligiamorpha they recognized three sections. 

Schmalfuss (1989) did not employ the infraorder 

level and instead divided all oniscideans 

among four major sections. More recent arrangements 

of the oniscidean families have been proposed 

by Erhard (1995) and Tabacaru and Danielopol 

(1996a, b; see also Roman and Dalens, 1999, 

who followed mostly Schmalfuss, 1989, and also 

Mattern and Schlegel, 2001). Many workers (e.g., 

Souza-Kury, 1998, in Young, 1998) list the oniscidean 

families alphabetically. 

We have maintained the two-infraorder system 

and have not recognized the new section Microchaeta 

proposed by Schmalfuss. The four families 

Helelidae, Irmaosidae, Pseudarmadillidae, and 

Scleropactidae have been removed from any infraorder 

or superfamily, as their status is indeterminate 

(R. Brusca, pers. comm.). For the currently 

accepted family names (as well as authors and 

dates, which were not included by Schmalfuss), we 

have had to rely primarily on the alphabetical list 

of oniscidean families maintained on the Smithsonian’s 

server (Kensley et al., 1998; URL http:// 

www/nmnh.si.edu/iz/isopod), which is based on 

Schmalfuss’ families (the terrestrial isopod list is 

also accessible via the Kensley et al. list of marine 

isopods, URL gopher://nmnhgoph.si.edu:70/11/. 

invertebrate/.crustaceans). Users of the terrestrial 

isopod list are strongly cautioned by the authors 

(Kensley et al., 1998): 

This list is thus intended as a rough guide to the astounding 

array of names and taxa in the Oniscidea. 

Synonymy will be rampant in the list. We have tried to 

use the most current interpretations of some genera and 

families. Nevertheless, we realise that in no way do we 

even begin to resolve the taxonomic confusion that 

reigns in this group. There is uncertainty regarding the 

familial placement of some genera, and there will certainly 

be repetition of the same specific name under 

different genera. There are omissions from the list, either 

of names of taxa that we’ve completely missed, or 

of authors and dates of publication and/or of localities 

that we have been unable to find. 

We are aware of only two newly described oniscidean 

families since 1982: Ferrara and Taiti 

(1983) described the family Irmaosidae, and 

Schultz (1995) described the Dubioniscidae (see 

Souza-Kury, in Young, 1998:656). Establishment of 

the family Platyarthridae is credited to Verhoeff 

(rather than to Vandel) by Ferrara and Taiti (1989), 

who also note that the families Bathytropidae and 

the Platyarthridae might coincide. G. Poore (pers. 

comm.) notes that the Styloniscidae Vandel, 1952, 

is a replacement name for the Patagoniscidae Verhoeff, 

1939, and is conserved under ICZN article 

40; he therefore recommends that the earlier date 

appear in parentheses, as Styloniscidae Vandel, 

1952 (1939). Characters that define the various 

groupings of the oniscideans are given by Roman 

and Dalens (1999), although workers should note 

that the characters and groupings based on them 

are, in some cases, not universally accepted. A recent 

molecular analysis (Mattern and Schlegel, 

2001) based on ssu rDNA suggests that Crinochaeta 

and Synochaeta are monophyletic, and that these 

groups together are the sister taxon to the Diplochaeta. 

ORDER TANAIDACEA 

Many of the major taxonomic changes suggested 

by the late J. Sieg were made prior to 1982 and 

were therefore incorporated into the Bowman and 

Abele classification. Subsequent to 1982, there were 

also some large-scale rearrangements suggested by 

Sieg (1983a, b, 1984, 1986a, b), but there has been 

almost no work done at higher levels of tanaid systematics 

since that time. Unfortunately, it now appears 

that many of the characters established or 

used by Sieg do not hold up well under scrutiny 

(see Larsen and Wilson, 1998), and it is not clear 

how many of Sieg’s characters or numerous classificatory 

assignments will survive. Kim Larsen (pers. 

comm.) is actively studying the group and has kindly 

updated us, as far as is possible pending a thorough 

revision of the group. Additionally, he has 

provided us with many suggested changes. An excellent 

and comprehensive web site maintained by 


Richard Heard and Gary Anderson now exists at 

URL http://tidepool.st.usm.edu/tanaids/index.html, 

and our arrangement of the group is the same as 

theirs. 

Authorship of the Tanaidacea is now credited to 

Dana (1849) rather than to Hansen (1895) (L. Holthuis, 

pers. comm.). A review by M. Gutu and the 

late Jürgen Sieg (Gutu and Sieg, 1999) additionally 

includes fossil taxa (most of which were added by 

Schram et al., 1983). The classification in Gutu and 

Sieg (1999) differs from ours in that we include the 

family Tanapseudidae, not listed in Gutu and Sieg 

(1999), and in that we have deleted the Leptognathiidae 

(see below). 

SUBORDER TANAIDOMORPHA 

The naturalness of the entire suborder Tanaidomorpha 

was questioned by Larsen and Wilson 

(1998), who noted that inconsistencies or contradictions 

in descriptions and illustrations of several 

authors ‘‘plague tanaidomorphan taxonomy.’’ Larsen 

and Wilson also noted that several of Sieg’s 

characters and subsequent classifications, which 

form the basis of our current understanding of tanaid 

systematics, have been found wanting. They 

conclude that ‘‘the current taxonomy . . . for the 

suborder Tanaidomorpha, heavily burdened by inconsistencies, 

is not useful at the present stage.’’ It 

seems unlikely that the situation for the other suborders 

would be any better. 

Within the Tanaidomorpha, the family Leptognathiidae 

was abandoned by Sieg (1986b) as it 

was found to be a junior synonym of Anarthruridae 

(Sieg, 1986b; see Larsen and Wilson, 1998). One 

of its constituent subfamilies was incorporated into 

the Anarthruridae Lang, and the other was elevated 

to familial rank (now the Typhlotanaidae Sieg). The 

family Agathotanaidae similarly was downgraded 

from a family to ‘‘tribe’’ status (Sieg, 1986b). Dates 

of establishment of the Nototanaidae and Pseudotanaidae 

(in the past, often credited to Sieg, 1973) 

have been changed from 1973 to 1976, as the 1973 

work is an unpublished thesis that did not appear 

in published form until three years later (Sieg, 

1976) (K. Larsen, pers. comm.). 

Additional tanaidomorphan families described 

subsequent to the Bowman and Abele (1982) list 

include the Pseudozeuxidae and Typhlotanaidae, 

described by Sieg (1982) and Sieg (1986b), respectively. 

SUBORDERS NEOTANAIDOMORPHA AND 

APSEUDOMORPHA 

Sieg (1983b) elevated to family status the Whiteleggiidae 

and placed within it the former family Leviapseudidae 

as a subfamily (Leviapseudinae) of the 

Whiteleggiidae. Sieg (1984) established the family 

Cyclopoapseudidae to accommodate a genus formerly 

in the Metapseudidae (Sieg, 1984; Larsen, 

pers. comm.), but the Cycloapseudidae is now considered 

a junior synonym of the Metapseudidae 

(Larsen, pers. comm.). The Parapseudidae was not 

accepted by Sieg (1986a, b) but has since been recognized 

as valid (see brief discussion in Gutu, 1996; 

K. Larsen, pers. comm.). See Gutu and Sieg (1999) 

for the most recent review. 

ORDER CUMACEA 

Our classification follows the World Wide Web list 

compiled by Watling and Kornfield (URL http:// 

nature.umesci.maine.edu/pub/Cumacea/data.html) 

as part of their National Science Foundation PEET 

training project. Their list is similar to that of Bowman 

and Abele, with two exceptions. First, the family 

Archaeocumatidae Băcescu, 1972, containing 

the single genus Archaeocuma, has been removed. 

Its establishment (in Băcescu, 1972) had been questioned 

earlier by Jones (1976), who felt that further 

confirmation was needed prior to accepting this 

family, and Watling (pers. comm.) informs us that 

this family is generally not recognized. However, 

the family is listed (considered valid) by Băcescu 

(1988) and by Băcescu and Petrescu (1999). Second, 

the family Gynodiastylidae Stebbing, 1912, 

has been included following Day (1980), Băcescu 

(1992), Băcescu and Petrescu (1999), and the 

above-mentioned web site. Thus, the total number 

of cumacean families remains at eight, as with the 

Bowman and Abele list, although the composition 

has changed. Watling (pers. comm.) also notes that 

the family Nannastacidae will very likely be split 

into two families in the near future. 

Relationships within the Cumacea have been tentatively 

suggested recently by Haye and Kornfield 

(1999) on the basis of somewhat limited molecular 

data. Their suggestion is that those families with an 

articulated telson (families Bodotriidae, Leuconidae, 

and Nannastacidea) form a clade that is distinct 

from a second lineage containing the five families 

without an articulated telson. This grouping is 

not reflected in the current classification, where all 

families are instead listed alphabetically. 

SUPERORDER EUCARIDA 

Most workers seem to be in agreement that the Eucarida 

is a valid (i.e., monophyletic) assemblage 

(but see arguments against a monophyletic Eucarida 

in Richter and Scholtz, in press). Schram (1984) 

noted that the eucarids ‘‘are destined for some kind 

of realignment,’’ and he later (1986) apparently 

abandoned the group in his classification, treating 

euphausiids, amphionidaceans, and decapods as 

separate orders within the Eumalacostraca 

(Schram, 1986:543). Yet his cladogram (Schram, 

1986:530) and his accompanying text (1986:529) 

depict the eucarid line as distinct, and he refers to 

the eucarids as one of the recognizable lines of eumalacostracan 

evolution. And indeed, most treatments 

consider the Eucarida a valid superorder of 

the subclass Eumalacostraca, as did Bowman and 

Abele (1982) and most treatments since then (e.g., 

Christoffersen, 1988; Ruppert and Barnes, 1994; 


Brusca and Brusca, 1990; Mayrat and Saint Laurent, 

1996; Camp, 1998 (in Camp et al., 1998); 

Young, 1998). But as with nearly all other crustacean 

assemblages, this grouping has its opponents 

as well. Most of the disagreement concerns whether 

the mysidaceans (i.e., either mysids, lophogastrids, 

or both) should be placed here (see earlier discussions 

on mysids and lophogastrids) and what the 

relationships are among the three currently recognized 

orders Euphausiacea, Amphionidacea, and 

Decapoda (see Jarman et al., 2000; Richter and 

Scholtz, in press). Eucarid relationships have been 

analyzed by Schram (1984) and by Christoffersen 

(1988). Our classification is consistent with both of 

these analyses at higher levels but differs in the constituent 

suborders. 

ORDER EUPHAUSIACEA 

The Euphausiacea still contains only the two families 

Bentheuphausiidae (monotypic) and Euphausiidae 

(all other species). The treatment by Baker et 

al. (1990) follows this arrangement as well. A recent 

analysis of 28S rDNA sequence data by Jarman 

et al. (2000) suggests that euphausiaceans may 

be more closely related to the Mysida than to the 

Decapoda. 

ORDER AMPHIONIDACEA 

This order remains monofamilial and monogeneric 

(Amphionides). 

ORDER DECAPODA 

The decapods have been the subject of more published 

papers than have all other crustacean groups 

combined. This popularity stems in part from the 

economic importance of many groups (especially 

penaeoid shrimps, palinurid and nephropid lobsters, 

and portunid and xanthoid crabs) but also in 

part because of their marvelous diversity. The convenient 

size of most decapods predisposed them to 

become subjects of some of the earliest papers using 

biochemical and molecular data to resolve crustacean 

relationships. Yet we are as far from reaching 

a consensus on the relationships among the decapods 

as we are for the more obscure groups, and 

opinions and datasets remain sharply divided. In 

the treatment that follows, we have tried to address 

the many changes and arrangements that have been 

suggested since 1982 under the taxonomic heading 

for each major group of decapods. However, we are 

certain to have missed several important papers, 

and we hasten to remind the reader that the literature 

on this topic is vast. In general, we have settled 

on a fairly conservative classification of the 

decapods, knowing that, as with all other crustacean 

taxa, this group is destined for revision. Some 

of the many reviews of decapod classification that 

have appeared since the Bowman and Abele (1982) 

classification are Felgenhauer and Abele (1983), 

Abele and Felgenhauer (1986), Kim and Abele 

(1990), Abele (1991), Holthuis (1993a), and 

Scholtz and Richter (1995). 

The creation of two major branches of decapods, 

Dendrobranchiata and Pleocyemata, by Martin 

Burkenroad (1963, 1981) was a rather bold departure 

from previous schemes of decapod classification. 

According to Fenner Chace (pers. comm.), T. 

Bowman more or less accepted Burkenroad’s arguments 

without much questioning, and thus the 

use of the Dendrobranchiata and Pleocyemata in 

the Bowman and Abele (1982) classification. Chace 

(pers. comm.) feels that there is ample evidence for 

elevating many of the major groups of the Decapoda 

as Burkenroad did with the penaeoids and 

that singling out the penaeoid shrimp was to assign 

that group an artificial distinction. He is not alone. 

Holthuis (1993a; see especially pages 11–13 for a 

concise historical overview of the many attempts to 

classify the decapods) felt that treating the penaeoids 

as a separate group (the Dendrobranchiata) 

equal in rank to the combined Natantia � Macrura 

Reptantia � Anomura � Brachyura (the Pleocyemata 

of Burkenroad) was unsatisfactory. Holthuis 

(1993a) proposed to revert to the older classifications 

and treated the Natantia and the Macrura 

Reptantia as ‘‘full suborders of equal rank with the 

Anomura and Brachyura.’’ In his own words (Holthuis, 

1993a:6): 

I know that this classification will generally be considered 

old-fashioned: in several modern handbooks the 

suborder Natantia has been abandoned altogether; a 

small part of it, namely the Penaeoidea, is elevated to 

the rank of a separate suborder Dendrobranchiata 

while the rest of the Natantia plus the Macrura Reptantia, 

plus the Anomura, plus the Brachyura are 

placed in a single suborder Pleocyemata. This to me 

seems a very artificial and unsatisfactory arrangement, 

and I therefore still keep to the old classification. 

This ‘‘old’’ classification to which he refers, probably 

because of its simplicity and relative lack of 

controversy, is often encountered in popular and 

lay versions of crustacean classification. As an example, 

the publishers of the BIOSIS and Zoological 

Record databases (see URL http://www.york. 

biosis.org/zrdocs for the BIOSIS/Zoological Record 

Taxonomic Hierarchy, Section 10, Crustacea) have 

‘‘thrown up their hands in despair’’ (Chace, pers. 

comm.) and have reverted to this older and simpler 

classification. There, Natantia is treated as a taxon 

containing all of the known shrimp groups (Penaeidea, 

Caridea, and Stenopodidea) and the Reptantia 

is treated as containing the anomurans, astacurans, 

brachyurans, and palinurans. 

Yet the distinct nature of the penaeoids (the Dendrobranchiata) 

has been supported by additional 

morphological (e.g., Schram, 1984, 1986), embryological, 

spermatological (e.g., see Jamieson, 

1991a), and molecular data. Kim and Abele (1990) 

reviewed previous schemes of decapod classification 

and concluded, based on somewhat limited 

data from 18S rRNA, that the penaeids were distinct 

from other decapods. This view was support- 


ed with additional sequence data and additional 

taxon sampling by Abele (1991), whose review of 

morphological and molecular data supported a distinct 

Dendrobranchiata (the penaeoids) clade and 

also three other distinct clades corresponding to (1) 

the Caridea (including the procaridoids), (2) the 

Stenopodidea, and (3) a ‘‘reptant’’ lineage. (The latter 

lineage is responsible for most of the more troublesome 

remaining problems in decapod classification. 

As Abele (1991) stated, ‘‘there seems to be as 

many groupings of these taxa as there are authors 

who have studied them.’’) The artificiality of the 

‘‘Natantia’’ is also pointed out by Christoffersen 

(1988a) and Scholtz and Richter (1995). 

Thus, there is no morphological or molecular 

support for a natural ‘‘natantian’’ clade that contains 

all shrimp-like forms. The features that seem 

to unite the natantians appear to be primitive characters 

that do not clearly define a monophyletic 

group. Consequently, we have recognized the Dendrobranchiata 

and Pleocyemata on the basis of 

what appear (to us) to be shared, derived features 

of both morphological and molecular data. 

Within the Dendrobranchiata, classification is 

relatively stable, mostly because there are relatively 

few taxa in this suborder. Relationships among the 

pleocyemate taxa are another story. If Caridea and 

Stenopodidea are treated as separate clades, then 

an argument could be made for recognition of the 

Reptantia (or Macrura Reptantia, following Holthuis, 

1993a) as a natural taxon based on the work 

of Schram (1984, 1986), Abele (1991), Scholtz and 

Richter (1995), and others. Scholtz (pers. comm.) 

argues that the evidence for a monophyletic Reptantia 

is at least as convincing as the evidence for 

recognition of Caridea and other decapod infraorders, 

and we tend to agree. Yet the Reptantia of 

Abele (1991) and Scholtz and Richter (1995) differ 

as to the constituent groups, and we have opted for 

treating the ‘‘reptant’’ infraorders (Astacidea, Thalassinidea, 

Palinura, Anomura, and Brachyura) separately 

rather than combining them in a taxon that 

would be the sister group to the stenopodidean and/ 

or caridean shrimps. Recognition of a natural 

‘‘Reptantia’’ would involve using this name at the 

level of infraorder and then ‘‘demoting’’ the above 

five groups to just below the infraorder level, which 

would add considerably to the confusion in an assemblage 

that already contains a large number of 

taxonomic subdivisions. 

Scholtz and Richter (1995) attempted to place 

the classification of the reptant decapods on firm 

cladistic footing. They argued (as did Christoffersen, 

1988a) that the Reptantia was a clearly defined 

monophyletic taxon and that its sister group was 

possibly the Stenopodidea (which, according to 

other authors, are members of the same clade Pleocyemata). 

Thus, the branching sequence of the 

decapods would be Penaeoidea (Dendrobranchiata), 

then Caridea, Stenopodidea, and Reptantia; 

this much at least is consistent with other bodies of 

data (e.g., Schram, 1984, 1986; Jamieson, 1991a; 

Abele, 1991) (although Christoffersen (1988a:342) 

suggested that Stenopodidea was the sister group to 

a Caridea � Reptantia clade). In light of this support, 

it is curious, and possibly a mistake, that we 

have not included the Reptantia as a monophyletic 

clade in our classification, although inclusion or exclusion 

of the stenopodideans is unresolved. Scholtz 

and Richter (1995) argued convincingly for monophyly 

of some of the constituent reptant groups, 

such as the Brachyura and Anomura, but other arguments 

are (to us) less convincing. The Scholtz 

and Richter (1995) classification also included several 

new group names, such as the Achelata, Fractosternalia, 

Meiura, etc., which we feel are unlikely 

to persist (but note that some of these taxon names 

already have been employed (although not necessarily 

endorsed) in the papers of, e.g., Schmidt and 

Harzsch, 1999; Suzuki and McLay, 1998; Sternberg, 

1996; Taylor et al., 1999; and Taylor and 

Schram, 1999). For reasons we feel are inappropriate 

for discussion in a review and compilation of 

this nature (mostly differences in how we would 

score certain morphological characters and the low 

number of specimens examined), we have not followed 

Scholtz and Richter here. In fairness, some 

of the characters proposed by Scholtz and Richter 

are well beyond our ability to comment on (such 

as the shape of thoracic and cephalic ganglia and 

the development of embryonic growth zones) and 

possibly provide fertile ground for further investigations. 

And we acknowledge and compliment 

them on an attempt to place decapod classification 

in a phylogenetic context, which our classification 

clearly does not do. But concerns raised by their 

questionable (to us) use of morphological characters 

caused sufficient doubt as to their overall 

scheme, and we have not accepted the Scholtz and 

Richter (1995) arrangement in the current classification. 

The date of establishment of the name Decapoda 

has been changed to Latreille (1802) rather than 

Latreille (1803) (L. Holthuis, pers. comm.; see earlier 

comments in the section Names, Dates, and the 

ICZN). 

SUBORDER DENDROBRANCHIATA 

Christoffersen (pers. comm.) would rather we employ 

the name Penaeidea Dana instead of Dendrobranchiata 

Bate, as the former name is older and 

‘‘perfectly legitimate.’’ Holthuis (pers. comm.) 

agrees but notes that ‘‘since Dendrobranchiata 

seems to [have] become generally accepted, I am 

quite willing to go along.’’ Within the group, there 

have been no significant family-level or higher 

changes proposed (to our knowledge) since the 

Bowman and Abele (1982) classification. Authorship 

of the family Solenoceridae has been credited 

to Wood-Mason rather than to Wood-Mason and 

Alcock (Kensley, pers. comm.). Thus, our classification 

of the Dendrobranchiata is the same as that 


employed recently by Pérez Farfante and Kensley 

(1997). 

SUBORDER PLEOCYEMATA 

The Pleocyemata contains all nonpenaeoid decapods, 

whether swimming (natant) or crawling (reptant). 

The group appears to be monophyletic based 

on morphological data (e.g., Schram, 1984, 1986; 

Scholtz and Richter, 1995) and molecular data 

(e.g., Kim and Abele, 1990; Abele, 1991). 

INFRAORDER STENOPODIDEA 

For this section, we followed the classification provided 

by Holthuis (1993a), which does not appear 

to be very controversial. Authorship of the taxon 

Stenopodidea is changed from Bate (1888) to Claus 

(1872) at the recommendation of M. Tavares (pers. 

comm.). To our knowledge, there has been only one 

new family-level taxon described since the Bowman 

and Abele (1982) work. Schram (1986) erected the 

family Spongicolidae, so that there are now two 

recognized families of extant stenopodideans (see 

also Holthuis, 1993a). Schram et al. (2000) recently 

described the first known fossil stenopodidean, also 

attributed to the Spongicolidae. 

INFRAORDER CARIDEA 

For the carideans, we followed, for the most part, 

the classification provided by Holthuis (1993a), 

which is very similar to that suggested by Chace 

(1992) (see also Vereshchaka, 1997b, for a key to 

caridean superfamilies modified slightly from Chace, 

1992). But in contrast with the relative lack of controversy 

over dendrobranchiate or stenopodidean 

classification, there is apparently no consensus on 

the relationships or even the names of the incredibly 

diverse families of caridean shrimps. There have 

been several cladistic analyses conducted on groups 

of caridean families by M. Christoffersen (see especially 

Christoffersen, 1990). These studies would, 

if accepted, rearrange large numbers of caridean 

families. For example, in his 1986 paper, Christoffersen 

placed seven families (oplophorids, atyids, 

pasiphaeids, alvinocarids, bresiliids, psalidopodids, 

and disciadids) in the superfamily Atyoidea, in contrast 

with Chace (1992) and Holthuis (1993a), who 

treated the Atyoidea as containing only the family 

Atyidae. Christoffersen points out (pers. comm.) 

that, among the ‘‘glaringly non-monophyletic assemblages’’ 

in our current classification, are the Alpheoidea, 

Hippolytidae, Pandaloidea, and Nematocarcinoidea. 

Adding to Christoffersen’s frustration 

(pers. comm.) is that, whereas many authors 

comment on the unsatisfactory state of current classifications, 

especially as concerns such ‘‘wastebasket’’ 

assemblages as the Hippolytidae and Pandaloidea, 

his own suggestions for novel arrangements 

have been slow to catch on. Chace (1997) recognizes 

the Hippolytidae Bate, and Holthuis (1993a) 

elected to synonymize a large number of Christof- 

fersen’s new taxa. Thus, we are left with the difficult 

task of following older yet clearly nonphylogenetic 

listings (e.g., Chace, 1992; Holthuis, 1993a) 

vs. cladistically generated phylogenetic arrangements 

(e.g., Christoffersen, 1987, 1988a, b, 1989a, 

b, 1990) that seem to have little following in the 

carcinological community and for which, in our estimation, 

some of the employed characters are 

questionable. We have followed Holthuis’s lead, 

more in deference to his vast knowledge of the carideans 

than for any other reason, while acknowledging 

that there have been alternative phylogenetically 

based ideas presented in the literature. Only 

those superfamilies for which there have been 

changes subsequent to Bowman and Abele (1982) 

are mentioned below. 

Superfamily Galatheacaridoidea 

The family Galatheacarididae and its superfamily 

Galatheacaridoidea were both described by Vereshchaka 

(1997b) for the species Galatheacaris abyssalis 

based on a single specimen. Additional specimens 

have since been found in the stomachs of 

deep-sea lancetfish (Chow et al., 2000). 

Superfamily Bresilioidea 

This assemblage has long been recognized as being 

an artificial group in dire need of revision (e.g., see 

Forest, 1977). Holthuis (1993a) elected to treat the 

Bresiliidae as a family and placed in synonymy 

some of the recently proposed families (Agostocaridae, 

Alvinocarididae). We have treated the group 

as an (admittedly) artificial superfamily containing 

five caridean families that may or may not be related. 

Three of these families are new (i.e., they 

were not included in the Bowman and Abele (1992) 

classification): the family Agostocarididae was 

erected by Hart and Manning (1986), the Alvinocarididae 

was proposed by Christoffersen (1986), 

and the Mirocarididae was described by Vereshchaka 

(1997a). 

Christoffersen’s (1986) family Alvinocarididae is 

recognized to accommodate the majority of the 

morphologically similar ‘‘bresilioid’’ shrimp from 

hydrothermal vents. The family was more thoroughly 

(although still somewhat incompletely) diagnosed 

by Segonzac et al. (1993) in a footnote and 

also by Vereshchaka (1996, 1997a) (see also Shank 

et al., 1999). Vereshchaka (1997a) created a new 

genus (Mirocaris) and family, the Mirocarididae, 

for the hydrothermal vent shrimp described originally 

as Chorocaris fortunata by Martin and Christiansen 

(1995b). 

Superfamily Campylonotoidea 

The family Bathypalaemonellidae was established 

(although without a description or diagnosis and 

without mention of the genus Bathypalaemonella; 

see Holthuis, 1993a:87) by Saint Laurent (1985). 

The family is placed in the superfamily Campylon- 


otoidea on the recommendation of L. Holthuis 

(1993a:87, and pers. comm.). 

Superfamily Palaemonoidea 

The family Euryrhynchidae Holthuis, 1950, was 

added on the recommendation of Holthuis (pers. 

comm.). The family Kakaducarididae was described 

by A. J. Bruce (1993) as a subfamily of the 

Palaeomonidae and is here treated as a family on 

the recommendation of L. Holthuis (pers. comm.). 

Superfamily Alpheoidea 

Authorship of the family Ogyrididae remains credited 

to Holthuis (1955). Although Hay and Shore 

(1918) established the family Ogyridae, as noted by 

M. Tavares (pers. comm.), L. Holthuis (pers. 

comm.) points out that they based it on the type 

genus Ogyris Stimpson, 1860, which is a junior 

homonym of Ogyris Westwood and is thus invalid. 

Stebbing (1914) proposed the replacement genus 

Ogyrides, and thus the family name is Ogyrididae, 

first used as such by Holthuis (1955). We have not 

followed Christoffersen’s (1987) suggestion to 

transfer the family Processidae to the Crangonoidea 

or to combine the alpheoids and crangonoids and 

pandaloids into one monophyletic taxon. Christoffersen 

(1987) also proposed the new alpheoid families 

Nauticarididae (to contain Nauticaris and Saron), 

Alopidae (to contain Chorismus, Alope, and 

Caridion), and Bythocarididae (to contain Bythocaris, 

Cryptocheles, and Bathyhippolyte). We have 

not followed these suggestions, nor have we recognized 

the families Merhippolytidae and Thoridae 

recognized by Christoffersen (e.g., Christoffersen 

1998). 

Christoffersen later (1987) also suggested the recognition 

of the family Barbouridae (spelling corrected 

to Barbouriidae by Christoffersen, 1990), to 

include the genera Barbouria, Janicea, and Parhippolyte. 

In his review of caridean shrimps of the Albatross 

Philippine Expedition, Chace (1997), although 

finding ‘‘no clear evidence to support the 

superfamilial categories suggested by Christoffersen 

(1987),’’ found ‘‘considerable reason to endorse his 

[Christoffersen’s] establishment of the Barbouriidae.’’ 

Chace refrained from treating these genera 

as Barbouriidae in that paper, but we have taken 

that step here and recognize the Barbouriidae. Inclusion 

of the family in the superfamily Alpheoida 

is because of the similarities to hippolytids (all three 

genera were formerly treated as members of the 

Hippolytidae). 

Superfamily Crangonoidea 

As noted above, Christoffersen (1987) proposed the 

family Barbouriidae for the genera Barbouria, Janicea, 

and Parhippolyte and originally placed the 

family in the superfamily Crangonoidea. We treat 

it here as a member of the Alpheoidea because of 

the similarities to the alpheoid family Hippolytidae 

(see Chace, 1997:40). 

Superfamily Pandaloidea 

Christoffersen (1989) suggested a new classification 

of this superfamily, wherein he proposed many significant 

changes. Three new families were proposed 

(Plesionikidae for the genus Plesionika, Heterocarpoididae 

for the genus Heterocarpoides, and Dorodoteidae 

for the genus Dorodotes). In addition, 

the family Physetocarididae was removed from its 

own superfamily and placed in the Pandaloidea, 

and the family Heterocarpidae was recognized. No 

diagnoses of the new taxa were provided (although 

character states were given), and we have opted to 

not recognize these changes for now. 

INFRAORDER ASTACIDEA 

Although we are not recognizing the ‘‘Macrura 

Reptantia’’ as a suborder (see above), for the most 

part, we have followed the admittedly conservative 

classification of Holthuis (1991) for the superfamilies 

and families of the Astacidea (see also Williams, 

1988, for classification of commercially important 

lobster families). Holthuis, who was at the 

time dealing only with the marine lobsters and so 

did not include the parastacoids and astacoids, 

treated marine astacideans as belonging to a single 

superfamily Nephropoidea containing two families, 

Thaumastochelidae and Nephropidae. Our classification 

differs only in the inclusion of the Enoplometopoidea 

(see below) and Gylpheoidea, the latter 

placed by Holthuis among the infraorder Palinura 

(his Palinuridea). Scholtz (1999) recently reviewed 

the freshwater crayfishes (Astacoidea and Parastacoidea) 

and argued that they are members of a distinct 

clade, Astacida, that is not closely related to 

clawed lobsters. However, strong molecular evidence 

suggests that clawed lobsters are indeed the 

sister group to the astacids (Crandall et al., 2000). 

Superfamily Glypheoidea 

The primitive family Glypheidae (the only extant 

family in the Glypheoidea) has been transferred to 

the Astacidea as per the recommendations of Forest 

and Saint Laurent (1989). The taxon name, credited 

to Zittel in Bowman and Abele (1982), has 

now been credited to the earlier usage by Winckler 

(M. Hendrickx, pers. comm.), following the usage 

in Glaessner (1969). 

Superfamily Enoplometopoidea 

The genus Enoplometopus was assigned its own superfamily 

and family (Enoplometopoidea, Enoplometopidae) 

by Saint Laurent (1988). 

Superfamily Nephropoidea 

Tshudy and Babcock (1997) examined fossil and 

extant clawed lobsters and indicated that the family 

Thaumastochelidae, at least as used previously, 

may be paraphyletic. We have not taken the extra 

step of deleting this family (which would result in 


the former thaumastochelids being treated as Nephropidae), 

as there was also strong support in 

their analysis for grouping at least some thaumastochelid 

genera together (Tshudy and Babcock, 

1997: fig. 1). 

Superfamilies Astacoidea and Parastacoidea 

Monophyly of the freshwater crayfishes now appears 

secure based on adult morphology, sperm ultrastructure, 

embryology, and molecular data (e.g., 

see Crandall, 1999; Crandall et al., 2000; Scholtz, 

1998, 1999). Scholtz (1998, 1999) reviews evolution 

of the crayfishes and confirms that there are 

two distinct clades within the group (i.e., within his 

Astacida) corresponding to the northern hemisphere 

Astacoidea (families Cambaridae and Astacidae, 

the latter of which is probably paraphyletic) 

and the southern hemisphere Parastacoidea (family 

Parastacidae). Crandall et al. (2000), using over 

3000 nucleotides from 3 different genes, have confirmed 

both the monophyly of the freshwater crayfishes 

(Astacoidea � Parastacoidea) as well as the 

monophyly of the astacoid and parastacoid clades. 

Thus, our current classification is misleading in that 

these two superfamilies (the Astacoidea and Parastacoidea) 

are still treated as being of equal rank 

with three other superfamilies in the Astacidea 

when in fact they need to be depicted as more closely 

related to each other than either is to any other 

astacidean superfamily. Scholtz (1999) also proposes 

that the crayfishes are not closely related to homarids 

(not supported by Crandall et al., 2000) but 

are instead members of ‘‘a large group including 

Thalassinida, Anomala and Brachyura’’ (see also 

Scholtz and Richter, 1995). Taylor et al. (1999) 

added some insights into evolution within the 

group based on well-preserved fossil material from 

China. 

INFRAORDER THALASSINIDEA 

Monophyly of the thalassinideans is uncertain; at 

least some morphological and molecular analyses 

indicate that the group is not monophyletic (e.g., 

Tudge, 1995; Morrison and Cunningham, 1999). 

The propensity to construct complex vertical burrows 

is one character that has been postulated as 

defining the group (Atkinson and Taylor, 1988; 

Griffis and Suchanek, 1991; Scholtz and Richter, 

1995), as has the presence of a dense row of long 

setae along the lower margin of the second leg 

(Poore, 1994, 1997). We have followed the revision 

by Poore (1994:92) (who also established the family 

Strahlaxiidae), with the only difference being 

that some of the authors and dates of some taxa 

have been changed to earlier usages according to 

L. Holthuis (pers. comm.). Relationships among the 

extant superfamilies, families, and genera were suggested 

by Poore (1994). Poore’s resulting classification 

(1997:92), like ours, does not adequately 

display all of the relationships suggested by his phylogenetic 

analysis (Poore, 1994:120). In particular, 

the Axioidea is the sister group to the Thalassinoidea 

� Callianassoidea in his phylogeny, whereas 

in his classification, all three are treated as superfamilies. 

The family Ctenochelidae is acknowledged 

by Poore (1994) to be paraphyletic (although Tudge 

et al., 2000, argued for ctenochelid monophyly). 

Poore (1997) subsequently addressed three of these 

families and their relationships in greater detail 

(Callianideidae Kossman, Micheleidae Sakai, and 

Thomassiniidae de Saint Laurent). 

In the superfamily Callianassoidea, the family 

Axianassidae was removed by Poore (1994), and 

the family Ctenochelidae was erected by Manning 

and Felder (1991). As noted above, Tudge et al. 

(2000) supported the monophyly of the family Callianassidae 

and the family Ctenochelidae (while 

noting that the latter includes, at least in their analysis, 

the genus Anacalliax, considered by some 

workers to belong to the Callianassidae). In the superfamily 

Axioidea, Sakai (1992) first established 

the subfamily Micheleinae, elevated to family level 

(Micheleidae Sakai) by Poore (1994), and Poore 

(1994) erected the Strahlaxiidae. Sakai (1999) recently 

has proposed some rather large-scale revisions 

within the Callianassidae; his revisions are apparently 

at odds with other analyses of the same or 

similar taxa (e.g., see Tudge et al., 2000). 

INFRAORDER PALINURA 

Holthuis (1991) referred to this assemblage as the 

Palinuridea, a spelling that would be consistent 

with some of our other infraorder names (such as 

Stenopodidea, Caridea) but not with others (Anomura, 

Brachyura). We have retained the spelling 

Palinura. Within the superfamily Palinuroidea, Davie 

(1990) felt that synaxids were not deserving of 

separate familial status and synonymized the family 

Synaxidae with the Palinuridae. However, Holthuis 

(1991) continued to recognize them as separate 

families, and we have maintained them as separate 

families here as well. The family Polychelidae has 

been recently reviewed and rediagnosed by Galil 

(2000). Removal of the glypheoids from this infraorder 

to the Astacidea has been noted above. 

INFRAORDER ANOMURA 

Our classification follows McLaughlin’s (1983b) 

fairly closely, with the exception of the use of the 

family name Pylochelidae replacing Pomatochelidae 

(following Forest, 1987). McLaughlin (1983a, 

b) employed the name Anomala De Haan (as had 

Burkenroad, 1981) rather than Anomura H. Milne 

Edwards, which had been used by Bowman and 

Abele (and many other workers). G. Scholtz (pers. 

comm.) also would prefer this usage (Anomala over 

Anomura), arguing that when the thalassinoid families 

are removed the taxon composition changes 

and thus the name Anomala is the more accurate. 

Use of Anomala over Anomura was reconsidered 

and discussed at length by McLaughlin and Holthuis 

(1985), who pointed out that both names 


have been used inconsistently in the past and that 

there are no rules governing the name given to a 

taxon above the family-group level. Thus, according 

to McLaughlin and Holthuis, the Rule of Priority 

need not be applied (Anomala is, strictly 

speaking, the older of the two names). Furthermore, 

they argued that, for stability, the name Anomura 

MacLeay, 1838, should be used for the taxa 

traditionally considered to belong to this group 

(lomisoids, galatheoids, paguroids, and hippoids), 

and we have followed their suggestion. Phylogenetic 

relationships within the Anomura remain largely 

unsettled; studies addressing this question include 

McLaughlin (1983b), Martin and Abele (1986), 

Cunningham et al. (1992), Tudge (1997b), Mc- 

Laughlin and Lemaitre (1997, 2000), and Morrison 

and Cunningham (1999). 

McLaughlin (1983a) recognized the unusual nature 

of Lomis hirta and placed it in its own family 

(Lomidae) and superfamily (Lomoidea) (corrected 

herein to Lomisidae and Lomisoidea, respectively). 

McLaughlin (1983b) concluded that the hermit 

crab families were monophyletic, and she therefore 

treated all six families as members of the superfamily 

Paguroidea. This arrangement has been adopted 

by a variety of workers (e.g., Tudge, 1991; Richter 

and Scholtz, 1994; Scholtz and Richter, 1995; Tudge, 

pers. comm.) and seems to us both logical and simple, 

and we have used it here. In his treatment of 

the Pylochelidae (treated as Pomatochelidae in 

Bowman and Abele, 1982), Forest (1987) indicated 

that the family is more closely allied with the Diogenidae 

than with other anomuran families, but 

we have not indicated this alliance pending formal 

recognition of that relationship. 

The family name Lomisidae and the superfamily 

name Lomisoidea, containing only the monotypic 

genus Lomis, occasionally have been spelled, beginning 

with Glaessner (1969), as Lomidae and Lomoidea 

(see especially McLaughlin, 1983a). However, 

the genus Lomis is not a Greek or Latin word, 

and thus it has no Greek or Latin stem (such as 

Lom-) to which the -idae ending can be added; the 

original author of Lomis, Bouvier, coined the 

French common name ‘‘Lomisinés’’ for these crabs 

(G. Poore, pers. comm.). Thus, the preferred spelling 

for the family is Lomisidae and for the superfamily 

is Lomisoidea. 

A recent analysis of anomuran phylogeny based 

on mitochondrial DNA gene rearrangements (Morrison 

and Cunningham, 1999; C. Morrison and C. 

Cunningham, pers. comm.) largely supports Mc- 

Laughlin’s (1983b) recognition of the major anomuran 

groups and their phylogeny. According to 

the findings of Morrison and Cunningham (1999), 

lithodids are strongly associated with pagurids and 

together these groups constitute a monophyletic 

clade (confirming the earlier report by Cunningham 

et al., 1992). The Hippoidea is also strongly supported 

as a monophyletic clade, and the Galatheoidea 

(including both Aegla and Lomis) is depicted 

as basal to the remaining Anomura. Thus, a clas- 

sification based on these data would differ from 

McLaughlin’s (1983a, b) in that the superfamily 

Lomisoidea would be removed, with the monotypic 

Lomisidae being placed within the Galatheoidea 

(which also contains the Aeglidae, Porcellanidae, 

Galatheidae, and Chirostylidae; see Baba (1988) 

for a thorough review of the latter family). However, 

support for this particular node (placement of 

Lomis) was not as strong in the Morrison and Cunningham 

tree, and indeed C. Morrison (pers. 

comm.) has suggested that we might be better off 

depicting a separate lineage for Aegla and Lomis 

from the remaining galatheoids. We have for now 

retained Lomis in its own family and superfamily, 

the Lomisoidea, which we have placed adjacent to 

the Galatheoidea as a concession to the new data. 

Similarly, we have moved the Paguroidea closer to 

the Hippoidea, also reflecting the findings of Morrison 

and Cunningham (1999). Several workers 

have discussed the fact that the lithodids (at least 

some of them) appear to have stemmed from within 

the Paguridae (Cunningham et al., 1992; Richter 

and Scholtz, 1994; Tudge, 1991, Tudge et al., 1998; 

C. Morrison, pers. comm.). Additionally, Cunningham 

(pers. comm.) suggested a rather close tie between 

the Aeglidae (restricted to freshwater streams 

and lakes in temperate South America) and the 

Lomisidae (a monotypic and exclusively marine 

family known only from Australia). According to 

Scholtz and Richter (1995), two groups of the Anomura, 

hippoids and galatheoids, share the apomorphic 

character of a telson stretch receptor not 

found in any other malacostracan group (Scholtz 

and Richter, 1995, citing Paul, 1989). 

In contrast with the phylogenetic hypotheses of 

McLaughlin (1983b) and Morrison and Cunningham 

(1999), evidence from sperm ultrastructure 

(reviewed in Tudge, 1997b) would suggest that the 

Anomura is not monophyletic, that Lomis does not 

belong to the Anomura sensu stricta, that at least 

some of the thalassinoids are within the Anomura, 

and that the superfamilies Thalassinoidea, Paguroidea, 

and Galatheoidea are not monophyletic. Because 

at this time the bulk of the evidence (i.e., 

adult morphology combined with molecular sequence 

and gene arrangement data) seems to support 

the more conservative approach of Mc- 

Laughlin (1983b), we have modified our arrangement 

of anomuran taxa only slightly. Our classification 

is therefore more in agreement with the 

findings of Morrison and Cunningham (1999) than 

with the sperm ultrastructural findings presented by 

Tudge (1997b). 

In the Bowman and Abele (1982) classification, 

the hermit crab families were divided among two 

superfamilies, Coenobitoidea and Paguroidea. The 

Coenobitoidea was removed following the suggestion 

of McLaughlin (1983b), and the family Coenobitidae 

is now treated within the superfamily Paguroidea. 

Thus, our infraorder Anomura contains 

four superfamilies: Lomisoidea (the distinctness of 

which is questionable in light of the Morrison and 


Cunningham (1999) data, which suggest placement 

in the galatheoid clade), Galatheoidea, Paguroidea, 

and Hippoidea (spermatozoal characters of which 

are described by Tudge et al., 1999). The paguroids 

(which in our scheme include the former coenobitoids) 

and hippoids should be considered sister taxa 

and together are the sister taxon to the Galatheoidea, 

according to Morrison and Cunningham 

(1999) and C. Morrison (pers. comm.). 

INFRAORDER BRACHYURA 

Subsequent to the Bowman and Abele (1982) classification, 

there has been relatively widespread use 

of a scheme first suggested by Guinot (1977, 1978, 

1979; see also Saint Laurent, 1979; Guinot and 

Bouchard, 1998) that recognizes three morphological 

‘‘grades’’ of brachyuran crabs (which she called 

the Podotremata, Heterotremata, and Thoracotremata) 

based mostly on the coxal vs. sternal location 

of the male and female genital apertures. Although 

Abele (1991) and Spears et al. (1992) found no molecular 

support for these divisions, some spermatological 

data seemed to support them (e.g., see Jamieson, 

1994; Jamieson et al., 1994a, b, 1995). The 

latter two groups (Heterotremata and Thoracotremata) 

were treated jointly as the Eubrachyura by 

Saint Laurent (1980a, b), and various authors (e.g., 

Schram, 1986) have followed this arrangement as 

well. At the same time, there is also growing evidence 

from molecular sequence data (e.g., Spears et 

al., 1992; Abele and Spears, 1997; Spears and 

Abele, 1999; Spears, pers. comm.) and from mitochondrial 

gene rearrangement data (Morrison and 

Cunningham, 1999; Morrison, pers. comm.) that 

the true crabs (Brachyura) can be divided into two 

major clades, one containing the dromiacean families 

and the other containing all ‘‘higher’’ crabs, 

and including the raninids. The two ideas are not 

totally incompatible, but at the same time, they 

cannot be completely reconciled. The main areas of 

disagreement concern the limits of the ‘‘true’’ crabs, 

the placement of several families traditionally 

thought of as being ‘‘primitive’’ (dromiids and raninids 

in particular), and the recognition of various 

assemblages (tribes, sections, etc.) within the major 

divisions. Evidence brought to bear on these issues 

has come from many fields, such as larval morphology 

(e.g., Rice, 1980, 1983, 1988; Martin, 

1988, 1991), sperm morphology (e.g., Jamieson, 

1991a, b, 1994), adult morphology (e.g., Sˇtevčić, 

1995, 1998; McLay, 1991, 1999; Guinot and Bouchard, 

1998), and molecular sequence data (e.g., 

Spears et al., 1992). 

Guinot (1977, 1978) originally defined the section 

Podotremata as containing the dromioids, 

homoloids, raninoids, and tymoloids. The Podotremata 

was suggested to be monophyletic on the basis 

of sperm ultrastructure (Jamieson, 1994) and yet 

paraphyletic on the basis of rRNA sequences 

(Spears and Abele, 1988; Spears et al., 1992). To 

quote Guinot and Bouchard (1998), ‘‘Monophyly 

versus paraphyly of the Podotremata and their possible 

placement as the sister group of the heterotreme-thoracotreme 

assemblage remain open questions.’’ 

Within the Podotremata, Guinot (1977, 

1978) recognized a subsection Dromiacea to contain 

two superfamilies, Dromioidea and Homolodromioidea, 

and a subsection Archaeobrachyura to 

contain the superfamilies Raninoidea, Homoloidea, 

and Tymoloidea. The molecular data (e.g., Spears 

et al., 1992; Spears and Abele, 1999; Morrison and 

Cunningham, 1999; Spears, pers. comm.) do not 

support this arrangement. Although one group of 

crabs, corresponding to the Dromiacea of Guinot 

and earlier workers, does appear separate from other 

‘‘higher’’ crabs, nearly all evidence to date points 

to the fact that the raninids are not members of this 

dromioid clade (in contrast with the conclusions of 

Sˇtevčić, 1973, 1995, 1998), and thus the Podotremata 

cannot be recognized as originally envisioned. 

Instead, the raninids appear to be basal members 

of the second ‘‘higher crab’’ clade. 

Thus, we have decided to abandon the concept 

of the Podotremata. The Brachyura is herein depicted 

as being composed of two major clades. The 

groups formerly treated as ‘‘podotremes’’ are split, 

with dromiaceans in one major clade and all other 

crabs in the other major clade. We are referring to 

the first clade as the section Dromiacea, a name 

that has much historical usage and that is well 

known among brachyuran researchers. This clade 

(section Dromiacea) is the sister group to all of the 

higher crab families. In our treatment, it contains 

the superfamily Homolodromioidea and its sole 

family Homolodromiidae, the superfamily Dromioidea 

containing the families Dromiidae and Dynomenidae, 

and the superfamily Homoloidea containing 

the Homolidae, Latreilliidae, and Poupiniidae 

(the latter established by Guinot, 1991). 

The second major clade (all other crab families 

and superfamilies) is then treated collectively as the 

section Eubrachyura, a name coined by Saint Laurent 

(1980a, b) for this assemblage (but now including 

the raninoids, which were excluded by 

Saint Laurent). We note, however, that Sˇtevčić 

(1973, 1995, 1998) would retain raninids with 

dromiids, and Jamieson et al. (1994b) argue, based 

on sperm morphology, against any raninid/higher 

crab sister group relationship. Inclusion of the raninoids 

among the Eubrachyura also might be questioned 

on the basis of the fact that they lack the 

‘‘sella turcica’’ of the endophragmal system (see Secretan, 

1998). Within this enormous clade Eubrachyura, 

we are recognizing three subsections. 

First, we are treating the raninids and their allies 

(the former tymolids, now treated as the Cyclodorippoidea; 

see below) as the subsection Raninoida. 

We could have used for this group the name Archaeobrachyura, 

a name that has been used previously 

for the assemblage that contained raninoids, 

homoloids, and tymoloids (Saint Laurent 1980a, b) 

while they were still considered members of the 

‘‘podotreme’’ lineage. However, use of the name Ar- 


chaeobrachyura would have been confusing, not 

only because the constituency and alliances have 

changed considerably from its original usage by 

Guinot but because the entire group has been 

moved to the other major crab clade. We also could 

have used the older name Gymnopleura, established 

by Bourne (1922) to accommodate the raninids 

and still used by some modern workers (e.g., 

Dai and Yang, 1991). But we have now placed the 

former tymoloids (now the Cyclodorippoidea) in 

this subsection with the raninids (which may be a 

mistake; see below). Hence, our use of the name 

Raninoida for the subsection. We have credited this 

higher taxon to the same authority (De Haan) who 

established the family Raninidae. The other two 

subsections (the subsections Heterotremata and 

Thoracotremata), jointly constituting the sister 

group to the Raninoida, are more or less as envisioned 

by Guinot (1977, 1978, 1979). Our adoption 

of Guinot’s scheme (minus the Podotremata) 

has meant that many formerly recognized ‘‘tribes’’ 

or ‘‘sections’’ among the higher crabs have been removed. 

This reflects not so much an advance in our 

knowledge of which families are closely related but 

rather knowledge concerning which ones are not. 

For example, the formerly recognized Oxyrhyncha 

appears to be an artificial assemblage (Sˇtevčić and 

Gore, 1982; Jamieson, 1991a, b, 1994; Spears et 

al., 1992), and there is no longer any justification 

for recognizing the Oxystomata, Brachyrhyncha, 

and other former sections or tribes (e.g., see Guinot, 

1977, 1978; Spears et al., 1992; Sˇtevčić, 1998). 

Thus, we have retained several of the crab superfamilies 

but have removed many of the sections that 

were found in the Bowman and Abele (1982) classification. 

Yet acceptance of the sections Heterotremata 

and Thoracotremata as natural monophyletic 

lineages is by no means universal. For one thing, 

Guinot herself never explicitly assigned every 

known family to one of her sections, leaving some 

families ‘‘orphaned’’ in her earlier publications. 

And as noted above, these groups are admittedly 

(Guinot 1977, 1978) ‘‘grades’’ rather than true 

monophyletic lineages (or at least, if they are monophyletic, 

this has yet to be demonstrated, although 

there are preliminary data from morphology (see 

below) and from 16S rDNA (Trisha Spears, pers. 

comm.) that at least the Thoracotremata may have 

some validity). While usage of these sections has 

become relatively widespread, it is unfortunate that 

many families were not explicitly mentioned by 

Guinot, such that users of her classification have 

been uncertain as to which families belonged to 

which section. Schram (1986) provided a more 

complete list of families (including some known 

only from fossils). 

Concerning monophyly of the Thoracotremata, 

dissections of the male reproductive tract of a series 

of freshwater crabs and some marine heterotremes 

and thoracotremes (during a search for the sister 

taxon of the freshwater crabs) has indicated that 

the Thoracotremata is a monophyletic group 

(Sternberg and Cumberlidge, 2001). One character 

uniting the thoracotremes is that the distal tracts of 

the vas deferentia pass through thoracic endosternite 

8 and contact the male pleopods via apertures 

on thoracic sternite 8. The situation in heterotremes 

is different, with the vas deferens passing through 

the musculature of endosternite 8 but also through 

the coxa of pereiopod 5 such that the male sexual 

tube contacts the pleopods via an aperture on the 

coxopodite. According to Sternberg et al. (1999), 

Sternberg and Cumberlidge (2001), and Cumberlidge 

and Sternberg (pers. comm.), the Eubrachyura 

(sensu Saint Laurent, 1980) are therefore defined by 

females with sternal vulvae and males with sexual 

tube outlets that open on the coxa of pereiopod 5. 

The Thoracotremata constitutes a monophyletic 

subset of the Eubrachyura characterized by male 

sexual tube outlets that unambiguously open on the 

sternum. 

Within these last two subsections (Heterotremata 

and Thoracotremata), many former subfamilies of 

crabs, notably in the Xanthoidea and Majoidea and 

some also in the Parthenopoidea, have been elevated 

to family status based on the publications of several 

workers (e.g., Serène, 1984, for xanthids; Hendrickx, 

1995, for majids). This is an ongoing trend 

that merely reflects our growing awareness of how 

incredibly diverse these taxa are. 

SECTION DROMIACEA 

In an early version of the updated classification, we 

had removed the dromiacean crabs from the Brachyura 

and had placed them instead among the Anomura. 

Larval characters have suggested this for 

years (e.g., see Williamson, 1976, 1982; Rice, 

1980, 1983; Martin, 1991), so much so that Williamson 

(1988a, b) invoked an unusual hypothesis 

of transspecific gene flow to account for it. Molecular 

(18S rRNA) evidence brought to bear by 

Spears et al. (1992) seemed to indicate that at least 

some dromiaceans are indeed closer to the Anomura 

than to the Brachyura sensu stricta based on 

these preliminary data, and early studies of dromiacean 

sperm morphology suggested their removal 

from the Brachyura as well (Jamieson 1990, 

1991a). Yet adult morphology has always suggested 

that dromiids are true crabs (e.g., see Sˇtevčić, 

1995), and moving the dromiids to the Anomura 

would raise many additional questions. Should all 

of the families associated with dromiids (i.e., the 

former Dromiacea, including dromiids, dynomenids, 

and homolodromiids) be moved to the Anomura, 

even though larval and molecular evidence 

are not in hand for all of them? Is the Dromiacea 

in fact a valid, monophyletic grouping? If that 

scheme were accepted, how many other ‘‘primitive’’ 

families should be moved? The fact that information 

on larval, molecular, and sperm morphology 

characters is still lacking for many members of this 

assemblage, plus more recent molecular data 

(Spears and Abele, 1999; T. Spears, pers. comm.), 


eventually led us to keep dromiids with the other 

‘‘primitive’’ brachyurans in our section Dromiacea, 

knowing that by so doing we are continuing to displease 

students of crab phylogeny who rely mostly 

on larval characters and that the current arrangement 

of primitive crabs is not completely in keeping 

with the molecular evidence in the Spears et al. 

(1992) study. A detailed discussion of the situation 

within the Dromiacea can be found in the review 

of the Dynomenidae by McLay (1999). 

Superfamily Dromioidea 

The families Dromiidae and Dynomenidae are still 

listed as valid families, although based on molecular 

data (Spears et al., 1992) and sperm morphology 

(Jamieson, 1994; Jamieson et al., 1995; Guinot 

et al., 1998), their monophyletic status has been 

questioned (but see McLay, 1991, 1999; Sˇtevčić, 

1995). Earlier classifications, some of which have 

included the Homolidae among the dromiacean 

families, are reviewed by Sˇtevčić (1995), Guinot 

and Richer de Forges (1995), and McLay (1999). 

Guinot et al. (1998) argue that the Dromioidea (referred 

to as Dromiacea in that paper, a lapsus calami, 

Guinot, pers. comm.), containing the three 

families Dromiidae, Dynomenidae, and Homolodromiidae, 

is a valid monophyletic superfamily, although 

they note the differences separating the 

homolodromiids. We have maintained the separate 

status of the homolodromiids (i.e., placing them in 

their own superfamily Homolodromioidea; see below) 

in light of the many morphological features of 

adults that seem to separate them from the dromiids 

and dynomenids. In doing so, we follow Guinot 

(1995), even though Guinot and Bouchard (1998) 

have reverted to treating all three of these families 

in one superfamily (their Dromiacea). The families 

were reviewed recently by McLay (1991, Dromiidae; 

1999, Dynomenidae) with special regard to 

their Indo-Pacific members. 

Superfamily Homolodromioidea 

Separate superfamily status for the Homolodromiidae 

appears warranted on the basis of larval and 

adult morphology (see Martin, 1991; Guinot, 

1995). Sˇtevčić (1998) considers the homolodromiids 

the most primitive extant family of brachyuran 

crabs. The date of Alcock’s establishment of the 

Homolodromiidae has been changed from 1899 to 

1900 following the revision by Guinot (1995). 

Superfamily Homoloidea 

The alliance of homolids with dromiids has been 

supported by ultrastructural characters of the 

sperm (Guinot et al., 1994; see also the extensive 

review by Guinot and Richer de Forges, 1995). The 

family Poupiniidae was added by Guinot (1991). 

SECTION EUBRACHYURA, SUBSECTION 

RANINOIDA 

Superfamily Raninoidea 

Within the Raninoidea, the subfamily Symethinae 

(monogeneric; Symethis Goeke) was elevated to 

family level by Tucker (1998), as had been suggested 

earlier by Guinot (1993). However, Tucker 

did not agree with the removal of the subfamily 

Cyrtorhininae from the Raninidae, which had been 

suggested as a possibility by Guinot (1993). 

Superfamily Cyclodorippoidea 

The superfamily Tymoloidea has been removed and 

in its place is the superfamily Cyclodorippoidea, as 

the family name Cyclodorippidae Ortmann has seniority 

over Tymolidae Alcock, according to Guinot 

(pers. comm.) and Tavares (1991, 1993). Tavares 

(1998) also established a new family, the Phyllotymolinidae, 

within the Cyclodorippoidea. Guinot 

and Bouchard (1998) continue to recognize the superfamily 

Cyclodorippoidea (as did Tavares, 1991, 

1993, 1998), stating that this was done ‘‘for convenience’’ 

while at the same time cautioning against 

possible paraphyly in the assemblage. 

Placement of this superfamily with the raninoids 

in the Raninoida is possibly a mistake; molecular 

data seem to indicate a placement somewhere between 

the raninids and the higher eubrachyurans 

(T. Spears, pers. comm.). 


HETEROTREMATA 

Superfamily Dorippoidea 

The family Orithyiidae Dana has been transferred 

to this superfamily based on the suggestion of Bellwood 

(1996, 1998; see below). 

Superfamilies Calappoidea and Leucosioidea 

The monophyly of the family Calappidae and its 

constituent subfamilies has been questioned recently. 

Bellwood (1996, 1998) has recommended that 

only the families Calappidae and Hepatidae be retained 

in the superfamily Calappoidea, with the 

Matutidae joining the leucosiids in the Leucosioidea 

and with the Orithyiidae transferred to the dorippoids. 

To some extent, these changes reflect earlier 

suggestions based on larval (Rice, 1980) and 

adult (Guinot, 1978; Seridji, 1993) morphology, 

and there is at least some fossil support for this 

arrangement as well (Feldmann and Hopkins, 

1999; Schweitzer and Feldmann, 2000). Sˇtevčić 

(1983) had earlier suggested recognition of the Matutidae 

and Orithyidae and their separation from 

other Calappidae as well. We have followed Bellwood’s 

(1996) recommendations while at the same 

time not agreeing with her that the Oxystomata be 

retained. Bellwood’s rearrangement of the calappids 

is not supported by recent molecular data (S. 

Boyce, unpublished). 


Superfamily Majoidea 

Hendrickx (1995, and pers. comm.) brought our 

attention to the elevation of several majid subfamilies 

to familial rank, such as the elevation of some 

inachoid groups by Drach and Guinot (1983), who 

recognized as families the Inachidae and Inachoididae. 

We have followed Hendrickx’s recognition of 

former majid subfamilies as families. To some degree, 

our treatment (and Hendrickx’s) of the majoid 

families follows the seven subfamilies proposed by 

Griffin and Tranter (1986) in their major revision 

of the Majidae of the Indo-West Pacific. Additional 

subfamilies have been proposed by other workers, 

including Sˇtevčić (1994), who disagreed with some 

of the subdivisions proposed by Griffin and Tranter 

(1986). Diversity of the former family Majidae is 

incredibly high, and recognition or treatment of the 

majoids as a superfamily has been noted or suggested 

by many earlier workers (e.g., Guinot, 1978; 

Drach and Guinot, 1983; Sˇtevčić, 1994; Clark and 

Webber, 1991, among others). M. Wicksten (pers. 

comm.) suggests that, if we elevate some of the former 

majid subfamilies to the family level, then we 

should recognize also the family Oregoniidae 

Garth, 1958, and possibly also the Macrocheiridae 

Balss, 1929, ‘‘for consistency.’’ Indeed, Clark and 

Webber (1991) proposed recognition of both of 

these families based on a reevaluation of the larval 

features of Macrocheira and suggested that extant 

majoids be partitioned among four families: Oregoniidae, 

Macrocheiridae, Majidae, and Inachidae. 

Larval morphology indicates the distinct nature, 

and presumed monophyly, of these groups as well 

(Pohle and Marques, 2000). We have not taken that 

step here, feeling that knowledge of larval majoids 

is still rather incomplete, and we recognize here 

only the families Epialtidae, Inachidae, Inachoididae, 

Majidae, Mithracidae, Pisidae, and Tychidae. 

Concerning phylogeny among the higher (heterotrematous) 

crabs, Rice (1983:326) depicts the Majidae 

(our Majoidea) as basal to the primitive xanthid 

stock, which in turn gives rise to all other crab 

families and superfamilies. A recent study based on 

larval characters (Pohle and Marques, 2000) suggests 

that, within the Majoidea, the Oregoniinae 

clade is most basal among those majoid families (or 

subfamilies) for which larval morphology is 

known. 

Superfamily Hymenosomatoidea 

According to Guinot and Richer de Forges (1997), 

members of the family Hymenosomatidae (sole 

member of this superfamily) are thought to be 

‘‘highly advanced Heterotremata and not Thoracotremata’’ 

(Guinot and Richer de Forges, 1997: 

454, English abstract). In addition, Guinot and 

Richer de Forges (1997) revive the idea that the 

closest relatives of the hymenosomatids may lie 

among the majoid family Inachoididae. The unusual 

sperm morphology of one species of the family, 

as reported by Richer de Forges et al. (1997), 

would seem to exclude the Hymenosomatidae from 

the Thoracotremata, and even casts doubts as to 

the family’s inclusion in the Heterotremata. 

Superfamily Parthenopoidea 

The superfamily Mimilambroidea and its sole family 

Mimilambridae, both originally erected by Williams 

(1979) to contain Mimilambrus, have been 

removed following the suggestion of Ng and Rodriguez 

(1986) that Mimilambrus can be accommodated 

within the Parthenopidae. Hendrickx (1995) 

again alerted us to the fact that several former subfamilies 

of crabs (in this case, former parthenopid 

subfamilies) had been suggested to be deserving of, 

or had actually been elevated to, family rank as 

long ago as 1978 (Guinot, 1978). Although several 

authors (e.g., Hendrickx, 1999) have attributed the 

family name Daldorfiidae to M. J. Rathbun, we 

have found no indication that the taxon was recognized 

by her. Ng and Rodriguez (1986) recognized 

the suggested parthenopoid groupings of 

Guinot as valid families and first used the names 

Daldorfiidae [as Daldorfidae] and Dairidae, and we 

have attributed these families to them. We have followed 

Guinot (1978) and Ng and Rodriguez (1986) 

and recognize the families Aethridae, Dairidae, 

Daldorfiidae, and Parthenopidae within a superfamily 

Parthenopoidea, although Hendrickx (1995) 

stopped short of treating all of these as valid families. 

Superfamily Retroplumoidea 

The family Retroplumidae was given its own superfamily 

by Saint Laurent (1989), and its placement 

among the Heterotremata is based on Saint 

Laurent (1989) and Guinot (pers. comm.) 

Superfamily Cancroidea 

The family Cheiragonidae Ortmann, 1893, containing 

the genera Telmessus and Erimacrus (formerly 

treated by most workers as atelecyclids), was 

resurrected and redescribed by Sˇtevčić (1988), and 

this has been followed by Peter Ng (1998, and pers. 

comm., 1997; see also Schweitzer and Salva, 2000), 

and so we have included it here as well. 

Superfamily Portunoidea 

The freshwater family Trichodactylidae has now 

been placed in this superfamily, where it joins the 

portunids and geryonids, based primarily on a recent 

morphological analysis (Sternberg et al., 1999; 

see also below under Potamoidea). Fundamental 

differences between trichodactylids and other freshwater 

crabs were recognized by several earlier 

workers. Rodriguez (1982, 1986, 1992), Magalhães 

and Türkay (1996a–c), Sternberg (1997), 

Sternberg et al. (1999), Christoph Schubart (pers. 

comm.), and Spears et al. (2000) all acknowledge 

the unique position of the Trichodactylidae and all 

consider the family monophyletic. The hypothesis 


that the trichodactylids may represent an independent 

lineage from any of the other freshwater crab 

families and that they are descended from portunoid 

stock is supported by a number of independent 

studies using morphological data (e.g., Rodriguez, 

1982; Magalhães and Türkay, 1996a–c; 

Sternberg, 1997; Sternberg et al., 1999; and Sternberg 

and Cumberlidge, in press). Possible corroboration 

from preliminary molecular evidence (18S, 

16S, and 12S rDNA), which is admittedly based on 

only a handful of freshwater and marine crab species, 

neither strongly supports nor falsifies this relationship 

(Abele et al., 1999; Spears et al., 2000). 

Based on the totality of the evidence available to 

us, we have transferred the freshwater crab family 

Trichodactylidae to the marine superfamily Portunoidea. 

Superfamily Bythograeoidea 

Since the discovery of crabs at hydrothermal vents 

and the erection of a new superfamily and family 

(Bythograeidae) to accommodate them (Williams, 

1980), there has been much discussion concerning 

the origins and affinities of these crabs (e.g., see 

Guinot, 1988, 1990; Hessler and Martin, 1989). 

Williams (1980) noted morphological similarities 

between bythograeids and portunoids, xanthoids, 

and potamoids. Guinot (1988) argued for a recent 

derivation of the hydrothermal crab fauna. Bythograeids 

are morphologically similar to certain xanthoids, 

and there are some spermatozoal similarities 

as well (Tudge et al., 1998). It may be that, at some 

point, the bythograeids should be transferred to the 

Xanthoidea. For now, we have left them in their 

own superfamily. 

Superfamily Xanthoidea 

The former xanthids are now treated as a superfamily 

containing 11 families, a recognition of the 

group’s diversity that many workers feel is long 

overdue. The former family Xanthidae contained a 

wide variety of disparate forms and was the largest 

single family of the Decapoda, with an estimated 

130 genera and over 1,000 species (Rice, 1980; 

Martin, 1988). Manning and Holthuis (1981) list 

no fewer than 32 family and subfamily names that 

have been proposed for various assemblages within 

the family. Our elevation of the former subfamilies 

follows mostly the recommendations of Guinot 

(1977, 1978). A similar subdivision was provided 

by Serène (1984), although his treatment was restricted 

to those taxa found in the Red Sea, and so 

some xanthoid groups (such as the Panopeidae) 

were not considered by him. Serène (1984) recognized 

a Xanthoidea containing only five families 

(Xanthidae, Trapeziidae, Pilumnidae, Carpiliidae, 

and Menippidae), most with a fairly large number 

of subfamilies, some of which we are now treating 

as families. There is recent molecular evidence suggesting 

that at least some of these former subfamilies 

are indeed distinct and warrant separate family 

status (e.g., see Schubart et al., 2000b, for the Panopeidae). 

Coelho and Coelho Filhol (1993) suggested 

splitting the former Xanthidae into four 

families (Carpiliidae, Xanthidae [containing the 

subfamilies Menippinae, Platyxanthinae, Xanthinae, 

and Eucratopsinae], Eriphiidae, and Pilumnidae 

[with subfamilies Trapeziinae and Pilumninae]). 

One of the problems in elevating the various 

xanthid groups is that currently there are no published 

lists of which genera should be included in 

which family. The field worker who previously 

could place any xanthoid crab in the Xanthidae is 

now faced with the rather challenging task of wading 

through a large amount of primary literature to 

locate the appropriate family; a further problem is 

that the primary literature often does not contain 

all of this information either. Like so many other 

groups of crustaceans, the ‘‘xanthoid’’ crabs are in 

need of revision, both taxonomic and phylogenetic 

(see also Coelho and Coelho Filhol, 1993). 

Peter Ng (pers. comm.) feels that the name Eriphiidae 

MacLeay, 1838, is a senior synonym and 

should be used instead of Menippidae Ortmann, 

1893, for this family, and indeed some workers 

(e.g., Ng, 1998) have employed the name Eriphiidae. 

Serène (1984) and other workers have occasionally 

treated the Eriphiinae as a subfamily of the 

Menippidae. The family Oziidae Dana, 1852, is apparently 

a senior synonym of Menippidae as well, 

as pointed out by Holthuis (1993b), and probably 

should be used in place of Menippidae if Ozius and 

Menippe are both considered members of this 

group. However, we continue to use Menippidae in 

this case because the current (fourth) edition of the 

ICZN allows continued recognition of a name that 

is enjoying ‘‘prevailing use,’’ and in our estimation, 

replacing Menippidae with Oziidae or Eriphiidae 

would cause more confusion than maintaining use 

of Menippidae. Hendrickx (1998) elevated the former 

goneplacid subfamily Pseudorhombilinae to 

family status to accommodate six goneplacid-like 

genera; hence, our inclusion of the family Pseudorhombilidae 

Alcock, 1900, among the xanthoids. 

The Eumedonidae, a family of crabs symbiotic 

on echinoderms, has at times been recognized as a 

distinct family (Lim and Ng, 1988; Sˇtevčić et al., 

1988; and P. Ng, pers. comm.; see Chia and Ng, 

2000), and it is often placed within the Xanthoidea, 

although exactly where it belongs in relation to other 

crab families is still somewhat uncertain. Most 

workers are in agreement that early attempts to 

place it among the parthenopoids were misguided 

(e.g., see Van Dover et al., 1986; Sˇtevčić et al., 

1988; Ng and Clark, 1999, 2000) and that it is 

probably a xanthoid (Sˇtevčić et al., 1988). Daniele 

Guinot (pers. comm.), who earlier listed the family 

in its own superfamily, the Eumedonoidea Miers 

(see Guinot, 1985), now also suggests that it might 

belong in the Xanthoidea, possibly close to the Pilumnidae, 

a view shared by Van Dover et al. (1986) 

based on larval evidence. Most recently, Ng and 

Clark (1999, 2000) have arrived at the conclusion 


(based primarily on additional strong larval evidence 

that has accrued since the Van Dover et al. 

(1986) paper) that eumedonids are simply a subfamily 

of the Pilumnidae (see also Lim and Ng, 

1988). Indeed, Ng (1983) considered it a pilumnid 

subfamily, as have several other workers (reviewed 

by Sˇtevčić et al., 1988). Yet Chia and Ng (2000) 

continue to recognize the family. For now, we have 

continued to treat the Eumedonidae as a separate 

family with clear affinities to the Pilumnidae, and 

thus we have placed it with the pilumnids among 

the xanthoids. 

Recognition of Halimede as different from other 

pilumnids goes back at least to the time of Alcock 

(1898), who recognized the ‘‘alliance’’ Halimedoida. 

More recent workers (e.g., Serène, 1984:11) 

have recognized the Halimedinae as a subfamily of 

the Pilumnidae. Although Bella Galil (pers. comm.) 

feels that the genus Halimede differs sufficiently 

from other xanthoids to warrant recognition of a 

separate family, the Halimedidae, we are not aware 

of any formal treatment or description of the family 

and how it differs from the other pilumnid groupings. 

At least some workers (e.g., R. von Sternberg, 

pers. comm.) would place the Hexapodidae in the 

Thoracotremata instead of among the xanthoid 

families in the Heterotremata; von Sternberg also 

suggests, based primarily on characters of the orbits, 

that the Goneplacidae may be more closely 

related to portunids than to other xanthoid families 

(see also Sternberg and Cumberlidge, in press). 

Concerning phylogeny of xanthoid crabs, Rice 

(1980, 1983) and Martin (1988) have postulated, 

based on larval features (zoeal and megalopal), that 

the ‘‘Group III’’ larvae (e.g., Homalaspis, Ozius, 

Eriphia) might be primitive; Martin et al. (1985) 

suggested that pilumnids might be the least derived 

assemblage. Guinot (1978) felt that pilumnids and 

panopeids were more derived than the other groupings. 

In the current classification, we have simply 

listed the families alphabetically within the Xanthoidea. 

Superfamily Potamoidea 

The higher taxonomy of the freshwater crabs has 

long been in a state of disarray, and there has been 

little agreement among authors as to the number of 

superfamilies and families (e.g., see Cumberlidge, 

1999, for a review; Bott, 1970a, b; Pretzmann, 

1973; Ng, 1988, 1998; Sternberg et al., 1999; Peter 

Ng, pers. comm.; Neil Cumberlidge, pers. comm.). 

Up to 3 superfamilies and 12 families are recognized, 

depending on the author and also on how 

far back in the literature one goes. Available higher 

classifications of the freshwater crabs are based 

largely on morphological data and, until recently 

(Rodríguez, 1992; Sternberg, 1997; Sternberg et al., 

1999; Sternberg and Cumberlidge, in press), few 

have been based on cladistic analyses. Many early 

freshwater crab systematists considered all the 

world’s freshwater crabs to comprise a single 

monophyletic family, Potamidae. Others (Bott, 

1970a, b; Pretzmann, 1973) recognized 11 families 

and 3 superfamilies, arguing that the group is polyphyletic 

(or at least paraphyletic) and that similarities 

represent convergent adaptations of different 

lineages to similar habitats. Investigations over the 

past two decades (e.g., Rodríguez, 1982; Ng, 1988; 

Guinot et al., 1997; Cumberlidge, 1999) have questioned 

the validity of several families, and these 

studies continue to reveal the fundamental artificiality 

of Bott’s (1970a,b) 11-family taxonomic arrangement. 

However, in the absence of a robust 

phylogenetic study, most authors (including Bowman 

and Abele, 1982) have adopted their own variant 

of Bott’s classification (albeit reluctantly), and 

this format is followed here. 

Underlying the above taxonomic instability is the 

unresolved question of the monophyly of the freshwater 

crabs. A growing body of recent research 

(Rodríguez, 1992; Sternberg, 1997; Sternberg et al., 

1999; Sternberg and Cumberlidge, in press) has falsified 

the monophyly of the entire group and supports 

paraphyly with two main lineages. The first 

lineage includes the Trichodactylidae, which may 

be descended from some portunoid stock (see 

above under superfamily Portunoidea), and thus 

represents an independent line from any of the ‘‘potamoid’’ 

stock. The second lineage includes the rest 

of the freshwater crab families. The work of Sternberg 

et al. (1999), Cumberlidge and Sternberg 

(1999), Abele et al. (1999), Spears et al. (2000), 

and Sternberg and Cumberlidge (2000a) indicates 

that the nontrichodactylid freshwater crabs (all of 

which are heterotremes) appear to be most closely 

related to a marine crab clade that includes ocypodids, 

grapsids, and possibly pinnotherids, with 

the grapsids providing the best candidate for a sister 

taxon (an odd result in light of the fact that 

currently the potamoids are treated as heterotremes 

whereas the grapsoids are thoracotremes). The hypothesis 

suggested by Sternberg et al. (1999), that 

most families of freshwater crabs form a single 

clade composed of New and Old World lineages, is 

a departure from the traditional view of the freshwater 

crab relationships and may lead to further 

alterations of the higher classification of the group. 

Some of the more recent evidence (see especially 

Abele et al., 1999; Spears et al., 2000) seems to 

indicate that the freshwater crabs may have arrived 

via two (and possibly more) invasions. One point 

of agreement seems to be that the New World pseudothelphusids 

represent a separate clade from the 

Old World potamoids. These New World crabs 

have long been thought to represent an independent 

lineage (sometimes referred to as the Pseudothelphusoidea; 

see below) from the rest of the world’s 

freshwater crabs (see also Sternberg and Cumberlidge, 

1999). However, even this idea is somewhat 

controversial concerning whether the trichodactylids 

belong to the New World clade or represent a 

separate, independent invasion. Sternberg et al. 

(1999), citing the works of Magalhães and Türkay 


(1996a–c), Rodríguez (1982, 1986, 1992), and 

Sternberg (1997), feel that there is ‘‘strong support 

for the idea that the Pseudothelphusidae and Trichodactylidae 

each form a natural group,’’ and 

Spears and Abele (1999) have suggested that the 

pseudothelphusids are deserving of superfamily status. 

Christoph Schubart (pers. comm.) also agrees 

that the former Potamoidea is polyphyletic, especially 

as concerns the South American lineages 

(families Pseudothelphusidae and Trichodactylidae). 

Our classification is in keeping with most of 

the above views. 

Thus, excluding the trichodactylids, we recognize 

three superfamilies of freshwater crabs: Potamoidea, 

Pseudothelphusoidea, and Gercarcinucoidea. 

Within the ‘‘potamoid’’ families (superfamily Potamoidea), 

the families Sinopotamidae and Isolapotamidae 

have been removed, as both are thought 

to fall within the limits of the existing Potamidae 

(Ng, 1988; Dai et al., 1995; Dai, 1997; Dai and 

Türkay, 1997). Sternberg and Cumberlidge (1999) 

have recently recognized the monogeneric Platythelphusidae 

Colossi, 1920, as a distinct potamoid 

family (see also Cumberlidge et al., 1999; Cumberlidge, 

1999) and at the same time suggested that 

the sister group of the platythelphusids is most likely 

the East African family Deckeniidae. The Potamonautidae, 

considered to belong to the Potamidae 

by Monod (1977, 1980) and Guinot et al. (1997), 

is recognized as an independent family following 

the works of Ng (1988), Ng and Takeda (1994), 

Stewart (1997), Cumberlidge (1999), and Sternberg 

et al. (1999). 

Thus, within the superfamily Potamoidea, we 

recognize only four families here, all of them Old 

World groups: Potamidae, Potamonautidae, Deckeniidae, 

and Platythelphusidae. 

Superfamily Gecarcinucoidea 

Only two of the three families originally included 

in this superfamily by Bott (1970a, b) are recognized 

here: Gecarcinucidae and Parathelphusidae. 

The family Sundathelphusidae has been removed, 

as that family is now considered a junior synonym 

of the Parathelphusidae (Peter Ng, pers. comm; see 

also Ng and Sket, 1996; Chia and Ng, 1998). The 

family Gecarcinucidae, although recognized as being 

artificial as currently defined and in need of revision 

(N. Cumberlidge, pers. comm.; and see Cumberlidge, 

1987, 1991, 1996a, b, 1999; Cumberlidge 

and Sachs, 1991), has been retained for now. Membership 

of the family, as currently defined, is likely 

to be altered radically in the near future (N. Cumberlidge, 

pers. comm.). For example, it is possible 

that the Gecarcinucidae will be shown to be restricted 

to the Indian subcontinent, Asia, and Australasia 

(see Cumberlidge, 1999; Martin and Trautwein, 

in press), and it is not represented on the 

African continent, despite reports to the contrary 

(e.g., Bott, 1970a, b). Evidence for maintaining this 

superfamily (Gecarcinucoidea) and for separating 

these two families (Gecarcinucidae and Parathelphusidae) 

from the four families in the Potamoidea 

is weak and controversial. Nevertheless, we are recognizing 

the distinctness of the Gecarcinucidae and 

Parathelphusidae from the four potamoid families 

until further evidence becomes available. 

Superfamily Pseudothelphusoidea 

Originally established by Bott (1970a, b) to include 

two families, Pseudothelphusidae and Potamocarcinidae, 

this New World superfamily is now restricted 

to a single family. The family Potamocarcinidae 

was removed by Rodríguez (1982), and its 

species are now included among the Pseudothelphusidae 

(see Sternberg et al., 1999). The monophyly 

of the family Pseudothelphusidae appears 

well established. As noted above, Sternberg et al. 

(1999), citing the works of Magalhães and Türkay 

(1996a–c), Rodríguez (1982, 1986, 1992), and 

Sternberg (1997), feel that there is ‘‘strong support 

for the idea that the Pseudothelphusidae and Trichodactylidae 

each form a natural group.’’ Spears 

and Abele (1999) also have suggested that the pseudothelphusids 

may be deserving of superfamily status, 

and most workers are in agreement that the 

pseudothelphusids are a natural (monophyletic) 

group (T. Spears, pers. comm.; C. Schubart, pers. 

comm.; Sternberg and Cumberlidge, 1999; Sternberg 

et al., 1999). We have retained this superfamily 

and its single family Pseudothelphusidae. 

Superfamily Cryptochiroidea 

Finally in the Heterotremata, the correct name for 

the superfamily and family of the coral gall crabs 

(Cryptochiroidea and Cryptochiridae, both credited 

to Paulson) was recognized by Kropp and Manning 

(1985, 1987), who replaced the name Hapalocarcinidae 

used previously for this group. 


THORACOTREMATA 

Superfamily Pinnotheroidea 

C. Schubart (pers. comm.) believes that the Pinnotheridae 

‘‘should remain in the Thoracotremata 

based on evidence from DNA sequencing.’’ Placement 

of the pinnotherids in the Thoracotremata 

was also advocated by Sˇtevčić (1998) based on 

morphological features. Thus, the pinnotherids are 

moved to within the Thoracotremata, although the 

author of the Thoracotremata does not agree with 

this placement (Guinot, pers. comm.) and feels that 

they fit better within the Heterotremata. Within the 

Pinnotheroidea, it is possible that an additional 

family will have to be erected to accommodate the 

genera Dissodactylus and Clypeasterophilus, which 

differ morphologically (larval characters) and genetically 

from other pinnotherids (J. Cuesta, pers. 

comm.). 


Superfamily Ocypodoidea 

Within the Ocypodoidea, Guinot (pers. comm.) 

questioned the inclusion of the Retroplumidae 

among the ocypodoids and also among the thoracotremes; 

she now feels that the family Retroplumidae 

‘‘probably belongs to the Heterotremata’’ 

(where we have now placed it, in its own superfamily 

following Saint Laurent, 1989). Also within the 

Ocypodoidea, Guinot (pers. comm.) questions the 

placement of the Palicidae and suggests that they 

be listed currently as incertae sedis; Guinot and 

Bouchard (1998) treat them as members of the Heterotremata. 

C. Schubart (pers. comm.) also questions 

the placement of the palicids based on results 

of his 16S mtDNA studies (Schubart et al., 1998). 

We have left the palicids among the Ocypodoids 

pending more firm suggestions as to where they 

might belong. We have also corrected authorship of 

the family Palicidae to Bouvier from Rathbun (as 

in Bowman and Abele, 1982, and most other earlier 

treatments), following the detailed explanation 

offered in Castro’s (2000) revision of the Palicidae 

of the Indo-West Pacific. The family Camptandriidae 

Stimpson is recognized by Ng (1988). Schubart 

(pers. comm.) points out that if we recognize the 

Camptandriidae, it would be logical also to elevate 

the other three ocypodid subfamilies (Macropthalminae, 

Dotillinae, and Heloeciinae) to family level, 

and apparently there is some preliminary data to 

support this from zoeal and adult morphology (C. 

Schubart, pers. comm.). This seems especially logical 

in light of the finding of Kitaura et al. (1998) 

that the Camptrandriinae (now Camptrandriidae) 

is more closely related to the Dotillinae (based on 

molecular studies) than to any other ocypodid 

group; however, we have not yet taken that step. 

Superfamily Grapsoidea 

It has been suggested that the former grapsid subfamilies 

(especially the Varuninae) should be elevated 

to family status based on a combination of 

morphological, larval, and molecular data (Cuesta 

and Schubart, 1999; Cuesta et al., 2000; Schubart, 

2000a–c; Spivak and Cuesta, 2000; Sternberg and 

Cumberlidge, 2000b). Schubart, Cuesta, and Felder 

(in press) review some of these arguments and establish, 

on the basis of adult and larval morphology 

and molecular sequence data, the validity of the 

Glyptograpsidae (containing only Glyptograpsus 

and Platychirograpsus); they also review relationships 

among other former grapsid subfamilies. On 

the basis of these papers, we recognize as valid families 

within the Grapsidoidea the Gecarcinidae, 

Glyptograpsidae, Grapsidae, Plagusiidae, Sesarmidae, 

and Varunidae. Comparing the families Grapsidae 

(as restricted; see Schubart, Cuesta, and Felder, 

in press, and Schubart, Cuesta, and Rodríguez, 

in press) and Gecarcinidae, Cuesta and Schubart 

stated (1999: 52) that there is ‘‘not a single larval 

morphological character that consistently distinguishes 

the Gecarcinidae from the Grapsidae.’’ 

However, J. Cuesta (pers. comm.) does not feel that 

the families are closely related and instead feels that 

larvae of the Gecarcinidae are more similar to larvae 

of the Varunidae and Sesarmidae. 


We have thoroughly enjoyed the discussions with, 

and suggestions from, fellow carcinologists during 

the compilation and editing of this classification. 

Doubtless we have pleased and angered some 

workers more than others in our ‘‘final’’ arrangement. 

We have been accused of making changes 

‘‘simply for the sake of change,’’ while at the same 

time we have been accused of ‘‘classificatory paralysis’’ 

in our ‘‘unwillingness to change.’’ The classification 

has been criticized as being ‘‘nonphylogenetic,’’ 

while at the same time parts of it have been 

criticized as relying too heavily on ‘‘recent lines of 

cladistic evidence’’ (for which read molecular systematics). 

We accept all such criticisms gladly; they 

are the signs of a growing and developing field of 

CONCLUDING REMARKS 

study and of a field that is of passionate interest to 

a large number of dedicated workers. We are proud 

to be your colleagues. 

It is our sincere hope that the classification that 

follows is used primarily as a starting point for future 

research. By comparing the new classification 

with that of Bowman and Abele and seeing where 

changes have, and have not, occurred, and by reading 

the various dissenting opinions that follow (in 

Appendix I), we hope that the weaknesses inherent 

in this classification will be more readily spotted. 

We further hope that knowledge of these weaknesses 

will in turn lead to further work on the Crustacea, 

the planet’s most morphologically diverse— 

and to us, the most interesting—group of organisms. 

Contributions in Science, Number 39 Concluding Remarks � 57

CLASSIFICATION OF RECENT CRUSTACEA 

Subphylum Crustacea Brünnich, 1772 

Class Branchiopoda Latreille, 1817 

Subclass Sarsostraca Tasch, 1969 

Order Anostraca Sars, 1867 

Family Artemiidae Grochowski, 1896 

Branchinectidae Daday, 1910 

Branchipodidae Simon, 1886 

Chirocephalidae Daday, 1910 

Polyartemiidae Simon, 1886 

Streptocephalidae Daday, 1910 

Thamnocephalidae Simon, 1886 

Subclass Phyllopoda Preuss, 1951 

Order Notostraca Sars, 1867 

Family Triopsidae Keilhack, 1909 

Order Diplostraca Gerstaecker, 1866 

Suborder Laevicaudata Linder, 1945 

Family Lynceidae Baird, 1845 

Suborder Spinicaudata Linder, 1945 

Family Cyzicidae Stebbing, 1910 

Leptestheriidae Daday, 1923 

Limnadiidae Baird, 1849 

Suborder Cyclestherida Sars, 1899 

Family Cyclestheriidae Sars, 1899 

Suborder Cladocera Latreille, 1829 

Infraorder Ctenopoda Sars, 1865 

Family Holopediidae Sars, 1865 

Sididae Baird, 1850 

Infraorder Anomopoda Stebbing, 1902 

Family Bosminidae Baird, 1845 

Chydoridae Stebbing, 1902 

Daphniidae Straus, 1820 

Macrothricidae Norman & Brady, 1867 

Infraorder Onychopoda Sars, 1865 

Family Cercopagididae Mordukhai-Boltovskoi, 1968 

Podonidae Mordukhai-Boltovskoi, 1968 

Polyphemidae Baird, 1845 

Infraorder Haplopoda Sars, 1865 

Family Leptodoridae Lilljeborg, 1900 

Class Remipedia Yager, 1981 

Order Nectiopoda Schram, 1986 

Family Godzilliidae Schram, Yager & Emerson, 1986 

Speleonectidae Yager, 1981 

Class Cephalocarida Sanders, 1955 

Order Brachypoda Birshteyn, 1960 

Family Hutchinsoniellidae Sanders, 1955 

Class Maxillopoda Dahl, 1956 

Subclass Thecostraca Gruvel, 1905 

Infraclass Facetotecta Grygier, 1985 

Infraclass Ascothoracida Lacaze-Duthiers, 1880 

Order Laurida Grygier, 1987 

Family Lauridae Gruvel, 1905 

Petrarcidae Gruvel, 1905 

Synagogidae Gruvel, 1905 

Order Dendrogastrida Grygier, 1987 

Family Ascothoracidae Grygier, 1987 

Ctenosculidae Thiele, 1925 

Dendrogastridae Gruvel, 1905 

Infraclass Cirripedia Burmeister, 1834 

Superorder Acrothoracica Gruvel, 1905 

58 � Contributions in Science, Number 39 Classification of Recent Crustacea

Order Pygophora Berndt, 1907 

Family Cryptophialidae Gerstaecker, 1866 

Lithoglyptidae Aurivillius, 1892 

Order Apygophora Berndt, 1907 

Family Trypetesidae Stebbing, 1910 

Superorder Rhizocephala Müller, 1862 

Order Kentrogonida Delage, 1884 

Family Lernaeodiscidae Boschma, 1928 

Peltogastridae Lilljeborg, 1860 

Sacculinidae Lilljeborg, 1860 

Order Akentrogonida Häfele, 1911 

Family Chthamalophilidae Bocquet-Védrine, 1961 

Clistosaccidae Boschma, 1928 

Duplorbidae Høeg & Rybakov, 1992 

Mycetomorphidae Høeg & Rybakov, 1992 

Polysaccidae Lützen & Takahashi, 1996 

Thompsoniidae Høeg & Rybakov, 1992 

Superorder Thoracica Darwin, 1854 

Order Pedunculata Lamarck, 1818 

Suborder Heteralepadomorpha Newman, 1987 

Family Anelasmatidae Gruvel, 1905 

Heteralepadidae Nilsson-Cantell, 1921 

Koleolepadidae Hiro, 1933 

Malacolepadidae Hiro, 1937 

Microlepadidae Zevina, 1980 

Rhizolepadidae Zevina, 1980 

Suborder Iblomorpha Newman, 1987 

Family Iblidae Leach, 1825 

Suborder Lepadomorpha Pilsbry, 1916 

Family Lepadidae Darwin, 1852 

Oxynaspididae Gruvel, 1905 

Poecilasmatidae Annandale, 1909 

Suborder Scalpellomorpha Newman, 1987 

Family Calanticidae Zevina, 1978 

Lithotryidae Gruvel, 1905 

Pollicipedidae Leach, 1817 

Scalpellidae Pilsbry, 1907 

Order Sessilia Lamarck, 1818 

Suborder Brachylepadomorpha Withers, 1923 

Family Neobrachylepadidae Newman & Yamaguchi, 1995 

Suborder Verrucomorpha Pilsbry, 1916 

Family Neoverrucidae Newman, 1989 

Verrucidae Darwin, 1854 

Suborder Balanomorpha Pilsbry, 1916 

Superfamily Chionelasmatoidea Buckeridge, 1983 

Family Chionelasmatidae Buckeridge, 1983 

Superfamily Pachylasmatoidea Utinomi, 1968 

Family Pachylasmatidae Utinomi, 1968 

Superfamily Chthamaloidea Darwin, 1854 

Family Catophragmidae Utinomi, 1968 

Chthamalidae Darwin, 1854 

Superfamily Coronuloidea Leach, 1817 

Family Chelonibiidae Pilsbry, 1916 

Coronulidae Leach, 1817 

Platylepadidae Newman & Ross, 1976 

Superfamily Tetraclitoidea Gruvel, 1903 

Family Bathylasmatidae Newman & Ross, 1971 

Tetraclitidae Gruvel, 1903 

Superfamily Balanoidea Leach, 1817 

Family Archaeobalanidae Newman & Ross, 1976 

Balanidae Leach, 1817 

Pyrgomatidae Gray, 1825 

Contributions in Science, Number 39 Classification of Recent Crustacea � 59

Subclass Tantulocarida Boxshall & Lincoln, 1983 

Family Basipodellidae Boxshall & Lincoln, 1983 

Deoterthridae Boxshall & Lincoln, 1987 

Doryphallophoridae Huys, 1991 

Microdajidae Boxshall & Lincoln, 1987 

Onceroxenidae Huys, 1991 

Subclass Branchiura Thorell, 1864 

Order Arguloida Yamaguti, 1963 

Family Argulidae Leach, 1819 

Subclass Pentastomida Diesing, 1836 

Order Cephalobaenida Heymons, 1935 

Family Cephalobaenidae Fain, 1961 

Reighardiidae Heymons, 1935 

Order Porocephalida Heymons, 1935 

Family Armilliferidae Fain, 1961 

Diesingidae Fain, 1961 

Linguatulidae Heymons, 1935 

Porocephalidae Fain, 1961 

Sambonidae Fain, 1961 

Sebekiidae Fain, 1961 

Subtriquetridae Fain, 1961 

Subclass Mystacocarida Pennak & Zinn, 1943 

Order Mystacocaridida Pennak & Zinn, 1943 

Family Derocheilocarididae Pennak & Zinn, 1943 

Subclass Copepoda Milne-Edwards, 1840 

Infraclass Progymnoplea Lang, 1948 

Order Platycopioida Fosshagen, 1985 

Family Platycopiidae Sars, 1911 

Infraclass Neocopepoda Huys & Boxshall, 1991 

Superorder Gymnoplea Giesbrecht, 1882 

Order Calanoida Sars, 1903 

Family Acartiidae Sars, 1900 

Aetideidae Giesbrecht, 1893 

Arietellidae Sars, 1902 

Augaptilidae Sars, 1905 

Bathypontiidae Brodsky, 1950 

Boholinidae Fosshagen & Iliffe, 1989 

Calanidae Dana, 1846 

Candaciidae Giesbrecht, 1893 

Centropagidae Giesbrecht, 1893 

Clausocalanidae Giesbrecht, 1893 

Diaixidae Sars, 1902 

Diaptomidae Baird, 1850 

Discoidae Gordejeva, 1975 

Epacteriscidae Fosshagen, 1973 

Eucalanidae Giesbrecht, 1893 

Euchaetidae Giesbrecht, 1893 

Fosshageniidae Suárez-Moráles & Iliffe, 1996 

Heterorhabdidae Sars, 1902 

Hyperbionychidae Ohtsuka, Roe & Boxshall, 1993 

Lucicutiidae Sars, 1902 

Mecynoceridae Andronov, 1973 

Megacalanidae Sewell, 1947 

Mesaiokeratidae Matthews, 1961 

Metridinidae Sars, 1902 

Nullosetigeridae Soh, Ohtsuka, Imbayashi & Suh, 1999 

Paracalanidae Giesbrecht, 1893 

Parapontellidae Giesbrecht, 1893 

Parkiidae Ferrari & Markhaseva, 1996 

Phaennidae Sars, 1902 

Pontellidae Dana, 1852 

Pseudocyclopidae Giesbrecht, 1893 


Pseudocyclopiidae Sars, 1902 

Pseudodiaptomidae Sars, 1902 

Rhincalanidae Geletin, 1976 

Ridgewayiidae Wilson, 1958 

Ryocalanidae Andronov, 1974 

Scolecitrichidae Giesbrecht, 1893 

Spinocalanidae Vervoort, 1951 

Stephidae Sars, 1902 

Sulcanidae Nicholls, 1945 

Temoridae Giesbrecht, 1893 

Tharybidae Sars, 1902 

Tortanidae Sars, 1902 

Superorder Podoplea Giesbrecht, 1882 

Order Misophrioida Gurney, 1933 

Family Misophriidae Brady, 1878 

Palpophriidae Boxshall & Jaume, 2000 

Speleophriidae Boxshall & Jaume, 2000 

Order Cyclopoida Burmeister, 1834 

Family Archinotodelphyidae Lang, 1949 

Ascidicolidae Thorell, 1860 

Buproridae Thorell, 1859 

Chordeumiidae Boxshall, 1988 

Cucumaricolidae Bouligand & Delamare-Deboutteville, 1959 

Cyclopidae Dana, 1846 

Cyclopinidae Sars, 1913 

Fratiidae Ho, Conradi & López-González, 1998 

Lernaeidae Cobbold, 1879 

Mantridae Leigh-Sharpe, 1934 

Notodelphyidae Dana, 1852 

Oithonidae Dana, 1852 

Ozmanidae Ho & Thatcher, 1989 

Speleoithonidae da Rocha & Iliffe, 1991 

Thaumatopsyllidae Sars, 1913 

Order Gelyelloida Huys, 1988 

Family Gelyellidae Rouch & Lescher-Moutoué, 1977 

Order Mormonilloida Boxshall, 1979 

Family Mormonillidae Giesbrecht, 1893 

Order Harpacticoida Sars, 1903 

Family Adenopleurellidae Huys, 1990 

Aegisthidae Giesbrecht, 1893 

Ambunguipedidae Huys, 1990 

Ameiridae Monard, 1927 

Ancorabolidae Sars, 1909 

Argestidae Por, 1986 

Balaenophilidae Sars, 1910 

Cancrincolidae Fiers, 1990 

Canthocamptidae Sars, 1906 

Canuellidae Lang, 1944 

Cerviniidae Sars, 1903 

Chappuisiidae Chappuis, 1940 

Cletodidae Scott, 1905 

Cletopsyllidae Huys & Williams, 1989 

Clytemnestridae Scott, 1909 

Cristacoxidae Huys, 1990 

Cylindropsyllidae Sars, 1909 

Darcythompsoniidae Lang, 1936 

Diosaccidae Sars, 1906 

Ectinosomatidae Sars, 1903 

Euterpinidae Brian, 1921 

Hamondiidae Huys, 1990 

Harpacticidae Dana, 1846 

Huntemanniidae Por, 1986 


Laophontidae Scott, 1905 

Laophontopsidae Huys & Willems, 1989 

Latiremidae Bozˇić, 1969 

Leptastacidae Lang, 1948 

Leptopontiidae Lang, 1948 

Longipediidae Sars, 1903 

Louriniidae Monard, 1927 

Metidae Sars, 1910 

Miraciidae Dana, 1846 

Neobradyidae Oloffson, 1917 

Normanellidae Lang, 1944 

Novocriniidae Huys & Iliffe, 1998 

Orthopsyllidae Huys, 1990 

Paramesochridae Lang, 1944 

Parastenheliidae Lang, 1936 

Parastenocarididae Chappuis, 1933 

Peltidiidae Sars, 1904 

Phyllognathopodidae Gurney, 1932 

Porcellidiidae Boeck, 1865 

Pseudotachidiidae Lang, 1936 

Rhizothricidae Por, 1986 

Rotundiclipeidae Huys, 1988 

Styracothoracidae Huys, 1993 

Superornatiremidae Huys, 1997 

Tachidiidae Boeck, 1865 

Tegastidae Sars, 1904 

Tetragonicipitidae Lang, 1944 

Thalestridae Sars, 1905 

Thompsonulidae Lang, 1944 

Tisbidae Stebbing, 1910 

Order Poecilostomatoida Thorell, 1859 

Family Anchimolgidae Humes & Boxshall, 1996 

Anomoclausiidae Gotto, 1964 

Antheacheridae Sars, 1870 

Anthessiidae Humes, 1986 

Bomolochidae Sumpf, 1871 

Catiniidae Bocquet & Stock, 1957 

Chitonophilidae Avdeev & Sirenko, 1991 

Chondracanthidae Milne Edwards, 1840 

Clausidiidae Embleton, 1901 

Clausiidae Giesbrecht, 1895 

Corallovexiidae Stock, 1975 

Corycaeidae Dana, 1852 

Echiurophilidae Delamare-Deboutteville & Nunes-Ruivo, 1955 

Entobiidae Ho, 1984 

Erebonasteridae Humes, 1987 

Ergasilidae von Nordmann, 1832 

Eunicicolidae Sars, 1918 

Gastrodelphyidae List, 1889 

Herpyllobiidae Hansen, 1892 

Intramolgidae Marchenkov & Boxshall, 1995 

Kelleriidae Humes & Boxshall, 1996 

Lamippidae Joliet, 1882 

Lernaeosoleidae Yamaguti, 1963 

Lichomolgidae Kossmann, 1877 

Lubbockiidae Huys & Böttger-Schnack, 1997 

Macrochironidae Humes & Boxshall, 1996 

Mesoglicolidae de Zulueta, 1911 

Micrallectidae Huys, 2001 

Myicolidae Yamaguti, 1936 

Mytilicolidae Bocquet & Stock, 1957 

Nereicolidae Claus, 1875 


Nucellicolidae Lamb, Boxshall, Mill & Grahame, 1996 

Octopicolidae Humes & Boxshall, 1996 

Oncaeidae Giesbrecht, 1893 

Paralubbockiidae Boxshall & Huys, 1989 

Pharodidae Illg, 1948 

Philichthyidae Vogt, 1877 

Philoblennidae Izawa, 1976 

Phyllodicolidae Delamare-Deboutteville & Laubier, 1961 

Polyankylidae Ho & Kim, 1997 

Pseudanthessiidae Humes & Stock, 1972 

Rhynchomolgidae Humes & Stock, 1972 

Sabelliphilidae Gurney, 1927 

Saccopsidae Lützen, 1964 

Sapphirinidae Thorell, 1860 

Serpulidicolidae Stock, 1979 

Shiinoidae Cressey, 1975 

Spiophanicolidae Ho, 1984 

Splanchnotrophidae Norman & Scott, 1906 

Synapticolidae Humes & Boxshall, 1996 

Synaptiphilidae Bocquet, 1953 

Taeniacanthidae Wilson, 1911 

Tegobomolochidae Avdeev, 1978 

Telsidae Ho, 1967 

Thamnomolgidae Humes & Boxshall, 1996 

Tuccidae Vervoort, 1962 

Urocopiidae Humes & Stock, 1972 

Vahiniidae Humes, 1967 

Ventriculinidae Leigh-Sharpe, 1934 

Xarifiidae Humes, 1960 

Xenocoelomatidae Bresciani & Lützen, 1966 

Order Siphonostomatoida Thorell, 1859 

Family Archidactylinidae Izawa, 1996 

Artotrogidae Brady, 1880 

Asterocheridae Giesbrecht, 1899 

Brychiopontiidae Humes, 1974 

Caligidae Burmeister, 1834 

Calverocheridae Stock, 1968 

Cancerillidae Giesbrecht, 1897 

Cecropidae Dana, 1849 

Codobidae Boxshall & Ohtsuka, 2001 

Coralliomyzontidae Humes & Stock, 1991 

Dichelesthiidae Milne Edwards, 1840 

Dichelinidae Boxshall & Ohtsuka, 2001 

Dinopontiidae Murnane, 1967 

Dirivultidae Humes & Dojiri, 1981 

Dissonidae Yamaguti, 1963 

Ecbathyriontidae Humes, 1987 

Entomolepididae Brady, 1899 

Eudactylinidae Wilson, 1922 

Euryphoridae Wilson, 1905 

Hatschekiidae Kabata, 1979 

Hyponeoidae Heegaard, 1962 

Kroyeriidae Kabata, 1979 

Lernaeopodidae Milne Edwards, 1840 

Lernanthropidae Kabata, 1979 

Megapontiidae Heptner, 1968 

Micropontiidae Gooding, 1957 

Nanaspididae Humes & Cressey, 1959 

Nicothoidae Dana, 1849 

Pandaridae Milne Edwards, 1840 

Pennellidae Burmeister, 1834 

Pontoeciellidae Giesbrecht, 1895 


Pseudocycnidae Wilson, 1922 

Rataniidae Giesbrecht, 1897 

Scottomyzontidae Ivanenko, Ferrari, & Smurov, 2001 

Sphyriidae Wilson, 1919 

Sponginticolidae Topsent, 1928 

Spongiocnizontidae Stock & Kleeton, 1964 

Stellicomitidae Humes & Cressey, 1958 

Tanypleuridae Kabata, 1969 

Trebiidae Wilson, 1905 

Order Monstrilloida Sars, 1901 

Family Monstrillidae Dana, 1849 

Class Ostracoda Latreille, 1802 

Subclass Myodocopa Sars, 1866 

Order Myodocopida Sars, 1866 

Suborder Myodocopina Sars, 1866 

Superfamily Cypridinoidea Baird, 1850 

Family Cypridinidae Baird, 1850 

Superfamily Cylindroleberidoidea Müller, 1906 

Family Cylindroleberididae Müller, 1906 

Superfamily Sarsielloidea Brady & Norman, 1896 

Family Philomedidae Müller, 1906 

Rutidermatidae Brady & Norman, 1896 

Sarsiellidae Brady & Norman, 1896 

Order Halocyprida Dana, 1853 

Suborder Cladocopina Sars, 1865 

Superfamily Polycopoidea Sars, 1865 

Family Polycopidae Sars, 1865 

Suborder Halocypridina Dana, 1853 

Superfamily Halocypridoidea Dana, 1853 

Family Halocyprididae Dana, 1853 

Superfamily Thaumatocypridoidea Müller, 1906 

Family Thaumatocyprididae Müller, 1906 

Subclass Podocopa Müller, 1894 

Order Platycopida Sars, 1866 

Family Cytherellidae Sars, 1866 

Punciidae Hornibrook, 1949 

Order Podocopida Sars, 1866 

Suborder Bairdiocopina Sars, 1865 

Superfamily Bairdioidea Sars, 1865 

Family Bairdiidae Sars, 1865 

Bythocyprididae Maddocks, 1969 

Suborder Cytherocopina Baird, 1850 

Superfamily Cytheroidea Baird, 1850 

Family Bythocytheridae Sars, 1866 

Cytheridae Baird, 1850 

Cytherideidae Sars, 1925 

Cytheromatidae Elofson, 1939 

Cytheruridae Müller, 1894 

Entocytheridae Hoff, 1942 

Eucytheridae Puri, 1954 

Hemicytheridae Puri, 1953 

Kliellidae Schäfer, 1945 

Krithidae Mandelstam, 1958 

Leptocytheridae Hanai, 1957 

Loxoconchidae Sars, 1925 

Microcytheridae Klie, 1938 

Neocytherideidae Puri, 1957 

Paradoxostomatidae Brady & Norman, 1889 

Pectocytheridae Hanai, 1957 

Protocytheridae Ljubimova, 1956 

Psammocytheridae Klie, 1938 

Schizocytheridae Howe, 1961 


Terrestricytheridae Schornikov, 1969 

Thaerocytheridae Hazel, 1967 

Trachyleberididae Sylvester-Bradley, 1948 

Xestoleberididae Sars, 1928 

Suborder Darwinulocopina Sohn, 1988 

Superfamily Darwinuloidea Brady & Norman, 1889 

Family Darwinulidae Brady & Norman, 1889 

Suborder Cypridocopina Jones, 1901 

Superfamily Cypridoidea Baird, 1845 

Family Candonidae Kaufmann, 1900 

Cyprididae Baird, 1845 

Ilyocyprididae Kaufmann, 1900 

Notodromadidae Kaufmann, 1900 

Superfamily Macrocypridoidea Müller, 1912 

Family Macrocyprididae Müller, 1912 

Superfamily Pontocypridoidea Müller, 1894 

Family Pontocyprididae Müller, 1894 

Suborder Sigilliocopina Martens, 1992 

Superfamily Sigillioidea Mandelstam, 1960 

Family Sigilliidae Mandelstam, 1960 

Class Malacostraca Latreille, 1802 

Subclass Phyllocarida Packard, 1879 

Order Leptostraca Claus, 1880 

Family Nebaliidae Samouelle, 1819 

Nebaliopsidae Hessler, 1984 

Paranebaliidae Walker-Smith & Poore, 2001 

Subclass Hoplocarida Calman, 1904 

Order Stomatopoda Latreille, 1817 

Suborder Unipeltata Latreille, 1825 

Superfamily Bathysquilloidea Manning, 1967 

Family Bathysquillidae Manning, 1967 

Indosquillidae Manning, 1995 

Superfamily Gonodactyloidea Giesbrecht, 1910 

Family Alainosquillidae Moosa, 1991 

Hemisquillidae Manning, 1980 

Gonodactylidae Giesbrecht, 1910 

Odontodactylidae Manning, 1980 

Protosquillidae Manning, 1980 

Pseudosquillidae Manning, 1977 

Takuidae Manning, 1995 

Superfamily Erythrosquilloidea Manning & Bruce, 1984 

Family Erythrosquillidae Manning & Bruce, 1984 

Superfamily Lysiosquilloidea Giesbrecht, 1910 

Family Coronididae Manning, 1980 

Lysiosquillidae Giesbrecht, 1910 

Nannosquillidae Manning, 1980 

Tetrasquillidae Manning & Camp, 1993 

Superfamily Squilloidea Latreille, 1802 

Family Squillidae Latreille, 1802 

Superfamily Eurysquilloidea Ahyong & Harling, 2000 

Family Eurysquillidae Manning, 1977 

Superfamily Parasquilloidea Ahyong & Harling, 2000 

Family Parasquillidae Manning, 1995 

Subclass Eumalacostraca Grobben, 1892 

Superorder Syncarida Packard, 1885 

Order Bathynellacea Chappuis, 1915 

Family Bathynellidae Chappuis, 1915 

Parabathynellidae Noodt, 1965 

Order Anaspidacea Calman, 1904 

Family Anaspididae Thomson, 1893 

Koonungidae Sayce, 1908 

Psammaspididae Schminke, 1974 


Stygocarididae Noodt, 1963 

Superorder Peracarida Calman, 1904 

Order Spelaeogriphacea Gordon, 1957 

Family Spelaeogriphidae Gordon, 1957 

Order Thermosbaenacea Monod, 1927 

Family Halosbaenidae Monod & Cals, 1988 

Monodellidae Taramelli, 1954 

Thermosbaenidae Monod, 1927 

Tulumellidae Wagner, 1994 

Order Lophogastrida Sars, 1870 

Family Eucopiidae Sars, 1885 

Lophogastridae Sars, 1870 

Order Mysida Haworth, 1825 

Family Lepidomysidae Clarke, 1961 

Mysidae Haworth, 1825 

Petalophthalmidae Czerniavsky, 1882 

Stygiomysidae Caroli, 1937 

Order Mictacea Bowman, Garner, Hessler, Iliffe & Sanders, 1985 

Family Hirsutiidae Sanders, Hessler & Garner, 1985 

Mictocarididae Bowman & Iliffe, 1985 

Order Amphipoda Latreille, 1816 

Suborder Gammaridea Latreille, 1802 

Family Acanthogammaridae Garjajeff, 1901 

Acanthonotozomellidae Coleman & Barnard, 1991 

Allocrangonyctidae Holsinger, 1989 

Amathillopsidae Pirlot, 1934 

Ampeliscidae Costa, 1857 

Amphilochidae Boeck, 1871 

Ampithoidae Stebbing, 1899 

Anamixidae Stebbing, 1897 

Anisogammaridae Bousfield, 1977 

Aoridae Walker, 1908 

Argissidae Walker, 1904 

Aristiidae Lowry & Stoddart, 1997 

Artesiidae Holsinger, 1980 

Bateidae Stebbing, 1906 

Biancolinidae Barnard, 1972 

Bogidiellidae Hertzog, 1936 

Bolttsiidae Barnard & Karaman, 1987 

Calliopidae Sars, 1893 

Carangoliopsidae Bousfield, 1977 

Cardenioidae Barnard & Karaman, 1987 

Caspicolidae Birstein, 1945 

Ceinidae Barnard, 1972 

Cheidae Thurston, 1982 

Cheluridae Allman, 1847 

Clarenciidae Barnard & Karaman, 1987 

Colomastigidae Stebbing, 1899 

Condukiidae Barnard & Drummond, 1982 

Corophiidae Leach, 1814 

Crangonyctidae Bousfield, 1973 

Cressidae Stebbing, 1899 

Cyphocarididae Lowry & Stoddart, 1997 

Cyproideidae Barnard, 1974 

Dexaminidae Leach, 1814 

Didymocheliidae Bellan-Santini & Ledoyer, 1986 

Dikwidae Coleman & Barnard, 1991 

Dogielinotidae Gurjanova, 1953 

Dulichiidae Dana, 1849 

Endevouridae Lowry & Stoddart, 1997 

Eophliantidae Sheard, 1936 

Epimeriidae Boeck, 1871 


Eusiridae Stebbing, 1888 

Exoedicerotidae Barnard & Drummond, 1982 

Gammaracanthidae Bousfield, 1989 

Gammarellidae Bousfield, 1977 

Gammaridae Latreille, 1802 

Gammaroporeiidae Bousfield, 1979 

Hadziidae Karaman, 1943 

Haustoriidae Stebbing, 1906 

Hyalellidae Bulycheva, 1957 

Hyalidae Bulycheva, 1957 

Hyperiopsidae Bovallius, 1886 

Iciliidae Dana, 1849 

Ipanemidae Barnard & Thomas, 1988 

Iphimediidae Boeck, 1871 

Isaeidae Dana, 1853 

Ischyroceridae Stebbing, 1899 

Kuriidae Walker & Scott, 1903 

Laphystiidae Sars, 1893 

Laphystiopsidae Stebbing, 1899 

Lepechinellidae Schellenberg, 1926 

Leucothoidae Dana, 1852 

Liljeborgiidae Stebbing, 1899 

Lysianassidae Dana, 1849 

Macrohectopidae Sowinsky, 1915 

Maxillipiidae Ledoyer, 1973 

Megaluropidae Thomas & Barnard, 1986 

Melitidae Bousfield, 1973 

Melphidippidae Stebbing, 1899 

Mesogammaridae Bousfield, 1977 

Metacrangonyctidae Boutin & Missouli, 1988 

Micruropidae Kamaltynov, 1999 

Najnidae Barnard, 1972 

Neomegamphopidae Myers, 1981 

Neoniphargidae Bousfield, 1977 

Nihotungidae Barnard, 1972 

Niphargidae Bousfield, 1977 

Ochlesidae Stebbing, 1910 

Odiidae Coleman & Barnard, 1991 

Oedicerotidae Lilljeborg, 1865 

Opisidae Lowry & Stoddart, 1995 

Pachyschesidae Kamaltynov, 1999 

Pagetinidae Barnard, 1931 

Paracalliopidae Barnard & Karaman, 1982 

Paracrangonyctidae Bousfield, 1982 

Paraleptamphopidae Bousfield, 1983 

Paramelitidae Bousfield, 1977 

Pardaliscidae Boeck, 1871 

Perthiidae Williams & Barnard, 1988 

Phliantidae Stebbing, 1899 

Phoxocephalidae Sars, 1891 

Phoxocephalopsidae Barnard & Drummond, 1982 

Phreatogammaridae Bousfield, 1982 

Platyischnopidae Barnard & Drummond, 1979 

Pleustidae Buchholz, 1874 

Plioplateidae Barnard, 1978 

Podoceridae Leach, 1814 

Podoprionidae Lowry & Stoddart, 1996 

Pontogammaridae Bousfield, 1977 

Pontoporeiidae Dana, 1853 

Priscomilitaridae Hirayama, 1988 

Pseudamphilochidae Schellenberg, 1931 

Pseudocrangonyctidae Holsinger, 1989 


Salentinellidae Bousfield, 1977 

Scopelocheiridae Lowry & Stoddart, 1997 

Sebidae Walker, 1908 

Sinurothoidae Ren, 1999 

Stegocephalidae Dana, 1853 

Stenothoidae Boeck, 1871 

Sternophysingidae Holsinger, 1992 

Stilipedidae Holmes, 1908 

Synopiidae Dana, 1853 

Talitridae Rafinesque, 1815 

Temnophliantidae Griffiths, 1975 

Trischizostomatidae Lilljeborg, 1865 

Tulearidae Ledoyer, 1979 

Typhlogammaridae, Bousfield, 1977 

Uristidae Hurley, 1963 

Urohaustoriidae Barnard & Drummond, 1982 

Urothoidae Bousfield, 1978 

Valettidae Stebbing, 1888 

Vicmusiidae Just, 1990 

Vitjazianidae Birstein & Vinogradov, 1955 

Wandinidae Lowry & Stoddart, 1990 

Zobrachoidae Barnard & Drummond, 1982 

Suborder Caprellidea Leach, 1814 

Infraorder Caprellida Leach, 1814 

Superfamily Caprelloidea Leach, 1814 

Family Caprellidae Leach, 1814 

Caprellinoididae Laubitz, 1993 

Caprogammaridae Kudrjaschov & Vassilenko, 1966 

Paracercopidae Vassilenko, 1968 

Pariambidae Laubitz, 1993 

Protellidae McCain, 1970 

Superfamily Phtisicoidea Vassilenko, 1968 

Family Phtisicidae Vassilenko, 1968 

Infraorder Cyamida Rafinesque, 1815 

Family Cyamidae Rafinesque, 1815 

Suborder Hyperiidea Milne Edwards, 1830 

Infraorder Physosomata Pirlot, 1929 

Superfamily Scinoidea Stebbing, 1888 

Family Archaeoscinidae Stebbing, 1904 

Mimonectidae Bovallius, 1885 

Proscinidae Pirlot, 1933 

Scinidae Stebbing, 1888 

Superfamily Lanceoloidea Bovallius, 1887 

Family Chuneolidae Woltereck, 1909 

Lanceolidae Bovallius, 1887 

Microphasmatidae Stephensen & Pirlot, 1931 

Infraorder Physocephalata Bowman & Gruner, 1973 

Superfamily Vibilioidea Dana, 1853 

Family Cystisomatidae Willemoes-Suhm, 1875 

Paraphronimidae Bovallius, 1887 

Vibiliidae Dana, 1853 

Superfamily Phronimoidea Rafinesque, 1815 

Family Dairellidae Bovallius, 1887 

Hyperiidae Dana, 1853 

Phronimidae Rafinesque, 1815 

Phrosinidae Dana, 1853 

Superfamily Lycaeopsoidea Chevreux, 1913 

Family Lycaeopsidae Chevreux, 1913 

Superfamily Platysceloidea Bate, 1862 

Family Anapronoidae Bowman & Gruner, 1973 

Lycaeidae Claus, 1879 

Oxycephalidae Dana, 1853 


Parascelidae Bate, 1862 

Platyscelidae Bate, 1862 

Pronoidae Dana, 1853 

Suborder Ingolfiellidea Hansen, 1903 

Family Ingolfiellidae Hansen, 1903 

Metaingolfiellidae Ruffo, 1969 

Order Isopoda Latreille, 1817 

Suborder Phreatoicidea Stebbing, 1893 

Family Amphisopodidae Nicholls, 1943 

Nichollsiidae Tiwari, 1955 

Phreatoicidae Chilton, 1891 

Suborder Anthuridea Monod, 1922 

Family Antheluridae Poore & Lew Ton, 1988 

Anthuridae Leach, 1814 

Expanathuridae Poore, 2001 

Hyssuridae Wägele, 1981 

Leptanthuridae Poore, 2001 

Paranthuridae Menzies & Glynn, 1968 

Suborder Microcerberidea Lang, 1961 

Family Atlantasellidae Sket, 1980 

Microcerberidae Karaman, 1933 

Suborder Flabellifera Sars, 1882 

Family Aegidae White, 1850 

Ancinidae Dana, 1852 

Anuropidae Stebbing, 1893 

Bathynataliidae Kensley, 1978 

Cirolanidae Dana, 1852 

Corallanidae Hansen, 1890 

Cymothoidae Leach, 1814 

Gnathiidae Leach, 1814 

Hadromastacidae Bruce & Müller, 1991 

Keuphyliidae Bruce, 1980 

Limnoriidae White, 1850 

Phoratopodidae Hale, 1925 

Plakarthriidae Hansen, 1905 

Protognathiidae Wägele & Brandt, 1988 

Serolidae Dana, 1852 

Sphaeromatidae Latreille, 1825 

Tecticepitidae Iverson, 1982 

Tridentellidae Bruce, 1984 

Suborder Asellota Latreille, 1802 

Superfamily Aselloidea Latreille, 1802 

Family Asellidae Latreille, 1802 

Stenasellidae Dudich, 1924 

Superfamily Stenetrioidea Hansen, 1905 

Family Pseudojaniridae Wilson, 1986 

Stenetriidae Hansen, 1905 

Superfamily Janiroidea Sars, 1897 

Family Acanthaspidiidae Menzies, 1962 

Dendrotiidae Vanhöffen, 1914 

Desmosomatidae Sars, 1899 

Echinothambematidae Menzies, 1956 

Haplomunnidae Wilson, 1976 

Haploniscidae Hansen, 1916 

Ischnomesidae Hansen, 1916 

Janirellidae Menzies, 1956 

Janiridae Sars, 1897 

Joeropsididae Nordenstam, 1933 

Katianiridae Svavarsson, 1987 

Macrostylidae Hansen, 1916 

Mesosignidae Schultz, 1969 

Microparasellidae Karaman, 1933 


Mictosomatidae Wolff, 1965 

Munnidae Sars, 1897 

Munnopsididae Sars, 1869 

Nannoniscidae Hansen, 1916 

Paramunnidae Vanhöffen, 1914 

Pleurocopidae Fresi & Schiecke, 1972 

Santiidae Wilson, 1987 

Thambematidae Stebbing, 1913 

Superfamily Gnathostenetroidoidea Kussakin, 1967 

Family Gnathostenetroididae Kussakin, 1967 

Protojaniridae Fresi, Idato & Scipione, 1980 

Vermectiadidae Just & Poore, 1992 

Suborder Calabozoida Van Lieshout, 1983 

Family Calabozoidae Van Lieshout, 1983 

Suborder Valvifera Sars, 1882 

Family Antarcturidae Poore, 2001 

Arcturidae Dana, 1849 

Arcturididae Poore, 2001 

Austrarcturellidae Poore & Bardsley, 1992 

Chaetiliidae Dana, 1849 

Holidoteidae Wägele, 1989 

Holognathidae Thomson, 1904 

Idoteidae Samouelle, 1819 

Pseudidotheidae Ohlin, 1901 

Rectarcturidae Poore, 2001 

Xenarcturidae Sheppard, 1957 

Suborder Epicaridea Latreille, 1831 

Superfamily Bopyroidea Rafinesque, 1815 

Family Bopyridae Rafinesque, 1815 

Dajidae Giard & Bonnier, 1887 

Entoniscidae Kossmann, 1881 

Superfamily Cryptoniscoidea Kossmann, 1880 

Family Asconiscidae Bonnier, 1900 

Cabiropidae Giard & Bonnier, 1887 

Crinoniscidae Bonnier, 1900 

Cryptoniscidae Kossmann, 1880 

Cyproniscidae Bonnier, 1900 

Fabidae Danforth, 1963 

Hemioniscidae Bonnier, 1900 

Podasconidae Bonnier, 1900 

Suborder Oniscidea Latreille, 1802 

Family Dubioniscidae Schultz, 1995 

Helelidae Ferrara, 1977 

Irmaosidae Ferrara & Taiti, 1983 

Pseudarmadillidae Vandel, 1973 

Scleropactidae Verhoeff, 1938 

Infraorder Tylomorpha Vandel, 1943 

Family Tylidae Dana, 1852 

Infraorder Ligiamorpha Vandel, 1943 

Section Diplocheta Vandel, 1957 

Family Ligiidae Leach, 1814 

Mesoniscidae Verhoeff, 1908 

Section Synocheta Legrand, 1946 

Superfamily Trichoniscoidea Sars, 1899 

Family Buddelundiellidae Verhoeff, 1930 

Trichoniscidae Sars, 1899 

Superfamily Styloniscoidea Vandel, 1952 

Family Schoebliidae Verhoeff, 1938 

Styloniscidae Vandel, 1952 

Titaniidae Verhoeff, 1938 

Tunanoniscidae Borutskii, 1969 

Section Crinocheta Legrand, 1946 


Superfamily Oniscoidea Latreille, 1802 

Family Bathytropidae Vandel, 1952 

Berytoniscidae Vandel, 1973 

Detonidae Budde-Lund, 1906 

Halophilosciidae Verhoeff, 1908 

Olibrinidae Vandel, 1973 

Oniscidae Latreille, 1802 

Philosciidae Kinahan, 1857 

Platyarthridae Vandel, 1946 

Pudeoniscidae Lemos de Castro, 1973 

Rhyscotidae Budde-Lund, 1908 

Scyphacidae Dana, 1852 

Speleoniscidae Vandel, 1948 

Sphaeroniscidae Vandel, 1964 

Stenoniscidae Budde-Lund, 1904 

Tendosphaeridae Verhoeff, 1930 

Superfamily Armadilloidea Brandt, 1831 

Family Actaeciidae Vandel, 1952 

Armadillidae Brandt, 1831 

Armadillidiidae Brandt, 1833 

Atlantidiidae Arcangeli, 1954 

Balloniscidae Vandel, 1963 

Cylisticidae Verhoeff, 1949 

Eubelidae Budde-Lund, 1904 

Periscyphicidae Ferrara, 1973 

Porcellionidae Brandt, 1831 

Trachelipodidae Strouhal, 1953 

Order Tanaidacea Dana, 1849 

Suborder Tanaidomorpha Sieg, 1980 

Superfamily Tanaoidea Dana, 1849 

Family Tanaidae Dana, 1849 

Superfamily Paratanaoidea Lang, 1949 

Family Anarthruridae Lang, 1971 

Leptochelidae Lang, 1973 

Nototanaidae Sieg, 1976 

Paratanaidae Lang, 1949 

Pseudotanaidae Sieg, 1976 

Pseudozeuxidae Sieg, 1982 

Typhlotanaidae Sieg, 1986 

Suborder Neotanaidomorpha Sieg, 1980 

Family Neotanaidae Lang, 1956 

Suborder Apseudomorpha Sieg, 1980 

Superfamily Apseudoidea Leach, 1814 

Family Anuropodidae Băcescu, 1980 

Apseudellidae Gutu, 1972 

Apseudidae Leach, 1814 

Gigantapseudidae Kudinova-Pasternak, 1978 

Kalliapseudidae Lang, 1956 

Metapseudidae Lang, 1970 

Pagurapseudidae Lang, 1970 

Parapseudidae Gutu, 1981 

Sphyrapidae Gutu, 1980 

Tanapseudidae Băcescu, 1978 

Tanzanapseudidae Băcescu, 1975 

Whiteleggiidae Gutu, 1972 

Order Cumacea Krøyer, 1846 

Family Bodotriidae Scott, 1901 

Ceratocumatidae Calman, 1905 

Diastylidae Bate, 1856 

Gynodiastylidae Stebbing, 1912 

Lampropidae Sars, 1878 

Leuconidae Sars, 1878 


Nannastacidae Bate, 1866 

Pseudocumatidae Sars, 1878 

Superorder Eucarida Calman, 1904 

Order Euphausiacea Dana, 1852 

Family Bentheuphausiidae Colosi, 1917 

Euphausiidae Dana, 1852 

Order Amphionidacea Williamson, 1973 

Family Amphionididae Holthuis, 1955 

Order Decapoda Latreille, 1802 

Suborder Dendrobranchiata Bate, 1888 

Superfamily Penaeoidea Rafinesque, 1815 

Family Aristeidae Wood-Mason, 1891 

Benthesicymidae Wood-Mason, 1891 

Penaeidae Rafinesque, 1815 

Sicyoniidae Ortmann, 1898 

Solenoceridae Wood-Mason, 1891 

Superfamily Sergestoidea Dana, 1852 

Family Luciferidae de Haan, 1849 

Sergestidae Dana, 1852 

Suborder Pleocyemata Burkenroad, 1963 

Infraorder Stenopodidea Claus, 1872 

Family Spongicolidae Schram, 1986 

Stenopodidae Claus, 1872 

Infraorder Caridea Dana, 1852 

Superfamily Procaridoidea Chace & Manning, 1972 

Family Procarididae Chace & Manning, 1972 

Superfamily Galatheacaridoidea Vereshchaka, 1997 

Family Galatheacarididae Vereshchaka, 1997 

Superfamily Pasiphaeoidea Dana, 1852 

Family Pasiphaeidae Dana, 1852 

Superfamily Oplophoroidea Dana, 1852 

Family Oplophoridae Dana, 1852 

Superfamily Atyoidea de Haan, 1849 

Family Atyidae de Haan, 1849 

Superfamily Bresilioidea Calman, 1896 

Family Agostocarididae Hart & Manning, 1986 

Alvinocarididae Christoffersen, 1986 

Bresiliidae Calman, 1896 

Disciadidae Rathbun, 1902 

Mirocarididae Vereshchaka, 1997 

Superfamily Nematocarcinoidea Smith, 1884 

Family Eugonatonotidae Chace, 1937 

Nematocarcinidae Smith, 1884 

Rhynchocinetidae Ortmann, 1890 

Xiphocarididae Ortmann, 1895 

Superfamily Psalidopodoidea Wood-Mason & Alcock, 1892 

Family Psalidopodidae Wood-Mason & Alcock, 1892 

Superfamily Stylodactyloidea Bate, 1888 

Family Stylodactylidae Bate, 1888 

Superfamily Campylonotoidea Sollaud, 1913 

Family Bathypalaemonellidae de Saint Laurent, 1985 

Campylonotidae Sollaud, 1913 

Superfamily Palaemonoidea Rafinesque, 1815 

Family Anchistioididae Borradaile, 1915 

Desmocarididae Borradaile, 1915 

Euryrhynchidae Holthuis, 1950 

Gnathophyllidae Dana, 1852 

Hymenoceridae Ortmann, 1890 

Kakaducarididae Bruce, 1993 

Palaemonidae Rafinesque, 1815 

Typhlocarididae Annandale & Kemp, 1913 

Superfamily Alpheoidea Rafinesque, 1815 


Family Alpheidae Rafinesque, 1815 

Barbouriidae Christoffersen, 1987 

Hippolytidae Dana, 1852 

Ogyrididae Holthuis, 1955 

Superfamily Processoidea Ortmann, 1890 

Family Processidae Ortmann, 1890 

Superfamily Pandaloidea Haworth, 1825 

Family Pandalidae Haworth, 1825 

Thalassocarididae Bate, 1888 

Superfamily Physetocaridoidea Chace, 1940 

Family Physetocarididae Chace, 1940 

Superfamily Crangonoidea Haworth, 1825 

Family Crangonidae Haworth, 1825 

Glyphocrangonidae Smith, 1884 

Infraorder Astacidea Latreille, 1802 

Superfamily Glypheoidea Winkler, 1883 

Family Glypheidae Winkler, 1883 

Superfamily Enoplometopoidea de Saint Laurent, 1988 

Family Enoplometopidae de Saint Laurent, 1988 

Superfamily Nephropoidea Dana, 1852 

Family Nephropidae Dana, 1852 

Thaumastochelidae Bate, 1888 

Superfamily Astacoidea Latreille, 1802 

Family Astacidae Latreille, 1802 

Cambaridae Hobbs, 1942 

Superfamily Parastacoidea Huxley, 1879 

Family Parastacidae Huxley, 1879 

Infraorder Thalassinidea Latreille, 1831 

Superfamily Thalassinoidea Latreille, 1831 

Family Thalassinidae Latreille, 1831 

Superfamily Callianassoidea Dana, 1852 

Family Callianassidae Dana, 1852 

Callianideidae Kossmann, 1880 

Ctenochelidae Manning & Felder, 1991 

Laomediidae Borradaile, 1903 

Thomassiniidae de Saint Laurent, 1979 

Upogebiidae Borradaile, 1903 

Superfamily Axioidea Huxley, 1879 

Family Axiidae Huxley, 1879 

Calocarididae Ortmann, 1891 

Micheleidae Sakai, 1992 

Strahlaxiidae Poore, 1994 

Infraorder Palinura Latreille, 1802 

Superfamily Eryonoidea de Haan, 1841 

Family Polychelidae Wood-Mason, 1874 

Superfamily Palinuroidea Latreille, 1802 

Family Palinuridae Latreille, 1802 

Scyllaridae Latreille, 1825 

Synaxidae Bate, 1881 

Infraorder Anomura MacLeay, 1838 

Superfamily Lomisoidea Bouvier, 1895 

Family Lomisidae Bouvier, 1895 

Superfamily Galatheoidea Samouelle, 1819 

Family Aeglidae Dana, 1852 

Chirostylidae Ortmann, 1892 

Galatheidae Samouelle, 1819 

Porcellanidae Haworth, 1825 

Superfamily Hippoidea Latreille, 1825 

Family Albuneidae Stimpson, 1858 

Hippidae Latreille, 1825 

Superfamily Paguroidea Latreille, 1802 

Family Coenobitidae Dana, 1851 


Diogenidae Ortmann, 1892 

Lithodidae Samouelle, 1819 

Paguridae Latreille, 1802 

Parapaguridae Smith, 1882 

Pylochelidae Bate, 1888 

Infraorder Brachyura Latreille, 1802 

Section Dromiacea de Haan, 1833 

Superfamily Homolodromioidea Alcock, 1900 

Family Homolodromiidae Alcock, 1900 

Superfamily Dromioidea de Haan, 1833 

Family Dromiidae de Haan, 1833 

Dynomenidae Ortmann, 1892 

Superfamily Homoloidea de Haan, 1839 

Family Homolidae de Haan, 1839 

Latreilliidae Stimpson, 1858 

Poupiniidae Guinot, 1991 

Section Eubrachyura de Saint Laurent, 1980 

Subsection Raninoida de Haan, 1839 

Superfamily Raninoidea de Haan, 1839 

Family Raninidae de Haan, 1839 

Symethidae Goeke, 1981 

Superfamily Cyclodorippoidea Ortmann, 1892 

Family Cyclodorippidae Ortmann, 1892 

Cymonomidae Bouvier, 1897 

Phyllotymolinidae Tavares, 1998 

Subsection Heterotremata Guinot, 1977 

Superfamily Dorippoidea MacLeay, 1838 

Family Dorippidae MacLeay, 1838 

Orithyiidae Dana, 1853 

Superfamily Calappoidea Milne Edwards, 1837 

Family Calappidae Milne Edwards, 1837 

Hepatidae Stimpson, 1871 

Superfamily Leucosioidea Samouelle, 1819 

Family Leucosiidae Samouelle, 1819 

Matutidae de Hann, 1841 

Superfamily Majoidea Samouelle, 1819 

Family Epialtidae MacLeay, 1838 

Inachidae MacLeay, 1838 

Inachoididae Dana, 1851 

Majidae Samouelle, 1819 

Mithracidae Balss, 1929 

Pisidae Dana, 1851 

Tychidae Dana, 1851 

Superfamily Hymenosomatoidea MacLeay, 1838 

Family Hymenosomatidae MacLeay, 1838 

Superfamily Parthenopoidea MacLeay, 1838 

Family Aethridae Dana, 1851 

Dairidae Ng & Rodriguez, 1986 

Daldorfiidae Ng & Rodriguez, 1986 

Parthenopidae MacLeay, 1838 

Superfamily Retroplumoidea Gill, 1894 

Family Retroplumidae Gill, 1894 

Superfamily Cancroidea Latreille, 1802 

Family Atelecyclidae Ortmann, 1893 

Cancridae Latreille, 1802 

Cheiragonidae Ortmann, 1893 

Corystidae Samouelle, 1819 

Pirimelidae Alcock, 1899 

Thiidae Dana, 1852 

Superfamily Portunoidea Rafinesque, 1815 

Family Geryonidae Colosi, 1923 

Portunidae Rafinesque, 1815 


Trichodactylidae Milne Edwards, 1853 

Superfamily Bythograeoidea Williams, 1980 

Family Bythograeidae Williams, 1980 

Superfamily Xanthoidea MacLeay, 1838 

Family Carpiliidae Ortmann, 1893 

Eumedonidae Dana, 1853 

Goneplacidae MacLeay, 1838 

Hexapodidae Miers, 1886 

Menippidae Ortmann, 1893 

Panopeidae Ortmann, 1893 

Pilumnidae Samouelle, 1819 

Platyxanthidae Guinot, 1977 

Pseudorhombilidae Alcock, 1900 

Trapeziidae Miers, 1886 

Xanthidae MacLeay, 1838 

Superfamily Bellioidea Dana, 1852 

Family Belliidae Dana, 1852 

Superfamily Potamoidea Ortmann, 1896 

Family Deckeniidae Ortmann, 1897 

Platythelphusidae Colosi, 1920 

Potamidae Ortmann, 1896 

Potamonautidae Bott, 1970 

Superfamily Pseudothelphusoidea Ortmann, 1893 

Family Pseudothelphusidae Ortmann, 1893 

Superfamily Gecarcinucoidea Rathbun, 1904 

Family Gecarcinucidae Rathbun, 1904 

Parathelphusidae Alcock, 1910 

Superfamily Cryptochiroidea Paulson, 1875 

Family Cryptochiridae Paulson, 1875 

Subsection Thoracotremata Guinot, 1977 

Superfamily Pinnotheroidea de Haan, 1833 

Family Pinnotheridae de Haan, 1833 

Superfamily Ocypodoidea Rafinesque, 1815 

Family Camptandriidae Stimpson, 1858 

Mictyridae Dana, 1851 

Ocypodidae Rafinesque, 1815 

Palicidae Bouvier, 1898 

Superfamily Grapsoidea MacLeay, 1838 

Family Gecarcinidae MacLeay, 1838 

Glyptograpsidae Schubart, Cuesta & Felder, 2001 

Grapsidae MacLeay, 1838 

Plagusiidae Dana, 1851 

Sesarmidae Dana, 1851 

Varunidae Milne Edwards, 1853 


Abele, L. G. 1987. Review of: Schram, F. R. 1986. Crustacea. 

Oxford University Press. Journal of Crustacean 

Biology 7:200–201. 

. 1991. Comparisons of morphological and molecular 

phylogeny of the Decapoda. In Proceedings of 

the 1990 International Crustacean Conference, ed. 

P. J. F. Davie and R. H. Quinn. Memoirs of the 

Queensland Museum 31:101–108. 

Abele, L. G., and B. E. Felgenhauer. 1986. Phylogenetic 

and phenetic relationships among the lower Decapoda. 

Journal of Crustacean Biology 6:385–400. 

Abele, L. G., W. Kim, and B. E. Felgenhauer. 1989. Molecular 

evidence for inclusion of the phylum Pentastomida 

in the Crustacea. Molecular Biology and 

Evolution 6:685–691. 

Abele, L. G., and T. Spears. 1997. Issues and answers in 

the molecular phylogeny of the Crustacea. Program 

and Abstracts, The Crustacean Society 1997 Summer 

Meeting, Mobile, Alabama: 13. 

Abele, L. G., T. Spears, and N. Cumberlidge. 1999. Biogeography 

and phylogeny of freshwater crabs based 

on molecular evidence. Program and Abstracts, The 

Crustacean Society 1999 Summer Meeting, Lafayette, 

Louisiana: 20. 

Abele, L. G., T. Spears, W. Kim, and M. Applegate. 1992. 

Phylogeny of selected maxillopodan and other crustacean 

taxa based on 18S ribosomal nucleotide sequences: 

a preliminary analysis. Acta Zoologica 73: 

271–392. 

Aguinaldo, A. M., J. M. Turbeville, L. S. Linford, M. C. 

Rivera, J. R. Garey, R. A. Raffe, and J. A. Lake. 

1997. Evidence for a clade of nematodes, arthropods, 

and other moulting animals. Nature 387:489– 

493. 

Ahyong, S. T. 1997. Phylogenetic analysis of the Stomatopoda 

(Malacostraca). Journal of Crustacean Biology 

17:695–715. 

. 2001. Revision of the Australian stomatopod 

Crustacea. Records of the Australian Museum, supplement 

26:1–326. 

Ahyong, S. T., and C. Harling. 2000. The phylogeny of 

the stomatopod Crustacea. Australian Journal of 

Zoology 48:607–642. 

Akam, M. 1998. Hox genes in arthropod development 

and evolution. Biological Bulletin 195:373–374 

Akam, M., M. Averof, J. Castelli-Gair, R. Dawes, F. Falciani, 

and D. Ferrier. 1994. The evolving role of Hox 

genes in arthropods. Development 1994(supplement):209–215. 

Alcock, A. 1898. Materials for a carcinological fauna of 

India. No. 3. The Brachyura Cyclometopa. Part I. 

The Family Xanthidae. Journal of the Asiatic Society 

of Bengal 68(part 2, no. 1):67–233. 

. 1900. Materials for a carcinological fauna of India. 

No. 5. The Brachyura Primigenia or Dromiacea. 

Journal of the Asiatic Society of Bengal 68(part 2, 

no. 3):123–169. 

Almeida, W. de O., and M. L. Christoffersen. 1999. A 

cladistic approach to relationships in Pentastomida. 

Journal of Parasitology 85:695–704. 

Alonso, M. 1996. Crustacea Branchiopoda. Fauna Iberica, 

vol. 7. Madrid: Museo Nacional de Ciencias Naturales, 

486 pp. 

Amado, M. A. P., J.-S. Ho, and C. E. F. da Rocha. 1995. 

Phylogeny and biogeography of the Ergasilidae (Copepoda, 

Poecilostomatoidea), with reconsideration 

LITERATURE CITED 

of the taxonomic status of the Vaigamidae. Contributions 

to Zoology 65:233–243. 

Amoros, C. 1996. Branchiopodes II. Ordre des Cténopodes, 

Anomopodes, Onychopodes et Haplopodes 

(Ctenopoda Sars, 1865—Anomopoda Sars, 1865— 

Onychopoda Sars, 1865—Haplopoda Sars, 1865). 

In Traité de Zoologie. Anatomie, Systématique, Biologie. 

Crustacés. Tome VII, Fascicule II. Généralités 

(suite) et Systématique, ed. J. Forest, 353–383. Paris: 

Masson, 1002 pp. 

Anderson, D. T. 1994. Barnacles. Structure, function, development 

and evolution. London: Chapman and 

Hall, 357 pp. 

Andronov, V. N. 1974. Phylogenetic relations of large taxa 

within the suborder Calanoida (Crustacea, Copepoda). 

Zoologischeskii Zhurnal 53:1002–1012. [In 

Russian, with English summary] 

Arbizu, P. M., and G. Moura. 1994. The phylogenetic position 

of the Cylindropsyllinae Sars (Copepoda, Harpacticoida) 

and the systematic status of the Leptopontiinae 

Lang. Zoologisch Beiträge 35:55–77. 

Arhat, A., and T. C. Kaufman. 1999. Novel regulation of 

the homeotic gene Scr associated with a crustacean 

leg-to-maxilliped appendage transformation. Development 

126:1121–1128. 

Atkinson, R. J. A., and A. C. Taylor. 1988. Physiological 

ecology of burrowing decapods. In Aspects of Decapod 

Crustacean Biology, ed. A. A. Fincham and P. 

Rainbow. Symposium of the Zoological Society of 

London 59:201–226. 

Avdeev, G. V., and B. I. Sirenko. 1991. Chitonophilidae 

fam. n., a new family of parasitic copepods from the 

chitons of the north-western Pacific. Parazitologiya 

25:370–374. 

Averof, M., and M. Akam. 1993. Hom/Hox genes of Artemia: 

implications for the origin of insect and crustacean 

body plans. Current Biology 3:73–78. 

. 1995a. Hox genes and the diversification of insect 

and crustacean body plans. Nature 376:420–423. 

. 1995b. Insect–crustacean relationships: insights 

from comparative developmental and molecular 

studies. Philosophical Transactions of the Royal Society 

of London 347B:293–303. 

Averof, M., and N. H. Patel. 1997. Crustacean appendage 

evolution associated with changes in Hox gene expression. 

Nature 388:682–686. 

Ax, P. 1999. Das system der Metazoa II. Ein Lehrbuch 

der phylogenetischen Systematik. Stuttgart: G. Fischer, 

383 pp. 

Baba, K. 1988. Chirostylid and galatheid crustaceans (Decapoda: 

Anomura) of the ‘‘Albatross’’ Philippine Expedition. 

Researches on Crustacea, special no. 2. Tokyo: 

The Carcinological Society of Japan, 203 pp. 

Băcescu, M. 1972. Archaeocuma and Schizocuma, new 

genera of Cumacea from the American tropical waters. 

Reue Roumaine de Biologie, serie Zoologie 17: 

241–245. 

. 1988. Cumacea I (Fam. Archaeocumatidae, Lampropidae, 

Bodotriidae, Leuconidae). In Crustaceourum 

catalogus pars 7, ed. H.-E. Gruner and L. B. 

Holthuis, 1–173. The Hague: SPB Academic Publishing. 

. 1992. Cumacea II (Fam. Nannastacidae, Diastylidae, 

Pseudocumatideae, Gynodiastylidae et Ceratocumatidae). 

In Crustaceorum catalogus pars 8, ed. 

76 � Contributions in Science, Number 39 Literature Cited

H.-E. Gruner and L. B. Holthuis, 1–468. The Hague: 

SPB Academic Publishing. 

Băcescu, M., and I. Petrescu. 1999. Ordre des Cumacés 

(Cumacea Kroyer, 1846). In Traité de Zoologie. Anatomie, 

Systématique, Biologie. Tome VII, Fascicule 

IIIA. Crustacés Péracarides, ed. J. Forest. Memoires 

de l’Institut Oceanographique Fondation Albert I er , 

Prince de Monaco, 19:391–428. 

Baird, W. 1850. The natural history of British Entomostraca. 

London: The Ray Society, 364 pp. 

Baker, A. de C., B. P. Boden, and E. Brinton. 1990. A 

practical guide to the euphausiids of the world. London: 

Natural History Museum Publications, The 

Natural History Museum, 96 pp. 

Barber, P. H., and M. V. Erdmann. 2000. Molecular systematics 

of the Gonodactylidae (Stomatopoda) using 

mitochondrial cytochrome oxidase C (subunit 1) 

DNA sequence data. Journal of Crustacean Biology, 

special no. 2, 20:20–36. 

Barnard, J. L., and J. Clark. 1984. Redescription of Phoxocephalopsis 

zimmeri with a new species, and establishment 

of the family Phoxocephalopsidae (Amphipoda) 

from Magellanic South America. Journal of 

Crustacean Biology 4:85–105. 

Barnard, J. L., and M. M. Drummond. 1982. Gammaridean 

amphipoda of Australia, part V: superfamily 

Haustorioidea. Smithsonian Contributions in Zoology 

360:1–148. 

Barnard, J. L., and G. S. Karaman. 1982. Classificatory 

revisions in gammaridean Amphipoda (Crustacea), 

part 2. Proceedings of the Biological Society of 

Washington 95:167–187. 

. 1987. Revisions in classification of gammaridean 

Amphipoda (Crustacea), part 3. Proceedings of the 

Biological Society of Washington 100:856–875. 

. 1991. The families and genera of marine gammaridean 

Amphipoda (except marine gammaroids). Records 

of the Australian Museum Supplement 

13(parts 1 and 2):1–866. 

Barnard, J. L., and J. D. Thomas. 1988. Ipanemidae, new 

family, Ipanema talpa, new genus and species, from 

the surf zone of Brazil (Crustacea: Amphipoda: 

Haustorioidea). Proceedings of the Biological Society 

of Washington 101:614–621. 

Barnes, R. D., and F. W. Harrison. 1992. Introduction. In 

Microscopic anatomy of invertebrates, vol. 9, Crustacea, 

ed. F. W. Harrison and A. G. Humes, 1–8. 

New York: Wiley-Liss, Inc. 

Bate, C. S. 1862. Catalogue of the specimens of amphipodous 

Crustacea in the collection of the British Museum. 

London: British Museum of Natural History, 

iv � 399 pages, plates. 1–58. 

Belk, D. 1996. Was sind ‘‘Urzeitkrebse’’? Stapfia 42, Zugleich 

Kataloge des O. Ö. Landesmuseums N. F. 

100:15–19. 

Bellan-Santini, D. 1999. Ordre des Amphipodes (Amphipoda 

Latreille, 1816). In Traité de Zoologie. Anatomie, 





Bellan-Santini, D., and M. Ledoyer. 1986. Gammariens 

(Crustacea—Amphipoda) des Iles Marion et Prince 

Edward. Boletino Museum Civico Storia Naturale 

Verona 13:349–435. 

Bellwood, O. 1996. A phylogenetic study of the Calappidae 

H. Milne Edwards 1837 (Crustacea: Brachyura) 

with a reappraisal of the status of the family. 

Zoological Journal of the Linnean Society 118:165– 

193. 

. 1998. Evolution and biogeography of the Calappidae 

(Decapoda: Brachyura). Proceedings and Abstracts 

of the 4th International Crustacean Congress, 

Amsterdam: 71 (abstract 101). 

Benton, M. J. (editor). 1993. The fossil record 2. London: 

Chapman and Hall, 845 pp. 

Berge, J., G. Boxshall, and W. Vader. 2000. Cladistic analysis 

of the Amphipoda. Abstracts of the 10th Colloqium 

on Amphipoda (http://www.odu.edu/ 

%7Ejrh100f/amphome): 1. 

Berge, J., W. Vader, and C. O. Coleman. 1998. Cladistic 

analysis of the amphipod families Odiidae and Ochlesidae. 

Proceedings and Abstracts of the 4th International 

Crustacean Congress, Amsterdam: 53 (abstract 

40). 

. 1999. A cladistic analysis of the amphipod families 

Ochlesidae and Odiidae, with description of a 

new species and genus. In Crustaceans and the biodiversity 

crisis. Proceedings of the 4th International 

Crustacean Congress, Amsterdam, vol. 1, ed. F. R. 

Schram and J. C. von Vaupel Klein, 239–265. Leiden: 

Brill. 

Bergström, J. 1992. The oldest arthropods and the origin 

of the Crustacea. Acta Zoologica 73:287–292. 

Boore, J. L., T. M. Collins, D. Stanton, L. L. Daehler, and 

W. M. Brown. 1995. Deducing arthropod phylogeny 

from mitochondrial DNA rearrangements. Nature 

376:163–165. 

Boore, J. L., D. Lavrov, and W. M. Brown. 1998. Gene 

translocation links insects and crustaceans. Nature 

392:667–668. 

Bott, R. 1970a. Die Süsswasserkrabben von Europa, 

Asien, Australien und ihre Stammesgeschichte. Eine 

Revision der Potamoidea und der Parathelphusoidea 

(Crustacea, Decapoda). Abhandlungen der Senckenbergischen 

Naturforschenden Gesellschaft 526:1– 

338. 

. 1970b. Betrachtungen uber die Entwicklungsgeschichte 

und Verbreitung der Süsswasser-krabben 

nach der Sammlung des Naturhistorischen Museums 

in Genf/Schweitz. Revue Suisse de Zoologie 77:327– 

344. 

Bourne, G. C. 1922. The Raninidae: a study in carcinology. 

Journal of the Linnean Society of London (Zoology) 

35:25–79. 

Bousfield, E. L. 1982a. Amphipoda. Gammaridea and Ingolfiellidea. 

In Synopsis and classification of living 

organisms, vol. 2, ed. S. B. Parker, 254–284, 293– 

294. New York: McGraw-Hill. 

. 1982b. Amphipoda (palaeohistory). In McGraw- 

Hill Yearbook of Science & Technology 1982–1983, 

96–100. New York: McGraw-Hill. 

. 1983. An updated phyletic classification and palaeohistory 

of the Amphipoda. In Crustacean Issues 

1. Crustacean Phylogeny, ed. F. R. Schram, 257–277. 

Rotterdam: A. A. Balkema Press, 372 pp. 

. 2000a. An updated commentary on the phyletic 

classification of amphipod crustaceans. Abstracts of 

the 10th Colloqium on Amphipoda (http://www. 

odu.edu/%7Ejrh100f/amphome): 1. 

. 2000b. The value of phyletic classification in biogeographical 

analyses. Abstracts of the 10th Colloqium 

on Amphipoda (http://www.odu.edu/%7Ejrh100f/ 

amphome): 1–2. 

. 2001. An updated commentary on phyletic classification 

of the amphipod Crustacea and its appli- 

Contributions in Science, Number 39 Literature Cited � 77

cability to the North American fauna. Amphipacifica 

3:49–120. 

Bousfield, E. L., and K. E. Conlan. 1990. Crustaceans. In 

The new Encyclopaedia Britannica, 15th edition, 

vol. 16, 840–859. Chicago: Encyclopaedia Brittanica, 

Inc. 

Bousfield, E. L., and E. A. Hendrycks. 1994. A revision 

of the family Pleustidae (Crustacea: Amphipoda: 

Leucothoidea). Systematics and biogeography of 

component subfamilies. Part I. Amphipacifica 1:17– 

58. 

. 1997. The amphipod superfamily Eusiroidea in 

the North American Pacific region. II. Family Calliopidae. 

Systematics and distributional ecology. Amphipacifica 

2:3–66. 

Bousfield, E. L., and P. M. Hoover. 1995. The amphipod 

superfamily Pontoporeioidea on the Pacific coast of 

North America. II. Family Haustoriidae. Genus Eohaustorius 

J. L. Barnard: systematics and distributional 

ecology. Amphipacifica 2:35–64. 

. 1997. The amphipod superfamily Corophioidea 

on the Pacific coast of North America. Part V. Family 

Corophiidae. Corophinae, new subfamily. Systematics 

and distributional ecology. Amphipacifica 2: 

67–139. 

Bousfield, E. L., and J. A. Kendall. 1994. The amphipod 

superfamily Dexaminoidea on the North American 

Pacific coast; families Atylidae and Dexaminidae: 

systematics and distributional ecology. Amphipacifica 

1:3–66. 

Bousfield, E. L., and C.-T. Shih. 1994. The phyletic classification 

of amphipod crustaceans: problems in resolution. 

Amphipacifica 1:76–134. 

Boutin, C., and M. Missouli. 1988. Metacrangonyx gineti, 

n. sp., d’une source du haut-Atlas Marocain det la 

famille des Metacrangonyctidae, n. fam. Vie et Milieu 

38:67–84. 

Bower, B. 1999. DNA’s evolutionary dilemma: genetic 

studies collide with the mystery of human evolution. 

Science News 155:88–90. 

Bowman, T. E., and L. G. Abele. 1982. Classification of 

the Recent Crustacea. In Systematics, the fossil record, 

and biogeography, ed. L. G. Abele, 1–27, vol. 

IofThe biology of Crustacea, ed. D. E. Bliss. New 

York: Academic Press. 

Bowman, T. E., S. P. Garner, R. R. Hessler, T. M. Iliffe, 

and H. L. Sanders. 1985. Mictacea, a new order of 

Crustacea Peracarida. Journal of Crustacean Biology 

5:74–78. 

Bowman, T. E., and H. E. Gruner. 1973. The families and 

genera of Hyperiidea. Smithsonian Contributions to 

Zoology 146:1–64. 

Bowman, T. E., and T. Iliffe. 1985. Mictocaris halope, a 

new unusual peracaridan crustacean from marine 

caves on Bermuda. Journal of Crustacean Biology 5: 

58–73. 

Boxshall, G. A. 1983. A comparative functional analysis 

of the major maxillopodan groups. In Crustacean Issues 

1. Crustacean Phylogeny, ed. F. R. Schram, 

121–143. Rotterdam: A. A. Balkema Press, 372 pp. 

. 1986. Panel discussion: copepod phylogeny. Syllogeus 

58:197–208. 

. 1988. A review of the copepod endoparasites of 

brittle stars (Ophiuroida). Bulletin of the British Museum 

of Natural History (Zoology) 54:261–270. 

. 1991. A review of the biology and phylogenetic 

relationships of the Tantulocarida, a subclass of 

Crustacea recognized in 1983. Verhandelungen der 

Deutschen Zoologischen Gesellschaft 84:271–279. 

. 1992. Synopsis of group discussion on the Maxillopoda. 

Acta Zoologica 73:335–337. 

. 1996. Classe de Tantulocarides (Tantulocarida 

Boxshall et Lincoln, 1983). In Traité de Zoologie. 

Anatomie, Systématique, Biologie. Crustacés. Tome 

VII, Fascicule II. Généralités (suite) et Systématique, 

ed. J. Forest, 399–408. Paris: Masson, 1002 pp. 

. 1999. Ordre des Spélaeogriphacés (Spelaeogriphacea 

Gordon, 1957). In Traité de Zoologie. Anatomie, 

Systématique, Biologie. Tome VII, Fascicule IIIA. 

Crustacés Péracarides, ed. J. Forest. Mémoires de 

l’Institut Océanographique Fondation Albert, I er , 


Boxshall, G. A., and D. Defaye. 1996. Classe des Mystacocarides 

(Mystacocarida Pennak et Zinn, 1943). In 

Traité de Zoologie. Anatomie, Systématique, Biologie. 




Boxshall, G. A., F. D. Ferrari, and H. Tiemann. 1984. The 

ancestral copepod: towards a consensus of opinion 

at the First International Conference on Copepoda. 

Crustaceana 7(supplement):68–84. 

Boxshall, G. A., and R. Huys. 1989a. New tantulocarid, 

Stygotantulus stocki, parasitic on harpacticoid copepods, 

with an analysis of the phylogenetic relationships 

within the Maxillopoda. Journal of Crustacean 

Biology 9:126–140. 

. 1989b. New family of deep-sea plankonic copepods, 

the Paralubbockiidae (Copepoda: Poecilostomatoida). 

Biological Oceanography 6:163–173. 

Boxshall, G. A., and D. Jaume. 1999. On the origin of 

misophrioid copepods from anchialine caves. Crustaceana 

72:957–963. 

. 2000. Discoveries of cave misophrioids (Crustacea: 

Copepoda) shed new light on the origin of anchialine 

faunas. Zoologische Anzeiger 239:1–19. 

Boxshall, G. A., and R. J. Lincoln. 1983. Tantulocarida, 

a new class of Crustacea ectoparasitic on other crustaceans. 


. 1987. The life cycle of the Tantulocarida (Crustacea). 

Philosophical Transactions of the Royal Society 


Boxshall, G. A., and S. Ohtsuka. 2001. Two new families 

of copepods (Copepoda: Siphonostomatoida) parasitic 

on echinoderms. Journal of Crustacean Biology 

21:96–105. 

Boxshall, G. A., J.-O. Strömberg, and E. Dahl (editors). 

1992. The Crustacea: origin and evolution. Acta 

Zoologica 73:271–392. 

Brady, G. S., and A. M. Norman, 1889. A monograph on 

the marine and freshwater Ostracoda of the North 

Atlantic and of northwestern Europe. Section I: Podocopa. 

Scientific Transactions of the Royal Dublin 

Society, series 2, 4:63–270. 

Braga, E., R. Zardoya, A. Meyer, and J. Yen. 1999. Mitochondrial 

and nuclear rRNA based copepod phylogeny 

with emphasis on the Euchaetidae (Calanoida). 

Marine Biology 133:79–90. 

Brandt, A., J. A. Crame, H. Polz, and M. R. A. Thomson. 

1999. Late Jurassic Tethyan ancestry of Recent 

southern high-latitude marine isopods (Crustacea, 

Malacostraca). Palaeontology 42:663–675. 

Brasil-Lima, I. M. 1998. Malacostraca—Peracarida. Isopoda. 

Epicaridea. In Catalogue of Crustacea of Bra- 


zil, ed. P. S. Young, 635–644. Rio de Janeiro: Museu 

Nacional. 

Briggs, D. E. G. 1983. Affinities and early evolution of the 

Crustacea: the evidence of the Cambrian fossils. In 

Crustacean issues 1. Crustacean phylogeny, ed. F. R. 

Schram, 1–22. Rotterdam: A. A. Balkema Press, 372 

pp. 

Briggs, D. E. G., D. H. Erwin, and F. J. Collier. 1994. The 

fossils of the Burgess Shale. Washington, D.C.: 

Smithsonian Institution Press, 238 pp. 

Briggs, D. E. G., and R. A. Fortey. 1989. The early radiation 

and relationships of the major arthropod 

groups. Science 246:241–243. 

. 1992. Chapter 10. The Early Cambrian radiation 

of arthropods. In Origin and early evolution of the 

Metazoa, ed. J. H. Lipps and P. W. Signor, 335–373. 

New York: Plenum Press. 

Briggs, D. E. G., R. A. Fortey, and M. A. Wills. 1992. 

Morphological disparity in the Cambrian. Science 

256:1670–1673. 

. 1993a. How big was the Cambrian explosion? A 

taxonomic and morphologic comparison of Cambrian 

and Recent arthropods. In Evolutionary patterns 

and processes, Linnean Society Symposium, ed. D. 

R. Lees and D. Edwards, 33–44. London: Linnean 

Society. 

Briggs, D. E. G., M. J. Weedon, and M. A. Whyte. 1993b. 

Arthropoda (Crustacea excluding Ostracoda). In 

The Fossil Record 2, ed. M. J. Benton, 321–342. 

London: Chapman and Hall. 

Briggs, D. E. G., and H. B. Whittington. 1981. Relationships 

of arthropods from the Burgess Shale and other 

Cambrian sequences. In Short Papers for the Second 

International Symposium on the Cambrian System, 

ed. M. E. Taylor, 38–41. U.S. Geological Survey 

Open-File Report 81-743:1–254. 

Brower, A. V. Z., R. DeSalle, and A. Vogler. 1996. Gene 

trees, species trees, and systematics: a cladistic perspective. 

Annual Review of Ecology and Systematics 

27:423–450. 

Brtek, J. 1964. Eine neue gattung und familie der ordnung 

Anostraca. Annotationes Zoologicae et Botanicae 

(Slovenské Národné Múzeum) 7:1–7. 

. 1995. Some notes on the taxonomy of the family 

Chirocephalidae (Crustacea, Branchiopoda, Anostraca). 

Zborník Slovenského Národného Mu´zea 

(Prirodné Vedy) 41:3–15. 

. 1997. Checklist of the valid and invalid names of 

the ‘‘large branchiopods’’ (Anostraca, Notostraca, 

Spinicaudata, and Laevicaudata), with a survey of 

the taxonomy of all Branchiopoda. Zborník Slovenského 

Národného Múzea (Prírondné Vedy) 43:1– 

66. 

Brtek, J., and G. Mura. 2000. Revised key to families and 

genera of the Anostraca with notes on their geographical 

distribution. Crustaceana 73:1037–1088. 

Brtek, J., and A. Thiéry. 1995. The geographic distribution 

of the European branchiopods (Anostraca, Notostraca, 

Spinicaudata, Laevicaudata). Hydrobiologia 298: 

263–280. 

Bruce, A. J. 1993. Kakadukaris glabra gen. nov., spec. 

nov., a new freshwater shrimp from the Kakadu National 

Park, Northern Territory, Australia (Crustacea: 

Decapoda: Palaemonidae) with designation of a 

new subfamily Kakaducaridinae. Hydrobiologia 

268:27–44. 

Bruce, N. L. 1984. A new family for the isopod crustacean 

genus Tridentella Richardson, 1905, with descrip- 

tion of a new species from Fiji. Zoological Journal 

of the Linnean Society 80:447–455. 

. 1988. Hadromastax merga, a new genus and species 

of marine isopod crustacean (Limnoriidae) from 

southeastern Australia, with discussion on the status 

of the families Keuphyliidae and Lynseiidae. Proceedings 

of the Biological Society of Washington 

101:346–353. 

. 1993. Two new genera of marine isopod crustaceans 

(Flabellifera: Sphaeromatidae) from southern 

Australia, with a reappraisal of the Sphaeromatidae. 

Invertebrate Taxonomy 7:151–171. 

Bruce, N. L., and H.-G. Müller. 1991. A new family for 

the isopod genus Hadromastax Bruce, 1988, with a 

description of a new species from the Society Islands. 


58. 

Brünnich, M. Th. 1772. Zoologiae fundamenta praelectionibus 

academicis accomodata. Grunde i Dyrelaeren. 

Hafniae et Lipsiae [� Copenhagen and Leipzig]: 

Apud Frider. Christ. Pelt., 254 pp. [not seen; as 

cited in Holthuis, 1991]. 

Brusca, R. C. 1990. Ferrara, F. (editor). Proceedings of the 

second symposium on the biology of terrrestrial isopods 

(Review). Journal of Crustacean Biology 10: 

568–570. 

. 2000. Unraveling the history of arthropod biodiversification. 

In Our unknown planet: recent discoveries 

and the future. Proceedings of the 45th Annual 

Systematics Symposium of the Missouri Botanical 

Garden. Annals of the Missouri Botanical Garden 

87:13–25. 

Brusca, R. C., and G. J. Brusca. 1990. Invertebrates. Sunderland, 

Massachusetts: Sinauer Associates, Inc., 

922 pp. 

Brusca, R. C., and G. D. F. Wilson. 1991. A phylogenetic 

analysis of the Isopoda with some classificatory recommendations. 

In Proceedings of the 1990 International 

Crustacean Conference, ed. P. J. F. Davie and 

R. H. Quinn. Memoirs of the Queensland Museum 

31:143–204. 

Buckeridge, J. S. 1983. Fossil barnacles (Cirripedia: Thoracica) 

of New Zealand and Australia. New Zealand 

Geological Survey Paleontological Bulletin 50:1– 

151. 

. 1995. Phylogeny and biogeography of the primitive 

Sessilia and a consideration of a Tethyan origin 

for the group. In Crustacean Issues 10. New Frontiers 

in Barnacle Evolution, ed. F. R. Schram and J. 

T. Høeg, 255–267. Rotterdam: A. A. Balkema Press, 

318 pp. 

Burkenroad, M. D. 1963. The evolution of the Eucarida 

(Crustacea, Eumalacostraca) in relation to the fossil 

record. Tulane Studies in Geology 2:3–16. 

. 1981. The higher taxonomy and evolution of Decapoda 

(Crustacea). Transactions of the San Diego 

Society of Natural History 19:251–268. 

Butterfield, N. J. 1994. Burgess Shale-type fossils from a 

Lower Cambrian shallow-shelf sequence in northwestern 

Canada. Nature 369:477–479. 

Cals, Ph. 1996. Classe des Rémipèdes (Remipedia Yager, 

1981). In Traité de Zoologie. Anatomie, Systématique, 

Biologie. Crustacés. Tome VII, Fascicule II. 

Généralités (suite) et Systématique, ed. J. Forest, 

385–397. Paris: Masson, 1002 pp. 

Cals, Ph., and Th. Monod. 1988. Évolution et biogéographie 

des Crustacés Thermosbénacés. Comptes Ren- 


dus Hebdomadaires des Seances de l’Academie des 

Sciences, series 3, 307:341–348. 

Camp, D. K., W. G. Lyons, and T. H. Perkins. 1998. 

Checklists of selected shallow-water marine invertebrates 

of Florida. Florida Marine Research Institute 

Technical Report TR-3, Florida Department of Environmental 

Protection, 1998:i–xv, 1–238. 

Cantino, P. D. 2000. Phylogenetic nomenclature: addressing 

some concerns. Taxon 49:85–93. 

Cantino, P. D., H. N. Bryant, K. de Queiroz, M. J. Donoghue, 

T. Eriksson, D. M. Hillis, and M. S. Y. Lee. 

1999. Species names in phylogenetic nomenclature. 

Systematic Biology 48:790–807. 

Cappola, V. A. 1999. Phylogeny of stomatopod crustaceans. 

Program and Abstracts, The Crustacean Society 

1999 Summer Meeting, Lafayette, Louisiana: 

25. 

Cappola, V. A., and R. B. Manning. 1998. Stomatopod 

superfamily Gonodactyloidea is not a natural group. 


Crustacean Congress, Amsterdam: 77 (abstract 121). 

Carpenter, J. H. 1999. Behavior and ecology of Speleonectes 

epilimnius (Remipedia, Speleonectidae) from 

surface water of an anchialine cave on San Salvador 

Island, Bahamas. Crustaceana 72:979–991. 

Carroll, S. B. 1995. Homeotic genes and the evolution of 

arthropods and chordates. Nature 376:479–485. 

Casanova, J.-P., L. De Jong, and E. Faure. 1998. Interrelationships 

of the two families constituting the Lophogastrida 

(Crustacea: Mysidacea) inferred from 

morphological and molecular data. Marine Biology 

132:59–65. 

Castro, P. 2000. Crustacea Decapoda: A revision of the 

Indo-West Pacific species of palicid crabs (Brachyura 

Palicidae). In Résultats des Campagnes Musorstom, 

vol. 21, ed. A. Crosnier. Mémoires du Museum National 

d’Histoire Naturelle, vol. 184, 437–610. 

Chace, F. A. 1992. On the classification of the Caridea 

(Decapoda). Crustaceana 63:70–80. 

. 1997. The caridean shrimps (Crustacea: Decapoda) 

of the Albatross Philippine Expedition, 1907– 

1910, Part 7: families Atyidae, Eugonatonotidae, 

Rhynchocinetidae, Bathypalaemonellidae, Processidae, 

and Hipploytidae. Smithsonian Contributions 

to Zoology 587:1–106. 

Chappuis, P. A. 1915. Bathynella natans und ihre Stellung 

im System. Zoologisches Jahrbüch der Systematik 

40:147–176. 

Chia, O. K. S., and P. K. L. Ng. 1998. Is the Sundathelphusidae 

Bott, 1969 a valid taxon? A cladistic appraisal. 


Crustacean Congress, Amsterdam: 72 (abstract 

103). 

Chia, D. G. B., and P. K. L. Ng. 2000. A revision of Eumedonus 

H. Milne Edwards, 1834 and Gonatonotus 

White, 1847 (Crustacea: Decapoda: Brachyura: Eumedonidae), 

two genera of crabs symbiotic with sea 

urchins. Journal of Natural History 34:15–56. 

Chow, S., M. Okazaki, M. Takeda, and T. Kubota. 2000. 

A rare abyssal shrimp, Galatheocaris abyssalis, 

found in the stomach of a lancetfish. Crustaceana 

73:243–246. 

Christoffersen, M. L. 1986. Phylogenetic relationships between 

Oplophoridae, Atyidae, Pasiphaeidae, Alvinocarididae 

fam. n., Bresiliidae, Psalidopodidae and 

Disciadidae (Crustacea Caridea Atyoidea). Bolletim 

de Zoologie (University Sao Paulo, Brazil) 10:273– 

281. 

. 1987. Phylogenetic relationships of hippolytid 

genera, with an assignment of new families for the 

Crangonoidea and Alpheoidea (Crustacea, Decapoda, 

Caridea). Cladistics 3:348–362. 

. 1988a. Phylogenetic systematics of the Eucarida 

(Crustacea Malacostraca). Revista Brasiliera de 

Zoologia 5:325–351. 

. 1988b. Genealogy and phylogenetic classification 

of the world Crangonidae (Crustacea, Caridea), with 

a new species and new records for the southwestern 

Atlantic. Revista Nordestina de Biologia 6:43–59. 

. 1989. Phylogeny and classification of the Pandaloidea 

(Crustacea, Caridea). Cladistics 5:259–274. 

. 1990. A new superfamily classification of the Caridea 

(Crustacea: Pleocyemata) based on phylogenetic 

pattern. Zeitschrift für Zoologische Systematik und 

Evolutionsforschung 28:94–106. 

. 1994. An overview of cladistic applications. Revista 

Nordestina de Biologia 9:133–141. 

. 1998. Malacostraca. Eucarida. Caridea. Crangonidea 

and Alpheoidea (except Glyphocrangoidae and 

Crangonidae). In Catalogue of Crustacea of Brazil, 

ed. P. S. Young, 351–372. Rio de Janeiro: Museu 

Nacional. 

Clark, P. F., and W. R. Webber. 1991. A redescription of 

Macrocheira kaempferi (Temminck, 1836) zoeas 

with a discussion of the classification of the Majoidea 

Samouelle, 1819 (Crustacea: Brachyura). Journal 

of Natural History 25:1259–1279. 

Coelho, P. A., and P. A. Coelho Filhol. 1993. Proposta de 

classificação da família Xanthidae (Crustacea, Decapoda, 

Brachyura) através da taxonomia numérica. 

Revista Brasileira de Zoologia 10:559–580. 

Cohen, A. C. 1982. Ostracoda. In Synopsis and classification 

of living organisms, ed. S. Parker, 181–202. 

New York: McGraw-Hill. 

Cohen, A. C., J. W. Martin, and L. S. Kornicker. 1998. 

Homology of Holocene ostracode biramous appendages 

with those of other crustaceans: the protopod, 

epipod, exopod, and endopod. Lethaia 31:251–265. 

Cohen, B. F. 1998. Dendrotiidae (Crustacea: Isopoda) of 

the southeastern Australian continental slope. Memoirs 

of the Museum of Victoria 57:1–38. 

Coineau, N. 1996. Sous-classe des Eumalacostracés (Eumalacostraca 

Grobben, 1892) super-ordre des Syncarides 

(Syncarida Packard, 1885). In Traité de 

Zoologie. Anatomie, Systématique, Biologie. Crustacés. 

Tome VII, Fascicule II. Généralités (suite) et 

Systématique, ed. J. Forest, 897–954. Paris: Masson, 

1002 pp. 

Coleman, C. O., and J. L. Barnard. 1991. Revision of 

Iphimediidae and similar families (Amphipoda: 

Gammaridea). Proceedings of the Biological Society 


Colgan, D. J., A. McLauchlan, G. D. F. Wilson, S. P. Livingston, 

G. D. Edgecomb, J. Macaranas, G. Cassis, 

and M. R. Gray. 1998. Histone H3 and U2 snRNA 

DNA sequences and arthropod molecular evolution. 

Australian Journal of Zoology 46:419–437. 

Cookson, L. J., and G. C. B. Poore. 1994. New species of 

Lynseia and transfer of the genus to Limnoriidae. 

Memoirs of the Museum of Victoria 54:179–189. 

Crandall, K. A. 1999. On the origin(s) of freshwater crayfishes. 


1999 Summer Meeting, Lafayette, Louisiana: 

27. 

Crandall, K. A., D. J. Harris, and J. W. Fetzner, Jr. 2000. 

The monophyletic origin of freshwater crayfish esti- 


mated from nuclear and mitochondrial DNA sequences. 

Proceedings of the Royal Society of London 

B 267:1679–1686. 

Crease, T. J., and D. J. Taylor. 1998. The origin and evolution 

of variable-region helices in V4 and V7 of the 

small-subunit ribosomal RNA of branchiopod crustaceans. 

Molecular Biology and Evolution 15:1430– 

1446. 

Cuesta, J. A., and C. D. Schubart. 1999. Proposed classification 

of the Grapsidae and Gecarcinidae (Decapoda, 

Brachyura) on the basis of larval morphology. 


1999 Summer Meeting, Lafayette, Louisiana: 52. 

Cuesta, J. A., C. D. Schubart, and A. Rodríguez. 2000. 

Larval development of Brachynotus sexdentatus 

(Risso, 1827) (Decapoda, Brachyura) reared under 

laboratory conditions, with notes on larval characters 

of the Varunidae. Invertebrate Reproduction 

and Development 38:207–223. 

Cumberlidge, N. 1987. Notes on the taxonomy of West 

African gecarcinucids of the genus Globonautes 

Bott, 1959 (Decapoda, Brachyura). Canadian Journal 

of Zoology 65:2210–2215. 

. 1991. The respiratory system of Globonautes macropus 

(Rathbun, 1898), a terrestrial freshwater crab 

from Liberia (Gecarcinucoidea, Gecarcinucidae). 

Crustaceana 61:69–80. 

. 1996a. A taxonomic revision of freshwater crabs 

(Brachyura, Potamoidea, Gecarcinucidae) from the 

Upper Guinea forest of West Africa. Crustaceana 69: 

681–695. 

. 1996b. On the Globonautinae Bott, 1969, freshwater 

crabs from West Africa (Brachyura: Potamoidea: 

Gecarcinucidae). Crustaceana 69:809–820. 

. 1999. The freshwater crabs of West Africa. Family 

Potamonautidae. Paris: IRD Éditions, Faune et 

Flore Tropicales 35, 1–382. 

Cumberlidge, N., and R. Sachs. 1991. Ecology, distribution, 

and growth in Globonautes macropus (Rathbun, 

1898), a tree-living freshwater crab from the 

rain forests of Liberia (Parathelphusoidea, Gecarcinucidae). 


Cumberlidge, N., and R. von Sternberg. 1999. Phylogenetic 

relationships of the freshwater crabs of Lake 

Tanganyika (Decapoda, Brachyura). In Crustaceans 

and the biodiversity crisis. Proceedings of the 4th 

International Crustacean Congress, Amsterdam, vol. 

1, ed. F. R. Schram and J. C. von Vaupel Klein, 405– 

422. Leiden: Brill. 

Cumberlidge, N., R. von Sternberg, I. R. Bills, and H. 

Martin. 1999. A revision of the genus Platythelphusa 

A. Milne-Edwards, 1887 from Lake Tanganyika, 

East Africa (Decapoda: Potamoidea: Platythelphusidae). 

Journal of Natural History 33:1487–1512. 

Cunningham, C. W., N. W. Blackstone, and L. W. Buss. 

1992. Evolution of king crabs from hermit crab ancestors. 

Nature 355:539–542. 

Dahl, E. 1983a. Alternatives in malacostracan evolution. 

Australian Museum Memoir 18:1–5. 

. 1983b. Malacostracan phylogeny and evolution. 

In Crustacean issues 1. Crustacean phylogeny, ed. F. 

R. Schram, 189–212. Rotterdam: A. A. Balkema 

Press, 372 pp. 

. 1987. Malacostraca maltreated—the case of the 

Phyllocarida. Journal of Crustacean Biology 7:721– 

726. 

. 1991. Crustacea Phyllopoda and Malacostraca: a 

reappraisal of cephalic dorsal shield and fold sys- 

tems. Philosophical Transactions of the Royal Society 


. 1992. Aspects of malacostracan evolution. Acta 

Zoologica 73:339–346. 

Dahl, E., and J.-O. Strömberg. 1992. Introduction: discovering 

crustacean diversity. Acta Zoologica 73: 

271–272. 

Dahl, E., and J.-W. Wägele. 1996. Sous-classe des Phyllocarides 

(Phyllocarida Packard, 1879). In Traité de 




1002 pp. 

Dahms, H.-U. 1990. Naupliar development of Harpacticoida 

(Crustacea, Copepoda) and its significance for 

phylogenetic systematics. Microfauna Marina 6: 

169–272. 

. 1993. Copepodid development in Harpacticoida 

(Crustacea, Copepoda). Microfauna Marina 8:195– 

245. 

Dai, A.-Y. 1997. A revision of freshwater crabs of the 

genus Nanhaipotamon Bott, 1968 from China 

(Crustacea: Decapoda: Brachyura: Potamidae). Raffles 

Bulletin of Zoology 45:209–235. 

Dai, A.-Y., and M. Türkay. 1997. Revision of the Chinese 

freshwater crabs previously placed in the genus Isolapotamon 

Bott, 1968 (Crustacea: Decapoda: Brachyura: 

Potamidae). Raffles Bulletin of Zoology 45: 

237–264. 

Dai, A.-Y., and S.-I. Yang. 1991. Crabs of the China Seas. 

Beijing: China Ocean Press, and Berlin: Springer Verlag, 

682 pp. [1991 English reprint of a 1984 Chinese 

volume] 

Dai, A.-Y., X. M. Zhou, and W. D. Peng. 1995. Eight new 

species of the genus Sinopotamon from Jiangxi Province, 

China (Crustacea, Decapoda, Brachyura, Potamidae). 

Beaufortia 45:61–76. 

Damkaer, D. M. 1996. Copepod taxonomy: discovery vs. 

recognition. Proceedings of the Biological Society of 


Dana, J. D. 1849. Conpsectus crustaceorum quae in orbis 

terrarum circumnavigatione, Carolo Wilkes e classe 

Reipublicae, Foederate Duce, lexit et descripsit (continued). 

American Journal of Sciences and Arts 8(2): 

424–428. 

. 1852. On the classification of the Crustacea Choristopoda 

or Tetradecapoda. American Journal of Sciences 

and Arts 14(2):297–316. 

Danielopol, D. L., and J. M. Betsch. 1980. Ostracodes 

terrestres de Madagascar: systematique, origine, adaptations. 

Revue d’Ecologie et de Biologie du Sol 17: 

87–123. 

da Rocha, C. E. F., and T. M. Iliffe. 1991. Speleoithonidae, 

a new family of Copepoda (Cyclopoida) from anchialine 

caves in the Bahama Islands. Sarsia 76:167– 

175. 

Davidson, E. H., K. J. Peterson, and R. A. Cameron. 

1995. Origin of bilaterian body plans: evolution of 

developmental regulatory mechanisms. Science 270: 

1319–1325. 

Davie, P. J. F. 1990. A new genus and species of marine 

crayfish, Palibythus magnificus, and new records of 

Palinurellus (Decapoda: Palinuridae) from the Pacific 

Ocean. Invertebrate Taxonomy 4:685–695. 

Day, J. 1980. Southern African Cumacea, part 4. Families 

Gynodiastylidae and Diastylidae. Annals of the 

South African Museum 82:187–292. 

De Broyer, C., and K. Jazdzewski. 1993. Contribution to 


the marine biodiversity inventory. A checklist of the 

Amphipoda (Crustacea) of the Southern Ocean. 

Documents de Travail de l’Institut Royal des Sciences 

Naturelles des Belgique 73:1–154. 

De Jong-Moreau, L., and J.-P. Casanova. 2001. The foreguts 

of the primitive families of the Mysida (Crustacea, 

Peracarida): a transitional link between those 

of the Lophogastrida (Crustacea, Mysidacea) and 

the most evolved Mysida. Acta Zoologica 82:137– 

147. 

Delle Cave, L., and A. M. Simonetta. 1991. Early Paleozoic 

arthropods and problems of arthropod phylogeny, 

with some notes on taxa of doubtful affinities. 

In The early evolution of Metazoa and the significance 

of problematic taxa, ed. A. M. Simonetta and 

S. Conway Morris, 189–244. London: Cambridge 

University Press. 

de Queiroz, K., and J. Gauthier. 1994. Toward a phylogenetic 

system of biological nomenclature. Trends in 

Ecology and Evolution 9:27–31. 

Diesing, K. M. 1835 (1836). Versuch einter Monographie 

der Gattung Pentastoma. Annalen des Wiener Museums 

der Naturgeschichte 1:1–32. 

Doyle, J. J. 1997. Trees within trees: genes and species, 

molecules and morphology. Systematic Biology 46: 

537–553. 

Drach, P., and D. Guinot. 1983. Les Inachoididae Dana, 

famille de Majoidea caractérisée par des connexions 

morphologiques d’un type nouveau entre carapace, 

pleurites, sternites, et pléon. Comptes Rendus Hebdomadaires 

des Seances de l’Academie des Sciences, 

series 3, 297:37–42. 

Dupuis, C. 1975. Objections aux propositions de Bousfield 

& Holthuis (1969) concernant une douzaine de 

genres d’Amphipodes. Z.N.(S.) 1879. Bulletin of 

Zoological Nomenclature 32:3–5. 

Edgecombe, G. D. (editor). 1998. Arthropod fossils and 

phylogeny. New York: Columbia University Press, 

347 pp. 

Edgecomb, G. D., G. D. F. Wilson, D. J. Colgan, M. R. 

Gray, and G. Cassis. 2000. Arthropod cladistics: 

combined analysis of histone H3 and U2 snRNA sequences 

and morphology. Cladistics 16:155–203. 

Eernisse, D. J. 1997. Arthropod and annelid relationships 

re-examined. In Arthropod relationships, ed. R. A. 

Fortey and R. H. Thomas, 43–56. Systematics Association 

Special Volume Series 55. London: Chapman 

and Hall. 

. 1998. A brief guide to phylogenetic software. 

Trends in Genetics 14:473–475. 

Elofsson, R. 1992. To the question of eyes in primitive 

crustaceans. Acta Zoologica 73:369–372. 

Emerson, M. J., and F. R. Schram. 1990. The origin of 

crustacean biramous appendages and the evolution 

of Arthropoda. Science 250:667–669. 

. 1991. Remipedia, part 2, paleontology. Proceedings 

of the San Diego Society of Natural History 7: 

1–52. 

. 1997. Theories, patterns, and reality: game plan 

for arthropod phylogeny. In Arthropod relationships, 

ed. R. A. Fortey and R. H. Thomas, 67–86. 

Systematics Association Special Volume series 55. 

London: Chapman and Hall. 

Erhard, F. 1995. Vergleichend- und funktionell-anatomische 

Untersuchungen am Pleon der Oniscidea (Crustacea, 

Isopoda) Zugleich ein Beitrag zu phylogenetischen 

Systematik der Landasseln. Zoologia 48:114 

pp. 

Fain, A. 1961. Les pentastomides de l’Afrique centrale. 

Annales de Musee Royale de l’Afrique Centrale, serie 

8, 92:1–115. 

Felder, D. L., J. W. Martin, and J. W. Goy. 1985. Patterns 

in early postlarval development of decapods. In 

Crustacean issues 2. Larval growth, ed. A. M. Wenner, 

163–225. Rotterdam: A. A. Balkema Press, 236 

pp. 

Feldmann, R. M., and C. S. Hopkins. 1999. Reconsideration 

of the family status of fossil and extant calappid 

crabs. Program and Abstracts, The Crustacean 

Society 1999 Summer Meeting, Lafayette, Louisiana: 

29. 

Felgenhauer, B. E., and L. G. Abele. 1983. Phylogenetic 

relationships among the shrimp-like decapods (Penaeoidea, 

Caridea, Stenopodidea). In Crustacean issues 

1. Crustacean phylogeny, ed. F. R. Schram, 291– 

311. Rotterdam: A. A. Balkema Press, 372 pp. 

Felgenhauer, B. E., L. G. Abele, and D. L. Felder. 1992. 

Remipedia. In Microscopic anatomy of invertebrates, 

vol. 9, Crustacea, ed. F. W. Harrison and A. 

G. Humes, 225–247. New York: Wiley-Liss. 

Ferrara, F. (editor). 1989. Proceedings of the second symposium 

on the biology of terrestrial isopods. Monitore 

Zoologico Italiano, n.s., Monografia 4:1–512. 

Ferrara, F., and S. Taiti. 1983. Isopodi terrestri. Contributions 

a l’etude de la faune granitiques de l’archipel 

des Seychelles. Annales du Musee Royale Afrique 

Centrale, Tervuren (serie in octavo), Sciences Zoologiques 

240:1–92. 

. 1989. A new genus and species of terrestrial isopod 

from Malaysia (Crustacea, Oniscidea, Platyarthridae). 


Ferrari, F. D. 1988. Evolutionary transformations and 

Dollo’s law. Journal of Crustacean Biology 8:618– 

619. 

Ferrari, F. D., and E. L. Markhaseva. 1996. Parkius karenwishnerae, 

a new genus and species of calanoid 

copepod (Parkiidae, new family) from benthopelagic 

waters of the eastern tropical Pacific Ocean. Proceedings 

of the Biological Society of Washington 

109:264–285. 

Field, K. G., G. J. Olsen, D. J. Lane, S. J. Giovanni, M. 

T. Ghiselin, E. C. Raff, N. R. Pace, and R. A. Raff. 

1988. Molecular analysis of the animal kingdom. 

Science 239:748–753. 

Fiers, F. 1990. Abscondicola humesi, n. gen., n. sp., from 

the gill chambers of land crabs and the definition of 

Cancrincolidae n. fam. (Copepoda, Harpacticoida). 

Bulletin d’Institute Royal de Sciences Naturelles Belgique 

(Biologie) 60:69–103. 

Forest, J. 1977. Un groupement injustifie: la superfamille 

des Bresilioidea. Remarques critiques sur le statut 

des familles sous ce nom (Crustacea Decapoda Caridea). 

Bulletin de Museum National Histoire Naturelle 

(Paris), series 3, no. 475, Zoologie 332:869– 

888. 

. 1987. Les Pylochelidae ou ‘‘Pagures symétriques’’ 

(Crustacea Coenobitoidea). In Résultats des Campagnes 

Musorstum, vol. 3, ed. A. Crosnier. Mémoires 

du Muséum National d’Histoire Naturelle, sére 

A, Zoologie, vol. 137, 1–254. 

. (editor). 1996. Traité de Zoologie. Anatomie, Systématique, 

Biologie. Crustacés. Tome VII. Fascicule 

II. Généralities (suite) et systématique. Paris: Masson, 

1002 pp, 426 plates and figures, 22 tables. 

. (editor). 1999. Traité de Zoologie. Anatomie, Systématique, 

Biologie. Tome VII, Fascicule IIIA. Crus- 


tacés Péracarides. Memoires de l’Institut Oceanographique 

Fondation Albert I er , Prince de Monaco, 

19:450 pp. 

Forest, J., and M. de St. Laurent. 1989. Nouvelle contribution 

à la connaissance de Neoglyphea inopinata 

Forest & de Saint Laurent, à propos de la description 

de la femelle adulte. In Résultats des Campagnes 

Musorstom, vol. 5, ed. J. Forest. Mémoires de Museum 

National d’Histoire Naturelle (Paris), sére A, 

vol. 144, 75–92. 

Fortey, R. A., D. E. G. Briggs, and M. A. Wills. 1997. The 

Cambrian evolutionary ‘explosion’ recalibrated. 

BioEssays 19:429–434. 

Fortey, R. A., and R. H. Thomas (editors). 1997. Arthropod 

relationships. Systematics Association Special 

Volume series 55. London: Chapman and Hall, 367 

pp. 

Fosshagen, A., and T. M. Iliffe. 1985. Two new genera of 

Calanoida and a new order of Copepoda, Platycopioida, 

from marine caves on Bermuda. Sarsia 70: 

345–358. 

France, S. C., and T. D. Kocher. 1996. DNA sequencing 

of formalin-fixed crustaceans from archival research 

collections. Molecular Marine Biology and Technology 

5:304–313. 

Fresi, E., E. Idato, and M. B. Scipione. 1980. The Gnathostenetroidea 

and the evolution of primitive asellote 

isopods. Monitore Zoologico Italiano (Nuovo 

Serie) 14:119–136. 

Frey, D. 1995. Changing attitudes toward chydorid anomopods 

since 1769. Hydrobiologia 307:43–45. 

Friedrich, M., and D. Tautz. 1995. Ribosomal DNA phylogeny 

of the major extant arthropod classes and the 

evolution of myriapods. Nature 376:165–167. 

Fryer, G. 1983. Functional ontogenetic changes in Branchinecta 

ferox (Milne-Edwards) (Crustacea: Anostraca). 



. 1985. Structure and habits of living branchiopods 

and their bearing on the interpretation of fossil 

forms. Transactions of the Royal Society of Edinburgh 

76:103–113. 

. 1987a. Morphology and classification of the socalled 

Cladocera. Hydrobiologia 145:19–28. 

. 1987b. The feeding mechanisms of the Daphniidae 

(Crustacea: Cladocera): recent suggestions and 

neglected considerations. Journal of Plankton Research 

9:419–432. 

. 1987c. A new classification of the branchiopod 

Crustacea. Zoological Journal of the Linnean Society 

91:357–383. 

. 1987d. Quantitative and qualitative: numbers and 

reality in the study of living organisms. Freshwater 

Biology 17:177–189. 

. 1992. The origin of the Crustacea. Acta Zoologica 

73:273–286. 

. 1995. Phylogeny and adaptive radiation within 

the Anomopoda: a preliminary exploration. Hydrobiologia 

307:57–68. 

. 1996a. Reflections on arthropod evolution. Biological 

Journal of the Linnean Society 58:1–55. 

. 1996b. The carapace of the branchiopod Crustacea. 



. 1997. A defence of arthropod polyphyly. In Arthropod 

relationships, ed. R. A. Fortey and R. H. 

Thomas, 23–33. Systematics Association Special 

Volume Series 55. London: Chapman and Hall. 

. 1999a. The case of the one-eyed brine shrimp: are 

ancient atavisms possible? Journal of Natural History 

33:791–798. 

. 1999b. A comment on a recent phylogenetic analysis 

of certain orders of the branchiopod Crustacea. 


. 1999c. Cambrian animals: evolutionary curiosities 

or the crucible of creation? Hydrobiologia 403: 

1–11. 

. 2001. The elucidation of branchiopod phylogeny. 


Funch, P., and R. M. Kristensen. 1995. Cycliophora is a 

new phylum with affinities to Entoprocta and Ectoprocta. 

Nature 378:711–714. 

Galil, B. 2000. Crustacea Decapoda: review of the genera 

and species of the family Polychelidae Wood-Mason, 

1874. In Résultats des Campagnes Musorstom, vol. 

21, ed. A. Crosnier. Mémoires du Muséum National 


García-Machado, E., M. Pempera, N. Dennebouy, M. Oliver-Suarez, 

J. C. Mounolou, and M. Monnerot. 

1999. Mitochondrial genes collectively suggest the 

paraphyly of Crustacea with respect to Insecta. Journal 

of Molecular Evolution 49:142–149. 

Garey, J. R. 2000. Invertebrate evolution. In McGraw-Hill 

Yearbook of Science and Technology 2001, 216– 

218. New York: McGraw-Hill. 

Garey, J. R., M. Krotec, D. R. Nelson, and J. Brooks. 

1996. Molecular analysis supports a tardigrade-arthropod 

association. Invertebrate Biology 115:79– 

88. 

Giribet, G., and C. Ribera. 2000. A review of arthropod 

phylogeny: new data based on ribosomal DNA sequences 

and direct character optimization. Cladistics 

16:204–231. 

Glaessner, M. F. 1969. Decapoda. In Treatise on invertebrate 

paleontology, part R, Arthropoda 4, ed. R. C. 

Moore, R399–R651. Lawrence, Kansas: The Geological 

Society of America and the University of Kansas 

Press. 

Glenner, H., M. J. Grygier, J. T. Høeg, P. G. Jensen, and 

F. R. Schram. 1995. Cladistic analysis of the Cirripedia 

Thoracica. Zoological Journal of the Linnean 

Society 114:365–404. 

Grenier, J. K., T. L. Garber, R. Warren, P. M. Whittington, 

and S. Carroll. 1997. Evolution of the entire arthropod 

Hox gene set predated the origin and radiation 

of the onychophoran/arthropod clade. Current Biology 

7:547–553. 

Griffin, D. J. G., and H. A. Tranter. 1986. The Decapoda 

Brachyura of the Siboga Expedition. Part VIII. Majidae. 

Siboga-Expeditie Monograph 39, C4, Livraison 

148:1–335. 

Griffis, R. B., and T. H. Suchanek. 1991. A model of burrow 

architecture and trophic models in thalassinidean 

shrimp (Decapoda: Thalassinidea). Marine 

Ecology Progress Series 79:171–183. 

Gruner, H. E. 1993. Lehrbuch der Speziellen Zoologie begründet 

von Alfred Kaestner. Band. 1: Wirbellose Tiere. 

Teil 4: Arthropoda (ohne Insecta). Jena: Gustav 

Fischer Verlag, 1209 pp. 

. 1996. Classe de Branchioures (Branchiura Thorell, 

1864). In Traité de Zoologie. Anatomie, Systématique, 


Généralités (suite) et Systématique, ed. J. Forest, 

739–754. Paris: Masson, 1002 pp. 

Grygier, M. J. 1981. Sperm of the ascothoracican parasite 

Dendrogaster, the most primitive found in Crusta- 


cea. International Journal of Invertebrate Reproduction 

3:65–73. 

. 1982. Sperm morphology in Ascothoracida (Crustacea: 

Maxillopoda): confirmation of generalized nature 

and phylogenetic importance. International 

Journal of Invertebrate Reproduction 4:323–332. 

. 1983a. Ascothoracida and the unity of Maxillopoda. 

In Crustacean issues 1. Crustacean phylogeny, 

ed. F. R. Schram, 73–104. Rotterdam: A. A. Balkema 


. 1983b. Revision of Synagoga (Crustacea: Maxillopoda: 

Ascothoracida). Journal of Natural History 

17:213–239. 

. 1985. Comparative morphology and ontogeny of 

the Ascothoracida, a step toward a phylogeny of the 

Maxillopoda. Dissertation Abstracts International 

45:2466B–2467B. 

. 1987a. New records, external and internal anatomy, 

and systematic position of Hansen’s y-larvae 

(Crustacea: Maxillopoda: Facetotecta). Sarsia 72: 

261–278. 

. 1987b. Classification of the Ascothoracida (Crustacea). 

Proceedings of the Biological Society of 


. 1987c. Nauplii, antennular ontogeny, and the position 

of the Ascothoracida within the Maxillopoda. 


. 1996a. Sous-classe des Facetotecta (Facetotecta 

Grygier, 1985). In Traité de Zoologie. Anatomie, 

Systématique, Biologie. Crustacés. Tome VII, Fascicule 

II. Généralités (suite) et Systématique, ed. J. 

Forest, 425–432. Paris: Masson, 1002 pp. 

. 1996b. Sous-classe de Ascothoracides (Ascothoracida 

Lacaze-Duthiers, 1880). In Traité de Zoologie. 

Anatomie, Systématique, Biologie. Crustacés. Tome 



Grygier, M. J., and T. E. Bowman. 1990. The correct family-level 

name for the ‘‘cryptoniscid’’ isopods (Epicaridea). 


. 1991. The authorship of Cryptoniscidae (Isopoda, 

Epicaridea); a correction. Crustaceana 61:106–107. 

Guinot, D. 1977. Propositions pour une nouvelle classification 

des Crustacés Décapodes Brachyoures. Comptes 

Rendus Hebdomadaires des Seances de 

l’Academie des Sciences, series 3, 285:1049–1052. 

. 1978. Principes d’une classification évolutive des 

Crustacés Décapodes Brachyoures. Bulletin Biologique 

de le France et de le Belgique (n. s.) 112:211– 

292. 

. 1979. Données nouvelles sur la morphologie, la 

phylogenèse et la taxonomie des Crustacés Décapodes 

Brachyoures. Mémoires du Muséum National 

d’Histoire Naturelle (Paris) sére A, Zoologie 112:1– 

354. 

. 1985. Crustacea. In French Polynesian coral reefs, 

reef knowledge and field guides, 5th International 

Coral Reef Congress, Tahiti, ed. G. Richard, 446– 

455. Paris: Muséum National d’Histoire Naturelle 

and École Pratique des Hautes Études Antenne de 

Tahiti. 

. 1988. Les crabes des sources hydrothermales de 

la dorsale du Pacifique oriental (Campagne BIO- 

CYARISE), 1984. Oceanologica Acta (special volume) 

8:109–118. 

. 1990. Austinograea alayseae sp. nov., crabe hydrothermal 

découvert dans le bassin de Lau, Pacifique 

sud-occidental (Crustacea Decapoda Brachy- 

ura). Bulletin du Museum National d’Histoire (Paris) 

11:879–903. 

. 1991. Établissement de la famille des Poupiniidae 

pour Poupinia hirsuta gen. nov., sp. nov. de Polynésie 

(Crustacea Decapoda Brachyura Homoloidea). 

Bulletin du Museum National d’Historie Naturelle 

(Paris) 12:577–605. 

. 1993. Donnees nouvelles sur les Raninoidea de 

Haan, 1841 (Crustacea Decapoda Brachyura Podotremata). 

Comptes Rendus Hebdomadaires des Seances 

de l’Academie des Sciences, series 3, 316: 

1324–1331. 

. 1995. Crustacea Decapoda Brachyura: révision 

des Homolodromiidae Alcock, 1900. In Résultats 

des Campagnes Musorstom, vol. 13, ed. A. Crosnier. 

Mémoires du Muséum National d’Histoire Naturelle, 

vol. 163, 155–282. 

Guinot, D., and J.-M. Bouchard. 1998. Evolution of the 

abdominal holding systems of brachyuran crabs 

(Crustacea, Decapoda, Brachyura). Zoosystema 20: 

613–694. 

Guinot, D., B. G. M. Jamieson, and B. Richer de Forges. 

1994. Relationship of Homolidae and Dromiidae— 

evidence from spermatozoal ultrastructure (Crustacea, 

Decapoda). Acta Zoologica 75:255–267. 

Guinot, D., B. G. M. Jamieson, B. Richer de Forges, and 

C. C. Tudge. 1998. Comparative spermatozoal ultrastructure 

of the three dromiacean families exemplified 

by Homolodromia kai (Homolodromiiidae), 

Sphaerodromia lamellata (Dromiidae), and Dynomene 

tanensis (Dynomenidae) (Podotremata: Brachyura). 


Guinot, D., B. G. M. Jamieson, and C. C. Tudge. 1997. 

Ultrastructure and relationships of spermatozoa of 

the freshwater crabs Potamon fluviatile and Potamon 

ibericum (Crustacea, Decapoda, Potamidae). 

Journal of Zoology 241:229–244. 

Guinot, D., and B. Richer de Forges. 1995. Crustacea Decapoda 

Brachyura: révision des Homolidae de Haan, 

1839. In Résultats des Campagnes Musorstom, vol. 

13, ed. A. Cronsnier. Mémoires de Muséum National 


. 1997. Affinités entre les Hymenosomatidae Mac- 

Leay, 1838 et les Inachoididae Dana, 1851 (Crustacea, 

Decapoda, Brachyura). Zoosystema 19:453– 

502. 

Gutu, M. 1996. Tanaidacea (Crustacea, Peracarida) from 

Brazil, with description of new taxa and systematical 

remarks on some families. Travaux du Muséum National 

d’Histoire Naturelle ‘‘Grigore Antipa’’ 36:23– 

133. 

. 1998. Spelaeogriphacea and Mictacea (partim) 

suborders of a new order, Cosinzeneacea (Crustacea, 

Peracarida). Travaux du Muséum National 

d’Histoire Naturelle ‘‘Grigore Antipa’’ 40:121–129. 

Gutu, M., and T. M. Iliffe. 1998. Description of a new 

hirsutiid (n. g., n. sp.) and reassignment of this family 

from Order Mictacea to the new order, Bochusacea 

(Crustacea, Peracarida). Travaux du Muséum 

d’Histoire Naturelle ‘‘Grigore Antipa’’ 40:93–120. 

Gutu, M., and J. Sieg. 1999. Ordre des Tanaïdacés (Tanaidacea 

Hansen, 1895). In Traité de Zoologie. Anatomie, 





Hanner, R., and M. Fugate. 1997. Branchiopod phyloge- 


netic reconstruction from 12S rDNA sequence data. 


Hansen, H. J. 1895. Isopoden, Cumaceen und Stomatopoden 

der Plankton Expedition. Ergebnisse der im 

Atlanishcen Ozean Plankton-Expedition der Humbolt-Stiftung, 

1889 2:1–105. 

Harling, C. 2000. Reexamination of eye design in the classification 

of stomatopod crustaceans. Journal of 


Harris, D. J., L. S. Maxson, L. F. Braithwaite, and K. A. 

Crandall. 2000. Phylogeny of the thoracican barnacles 

based on 18S rDNA sequences. Journal of Crustacean 

Biology 20:393–398. Harrison, K., and J. P. 

Ellis. 1991. The genera of the Sphaeromatidae (Crustacea: 

Isopoda): a key and distribution list. Invertebrate 

Taxonomy 5:915–952. 

Hart, C. W. J., and R. B. Manning. 1986. Two new 

shrimps (Procarididae and Agostocarididae, new 

family) from marine caves of the western North Atlantic. 


Hartmann, G., and M.-C. Guillaume. 1996. Classe des 

Ostracodes (Ostracoda Latreille, 1802). In Traité de 




1002 pp. 

Harvey, A. H., J. W. Martin, and R. Wetzer. In press. 

Crustacea. In Atlas of marine invertebrate larvae, ed. 

C. Young, M. Sewell, and M. Rice. London: Academic 

Press. 

Harzsch, S., and D. Walossek. 2001. Neurogenesis in the 

developing visual system of the branchiopod crustacean 

Triops longicaudatus (LeConte, 1846): corresponding 

patterns of compound-eye formation in 

Crustacea and Insecta? Developmental Genes and 


Hay, W. P., and C. A. Shore. 1918. The decapod crustaceans 

of Beaufort, N. C., and the surrounding regions. 

Bulletin of the U.S. Fisheries Bureau 35(for 

1915 and 1916):369–475. 

Haye, P., and I. Kornfield. 1999. Familial relationships 

within the Cumacea: preliminary molecular findings. 


1999 Summer Meeting, Lafayette, Louisiana: 33. 

Hendrickx, M. E. 1995. Checklist of brachyuran crabs 

(Crustacea: Decapoda) from the eastern tropical Pacific. 

Bulletin de l’Institut Royal des Sciences Naturelles 

de Belgique, Biologie 65:125–150. 

. 1998. A new genus and species of ‘‘goneplacidlike’’ 

brachyuran crab (Crustacea: Decapoda) from 

the Gulf of California, Mexico, and a proposal for 

the use of the family Pseudorhombilidae Alcock, 

1900. Proceedings of the Biological Society of Washington 

111:634–644. 

. 1999. Los Cangrejos Braquiros (Crustacea: 

Brachyura: Majoidea y Parthenopoidea) del Pacifico 

Mexicano. Instituto de Ciencias del Mar y Limnología, 

Universidad Nacional Autónoma de México: 

CONABIO, 274 pp., 13 plates. 

Hessler, R. R. 1982. The structural morphology of walking 

mechanisms in eumalacostracan crustaceans. 

Philosophical Transactions of the Royal Society of 

London 296B:245–298. 

. 1983. A defense of the caridoid facies; wherein 

the early evolution of the Eumalacostraca is discussed. 

In Crustacean issues 1. Crustacean phylogeny, 

ed. F. R. Schram, 145–164. Rotterdam: A. A. 

Balkema Press, 372 pp. 

. 1984. Dahlella caldariensis, new genus, new species: 

a leptostracan (Crustacea, Malacostraca) from 

deep-sea hydrothermal vents. Journal of Crustacean 

Biology 4:655–664. 

. 1985. Swimming in Crustacea. Transactions of 

the Royal Society of Edinburgh 76:115–122. 

. 1992. Reflections on the phylogenetic position of 

the Cephalocarida. Acta Zoologica 73:315–316. 

. 1999. Ordre des Mictacés (Mictacea Bowman, 

Garner, Hessler, Iliffe, Sanders, 1985). In Traité de 

Zoologie. Anatomie, Systématique, Biologie. Tome 

VII, Fascicule IIIA. Crustacés Péracarides, ed. J. Forest. 

Memoires de l’Institut Oceanographique Fondation 

Albert I er , Prince de Monaco, 19:87–91. 

Hessler, R. R., and R. Elofsson. 1996. Classe de Céphalocarides 

(Cephalocarida Sanders, 1955). In Traité de 

Zoologie. Anatomie, Systematique, Biologie. Tome 



Hessler, R. R., and J. W. Martin. 1989. Austinograea williamsi, 

new genus, new species, a hydrothermal vent 

crab (Decapoda: Bythograeidae) from the Mariana 

Back-Arc Basin, western Pacific. Journal of Crustacean 

Biology 9:645–661. 

Hessler, R. R., and L. Watling. 1999. Les Péracarides: un 

groupe controversé. In Traité de Zoologie. Anatomie, 





Heymons, R. 1935. Pentastomida. In Bronns Klassen und 

Ordnungen das Tierreichs, vol. 5, 1–268. Leipzig: 

Akademische Verlagsgesellschaft. 

Hibbett, D. S., and M. J. Donoghue. 1998. Integrating 

phylogenetic analysis and classification in fungi. Mycologia 

90:347–356. 

Hickman, C. P., Jr., L. S. Roberts, and A. Larson. 1996. 

Integrated principles of zoology. Dubuque, Iowa: 

Wm. C. Brown Publishers, Times Mirror Higher Education 

Group, 901 pp. 

Ho, J.-S. 1984. New family of poecilostomatoid copepods 

(Spiophanicolidae) parasitic on polychaetes from 

southern California, with a phylogenetic analysis of 

nereicoliform families. Journal of Crustacean Biology 

4:134–146. 

. 1990. Phylogenetic analysis of copepod orders. 


. 1991. Phylogeny of Poecilostomatoida; a major 

order of symbiotic copepods. Proceedings of the 4th 

International Conference on Copepoda. Bulletin of 

the Plankton Society of Japan, Special Volume, 25– 

48. 

. 1994a. Copepod phylogeny: a reconsideration of 

Huys & Boxshall’s ‘‘parsimony versus homology.’’ 

Hydrobiologia 292/293:31–39. 

. 1994b. Origin and evolution of the parasitic cyclopoid 

copepods. International Journal for Parasitology 

24:1293–1300. 

Ho, J.-S., M. Conradi, and P. J. López-González. 1998. A 

new family of cyclopoid copepods (Fratiidae) symbiotic 

in the ascidian (Clavellina dellavallei) from 

Cádiz, Spain. Journal of Zoology 246:39–48. 

Ho, J.-S., and I. H. Kim. 1997. A new family of poecilostomatoid 

copepods (Polyankyliidae) from a tide pool 

on a mud flat in Korea. Korean Journal of Biological 

Sciences 1:429–434. 

Ho, J.-S., and V. E. Thatcher. 1989. A new family of cyclopoid 

copepods (Ozmanidae) parasitic in the he- 


mocoel of a snail from the Brazilian Amazon. Journal 

of Natural History 23:903–911. 

Høeg, J. T. 1992a. The phylogenetic position of the Rhizocephala: 

are they truly barnacles? Acta Zoologica 

73:271–392. 

. 1992b. Rhizocephala. In Microscopic anatomy of 

invertebrates. vol. 9, Crustacea, ed F. W. Harrison 

and A. G. Humes, 313–345. New York; Wiley-Liss. 

. 1995. Sex and the single cirriped: a phylogenetic 

perspective. In Crustacean issues 10. New frontiers 

in barnacle evolution, ed. F. R. Schram and J. T. 

Høeg, 195–207. Rotterdam: A. A. Balkema Press, 

318 pp. 

Høeg, J. T., B. Hosfeld, and P. G. Jensen. 1998. TEM 

studies on the lattice organs of cirripede cypris larvae 

(Crustacea, Thecostraca, Cirripedia). Zoomorphology 

118:195–205. 

Høeg, J. T., and J. Lützen. 1993. Comparative morphology 

and phylogeny of the family Thompsoniidae 

(Cirripedia, Rhizocephala, Akentrogonida), with descriptions 

of three new genera and seven new species. 

Zoologica Scripta 22:363–386. 

. 1996. Super-ordre des Rhizocephales (Rhizocephala 

F. Müller, 1862). In Traité de Zoologie. Anatomie, 

Systématique, Biologie. Crustacés. Tome VII, 

Fascicule II. Généralités (suite) et Systématique, ed. 

J. Forest, 541–570. Paris: Masson, 1002 pp. 

Høeg, J. T., and A. V. Rybakov. 1992. Revision of the 

Rhizocephala Akentrogonida (Cirripedia), with a list 

of all the species and a key to the identification of 

families. Journal of Crustacean Biology 12:600–609. 

Høeg, J. T., M. A. Whyte, H. Glenner, and F. R. Schram. 

1999. New evidence on the basic phylogeny of the 

Cirripedia Thoracia. In Crustaceans and the biodiversity 

crisis. Proceedings of the 4th International 

Crustacean Congress, Amsterdam, ed. F. R. Schram 

and J. C. von Vaupel Klein, 101–149. Leiden: Brill. 

Hof, C. H. J. 1998a. Mesozoic stomatopods. Proceedings 

and Abstracts of the 4th International Crustacean 

Congress, Amsterdam: 78 (abstract 123). 

. 1998b. Fossil stomatopods (Crustacea: Malacostraca) 

and their phylogenetic impact. Journal of Natural 

History 32:1567–1576. 

Hof, C. H. J., and F. R. Schram. 1999. The phylogenetic 

position and evolution of Hoplocarida. Program and 

Abstracts, The Crustacean Society 1999 Summer 

Meeting, Lafayette, Louisiana: 35. 

Hogans, W. E., and G. W. Benz. 1990. A new family of 

parasitic copepods, the Lernaeosoleidae (Poecilostomatoida), 

from demersal fishes in the northwest Atlantic, 

with a description of Bobkabata kabatabobbus 

n. gen., n. sp. and a redescription of Lernaeosolea 

lycodis Wilson, 1944. Canadian Journal of Zoology 

68:2483–2488. 

Holdich, D., R. Lincoln, and J. Ellis. 1984. The biology 

of terrestrial isopods: terminology and classification. 

Symposium of the Zoological Society of London 53: 

1–6. 

Holsinger, J. R. 1989. Allocrangonyctidae and Pseudocrangonyctidae, 

two new families of Holarctic subterranean 

amphipod crustaceans (Gammaridea), 

with comments on their phylogenetic and zoogeographic 

relationships. Proceedings of the Biological 

Society of Washington 102:947–959. 

. 1992. Sternophysingidae, a new family of subterranean 

amphipods (Gammaridea: Crangonyctoidea) 

from South Africa, with descriptions of Sternophysinx 

calceola, new species, and comments on phy- 

logenetic and biogeographic relationships. Journal of 


Holthuis, L. B. 1955. The Recent genera of the caridean 

and stenopodidean shrimps (Class Crustacea, Order 

Decapoda, supersection Natantia) with keys for their 

determination. Zoologische Verhandelingen 26:1– 

157. 

. 1991. Marine lobsters of the world. An annotated 

and illustrated catalogue of species of interest to fisheries 

known to date. FAO species catalogue, vol. 13, 

and FAO Fisheries Synopsis no. 125. Rome: FAO, i– 

vii � 292 pp. 

. 1993a. The Recent genera of the caridean and 

stenopodidean shrimps (Crustacea, Decapoda) with 

an appendix on the order Amphionidacea. Leiden: 

Nationaal Natuurhistorisch Museum, 328 pp. 

. 1993b. The non-Japanese new species established 

by W. de Haan in the Crustacea volume of Fauna 

Japonica (1833–1850). In Ph. F. von Siebold and 

Natural History of Japan, Crustacea, ed. T. Yamaguchi, 

599–642. Tokyo: The Carcinological Society 

of Japan. 

Hou X., and J. Bergström. 1991. The arthropods of the 

Lower Cambrian Chengjiang fauna, with relationships 

and evolutionary significance. In The early evolution 

of Metazoa and the significance of problematic 

taxa, ed. A. M. Simonetta and S. Conway Morris, 

179–188. London: Cambridge University Press. 

. 1997. Arthropods of the Lower Cambrian Chengjiang 

fauna of southwest China. Fossils and Strata 

45:1–116. 

Humes, A. G. 1986. Myicola metisiensis (Copepoda: Poecilostomatoida), 

a parasite of the bivalve Mya arenaria 

in eastern Canada, redefinition of the Myicolidae, 

and diagnosis of the Anthessiidae n. fam. 

Canadian Journal of Zoology 64:1021–1033. 

. 1987. Copepoda from deep-sea hydrothermal 

vents. Bulletin of Marine Science 41:645–788. 

Humes, A. G., and G. A. Boxshall. 1996. A revision of 

the lichomolgoid complex (Copepoda: Poecilostomatoidea), 

with the recognition of six new families. 


Humes, A. G., and J. H. Stock. 1991. Coralliomyzontidae, 

fam. n. (Copepoda: Siphonostomatoida), associated 

with scleractinian corals in Madagascar. Bulletin 

Zoölogisch Museum (Universiteit van Amsterdam) 

13:17–24. 

Huvard, A. 1990. Ultrastructural study of the naupliar eye 

of the ostracode Vargula graminicola (Crustacea, 

Ostracoda). Zoomorphology 110:47–51. 

Huys, R. 1988a. Gelyelloida, a new order of stygobiont 

copepods from European karstic systems. Hydrobiologia 

167/168:485–495. 

. 1988b. On the identity of the Namakosiramiidae 

Ho & Perkins, 1977 (Crustacea, Copepoda), including 

a review of harpacticoid associates of Echinodermata. 


. 1990a. Adenopleurella, new genus, Proceropes, 

new genus, Sarsocletodes Wilson (ex Laophontidae), 

and Miroslavia Apostolov (ex Cletodidae): representatives 

of a new family (Copepoda: Harpacticoida). 


. 1990b. A new harpacticoid copepod family collected 

from Australian sponges and the status of the 

subfamily Rhynchothalestrinae Lang. Zoological 


. 1990c. Amsterdam expeditions to the West Indian 

Islands, Report 64. A new family of harpacticoid co- 


pepods and an analysis of the phylogenetic relationships 

within the Laophontoidea T. Scott. Bijdragen 

tot de Dierkunde 60:79–120. 

. 1990d. Allocation of the Mantridae Leigh-Sharpe 

to the Cyclopoida (Crustacea: Copepoda) with notes 

on Nearchinotodelphys Ummerkutty. Bijdragen tot de 

Dierkunde 60:283–291. 

. 1990e. Campyloxiphos dineti gen. et spec. nov. 

from off Namibia and a redefinition of the Deoterthridae 

Boxshall and Lincoln (Crustacea: Tantulocarida). 


. 1991. Tantulocarida (Crustacea: Maxillopoda): a 

new taxon from the temporary meiobenthos. Marine 

Ecology (Publicazioni della Stazione Zoologica di 

Napoli I) 12:1–34. 

. 1992. The amphiatlantic distribution of Leptastacus 

macronyx (T. Scott, 1892) (Copepoda: Harpacticoida): 

a paradigm of taxonomic confusion; 

and, a cladistic approach to the classification of the 

Leptastacidae Land, 1948. Mededelingen Koninklijke 

Academie voor Wetenschappen, Letteren en 

Schone Kunsten van Belgie 54:21–196. 

. 1993. Styracothoracidae (Copepoda: Harpacticoida), 

a new family from the Philippine deep sea. 


. 1997. Superornatiremidae fam. nov. (Copepoda: 

Harpacticoida): an enigmatic family from North Atlantic 

anchialine caves. Scientia Marina 60:497–542. 

. 2001. Splanchnotrophid systematics: a case of polyphyly 

and taxonomic myopia. Journal of Crustacean 

Biology 21:106–156. 

Huys, R., and R. Böttger-Schnack. 1997. On the diphyletic 

origin of the Oncaeidae Giesbrech, 1892 (Copepoda: 

Poecilostomatoida) with a phylogenetic 

analysis of the Lubockiidae fam. nov. Zoologischer 

Anzeiger 235:243–261. 

Huys, R., and G. A. Boxshall. 1991. Copepod evolution. 

London: The Ray Society, 468 pp. 

Huys, R., G. A. Boxshall, and R. J. Lincoln. 1993. The 

tantulocarid life cycle: the circle closed? Journal of 


Huys, R., and J. M. Gee. 1996. A revision of Danielssenia 

Boeck and Psammis Sars with the establishment of 

two new genera Archisenia and Bathypsammis (Harpacticoida: 

Paranannopidae). Bulletin National 

d’Histoire Naturelle (Zoologie) 59:45–81. 

Huys, R., J. M. Gee, C. G. Moore, and R. Hammond. 

1996. Marine and brackish water harpacticoid copepods. 

Part 1. Keys and notes for identification of 

the species. Synopses of the British Fauna (New Series). 

London: Academic Press, for the Linnean Society 

of London, 352 pp. Huys, R., and T. M. Iliffe. 

1998. Novocriniidae, a new family of harpacticoid 

copepods from anchialine caves in Belize. Zoologica 

Scripta 27:1–15. 

Huys, R., and W. Lee. 1999. On the relationships of the 

Normanellidae and the recognition of Cletopsyllidae 

grad. nov. (Copepoda, Harpacticoida). Zoologischer 

Anzeiger 237:267–290. 

Huys, R., S. Ohtsuka, and G. A. Boxshall. 1994. A new 

tantulocaridan (Crustacea: Maxillopoda) parasitic 

on calanoid, harpacticoid, and cyclopoid copepods. 

Publications of the Seto Marine Biological Laboratory 

36:197–209. 

Huys, R., and K. A. Willems. 1989. Laophontopsis Sars 

and the taxonomic concept of the Normanellinae 

(Copepoda: Harpacticoida): a revision. Bijdragen tot 

de Dierkunde 59:203–227. 

Illg, P. L., and P. L. Dudley. 1980. The family Ascidicolidae 

and its subfamilies (Copepoda, Cyclopoida), 

with descriptions of new species. Memoires du Museum 

National d’Histoire Naturelle, Nouvelle Serie 

A, Zoologie 117:1–192. 

International Commission on Zoological Nomenclature. 

1985a. International Code of Zoological Nomenclature, 

3rd edition. London: The International Trust 

for Zoological Nomenclature and The British Museum 

(Natural History), 338 pp. 

. 1985b. Opinion 1357. Anuropodidae Băcescu, 

1980 (Crustacea, Tanaidacea) and Anuropodidae 

Stebbing, 1893 (Crustacea, Isopoda); a ruling to remove 

the homonymy. Bulletin of Zoological Nomenclature 

42:338–340. 

. 1999. International Code of Zoological Nomenclature, 

4th edition. London: The International Trust 

for Zoological Nomenclature and The Natural History 

Museum, 306 pp. 

International Code of Zoological Nomenclature, 3rd edition, 

adopted by the XX General Assembly of the 

International Union of Biological Sciences. 1985. 

Berkeley, California: University of California Press 

and International Trust for Zoological Nomenclature, 

338 pp. 

Ishimaru, S. 1994. A catalogue of gammaridean and ingolfiellidean 

Amphipoda recorded from the vicinity 

of Japan. Reports of the Seto Marine Biological Laboratory 

24:29–846. 

Itô, T. 1989. Origin of the basis in copepod limbs, with 

reference to remipedian and cephalocarid limbs. 


Itô, T., and F. R. Schram. 1988. Gonopores and the reproductive 

system of nectiopodan Remipedia. Journal 

of Crustacean Biology 8:250–253. 

Ivanenko, V. N., F. D. Ferrari, and A. V. Smurov. 2001. 

Nauplii and copepodids of Scottomyzon gibberum 

(Copepoda: Siphonostomatoida: Scottomyzontidae, 

a new family), a symbiont of Asterias rubens (Asteroida). 

Proceedings of the Biological Society of Washington 

114:237–261. 

Iverson, E. 1982. Revision of the isopod family Sphaeromatidae 

(Crustacea: Isopoda: Flabellifera) I. Subfamily 

names with diagnoses and key. Journal of Crustacean 

Biology 2:248–254. 

Izawa, K. 1996. Archidactylina myxinicola, new genus, 

new species (Siphonostomatoida), in a new family of 

Copepoda parasitic on hagfishes (Agnatha: Myxiniformes) 

from Japan. Journal of Crustacean Biology 

16:406–417. 

Jamieson, B. G. M. 1987. The ultrastructure and phylogeny 

of insect spermatozoa. London: Cambridge University 


. 1989a. The ultrastructure of the spermatozoa of 

four species of xanthid crabs (Crustacea, Brachyura, 

Xanthidae). Journal of Submicroscopic Cytology 

and Pathology 21:589–584. 

. 1989b. Ultrastructural comparison of the spermatozoa 

of Ranina ranina (Oxystomata) and of other 

crabs exemplified by Portunus pelagicus (Brachygnatha) 

(Crustacea, Brachyura). Zoomorphology 

109:103–111. 

. 1989c. A comparison of the spermatozoa of Oratosquilla 

stephensoni and Squilla mantis (Crustacea, 

Stomatopoda) with comments on the phylogeny 

of the Malacostraca. Zoologica Scripta 18:509–517. 

. 1990. The ultrastructure of the spermatozoa of 

Petalomera lateralis (Gray) (Crustacea, Brachyura, 


Dromiacea) and its phylogenetic significance. Invertebrate 

Reproduction and Development 17:39–45. 

. 1991a. Ultrastructure and phylogeny of crustacean 

spermatozoa. In Proceedings of the 1990 International 

Crustacean Conference, ed. P. J. F. Davie 

and R. H. Quinn. Memoirs of the Queensland Museum 

31:109–142. 

. 1991b. Sperm and phylogeny in the Brachyura 

(Crustacea). In Comparative spermatology 20 years 

after, ed. B. Bacetti, 967–972. New York: Serono 

Symposia Publications from Raven Press, vol. 75. 

. 1993. Spermatological evidence for the taxonomic 

status of Trapezia (Crustacea: Brachyura: Heterotremata). 

Memoirs of the Queensland Museum 33: 

225–234. 

. 1994. Phylogeny of the Brachyura with particular 

reference to the Podotremata: evidence from a review 

of spermatozoal ultrastructure. Philosophical 

Transactions of the Royal Society 345B:373–393. 

Jamieson, B. G. M., D. Guinot, and B. Richer de Forges. 

1993a. The ultrastructure of the spermatozoon of 

Paradynomene tuberculata Sakai, 1963 (Crustacea, 

Brachyura, Dynomenidae): synapomorphies with 

dromiid sperm. Helgoländer Meeresuntersuchungen 

47:311–322. 

. 1993b. Spermatozoal ultrastructure in 4 genera 

of Homolidae (Crustacea, Decapoda)—exemplified 

by Homologenus sp., Latreillopsis sp., Homolomannia 

sibogae, and Paromolopsis boasi. Helgoländer 

Meeresuntersuchungen 47:323–334. 

. 1993c. The spermatozoon of Calocarcinus africanus 

(Heterotremata, Brachyura, Crustacea)—ultrastructural 

synapomorphies with xanthid sperm. 

Invertebrate Reproduction and Development 24: 

189–196. 

. 1994a. Relationships of the Cyclodorippoidea 

Ortmann—evidence from spermatozoal ultrastructure 

in the genera Xeinostoma, Tymolus, and Cymonomus 

(Crustacea, Decapoda). Invertebrate Reproduction 


. 1994b. Podotreme affinities of Raninoides sp. and 

Lyreidus brevifrons—evidence from spermatozoal 

ultrastructure (Crustacea, Brachyura, Raninoidea). 


. 1995. Phylogeny of the Brachyura: evidence from 

spermatozoal ultrastructure (Crustacea, Decapoda). 

In Advances in spermatozoal phylogeny and taxonomy, 

ed. B. G. M. Jamieson, J. Ausio, and J.-L. Justine 


vol. 166, 265–283. 

. 1996. Contrasting spermatozoal ultrastructure in 

two thoracotreme crabs, Cardisoma carnifex (Gecarcinidae) 

and Varuna litterata (Grapsidae) (Crustacea: 

Brachyura). Invertebrate Reproduction and 

Development 29:111–126. 

Jamieson, B. G. M., D. Guinot, C. C. Tudge, and B. Richer 

de Forges. 1997. Ultrastructure of the spermatozoa 

of Corystes cassivelaunus (Corystidae), Platepistoma 

nanum (Cancridae), and Cancer pagurus (Cancridae) 

supports recognition of the Corystoidea (Crustacea, 

Brachyura, Heterotremata). Helgoländer 

Meeresuntersuchungen 51:83–93. 

Jamieson, B. G. M., and C. C. Tudge. 1990. Dorippids 

are Heterotremata: evidence from ultrastructure of 

the spermatozoa of Neodorippe astuta (Dorippidae) 

and Portunus pelagicus (Portunidae) (Brachyura: Decapoda). 


Jamieson, B. G. M., C. C. Tudge, and D. M. Scheltinga. 

1993. The ultrastructure of the spermatozoan of 

Dromidiopsis edwardsi Rathbun, 1919 (Crustacea: 

Brachyura: Dromiidae): confirmation of a dromiid 

sperm type. Australian Journal of Zoology 41:537– 

548. 

Jarett, N. E., and E. L. Bousfield. 1994a. The amphipod 

superfamily Phoxocephaloidea on the Pacific Coast 

of North America. Family Phoxocephalidae. Part I. 

Metharpiniinae, new subfamily. Amphipacifica 1: 

58–140. 

. 1994b. The amphipod superfamily Phoxocephaloidea 

on the Pacific Coast of North America. Family 

Phoxocephalidae. Part II. Subfamilies Pontharpiniinae, 

Parharpiniinae, Brolginae, Phoxocephalinae, 

and Harpiniinae. Systematics and distributional ecology. 

Amphipacifica 1:71–150. 

Jarman, S. N., S. Nicol, N. G. Elliott, and A. McMinn. 

2000. 28S rDNA evolution in the Eumalacostraca 

and the phylogenetic position of krill. Molecular 

Phylogenetics and Evolution 17:26–36. 

Jenner, R. A., C. H. J. Hof, and F. R. Schram. 1998. Palaeo- 

and archaeostomatopods (Hoplocarida, Crustacea) 

from the Bear Gulch Limestone, Mississippian 

(Namurain), of central Montana. Contributions to 

Zoology (Amsterdam: SPB Academic Publishing) 67: 

155–185. 

Jensen, P. G., J. T. Høeg, S. Bower, and A. V. Rybakov. 

1994a. Scanning electron microscopy of lattice organs 

in cyprids of the Rhizocephala Akentrogonida 

(Crustacea: Cirripedia). Canadian Journal of Zoology 

72:1018–1026. 

Jensen, P. G., J. Moyse, J. T. Høeg, and H. Al-Yahya. 

1994b. Comparative SEM studies of lattice organs: 

putative sensory structures on the carapace of larvae 

from Ascothoracida and Cirripedia (Crustacea Maxillopoda 

Theocostraca). Acta Zoologica 75:125– 

142. 

Jespersen, A˚ . 1979. Spermiogenesis in two species of Nebalia 

Leach (Crustacea, Malacostraca, Phyllocarida). 

Zoomorphology 93:87–97. 

Jones, M. L. 1961. Lightiella serendipita gen. nov., sp. 

nov., a cephalocarid from San Francisco Bay, California. 


Jones, N. S. 1976. British Cumaceans. Synopses of the 

British Fauna No. 7. London: Academic Press, for 

the Linnean Society of London, 66 pp. 

Just, J. 1990. Abyssianiridae, a synonym of Paramunnidae 

(Crustacea: Isopoda: Asellota) with two new species 

of Abyssianira from south-eastern Australia. Memoirs 

of the Museum of Victoria 50:403–416. 

Just, J., and G. C. B. Poore. 1988. Second record of Hirsutiidae 

(Peracarida: Mictacea): Hirsutia sandersetalia, 

new species, from southeastern Australia. Journal 


. 1992. Vermectiadidae, a new primitive asellote 

isopod family with important phylogenetic implications. 


Kamaltynov, R. M. 1999. On the higher classification of 

Lake Baikal amphipods. Crustaceana 72:933–944. 

Kensley, B. 1998. Estimates of species diversity of freeliving 

isopod crustaceans on coral reefs. Coral Reefs 

17:83–88. 

Kensley, B., M. Schotte, and S. Schilling. 1996–1998. 

World list of marine, freshwater and terrestrial isopod 

crustaceans. URL gopher://nmnhgoph.si.edu:70/ 

11/.invertebrate/.crustaceans; URL http://www. 

nmnh.si.edu/iz/isopod/about.html#org. 

Kim, C. B., and W. Kim. 1993. Phylogenetic relationships 


among gammaridean families and amphipod suborders. 


Kim, W., and L. G. Abele. 1990. Molecular phylogeny of 

selected decapod crustaceans based on 18S rRNA 

nucleotide sequences. Journal of Crustacean Biology 

10:1–13. 

Kitaura, J., K. Wada, and M. Nishida. 1998. Molecular 

phylogeny and evolution of unique mud-using territorial 

behavior in ocypodid crabs (Crustacea: Brachyura: 

Ocypodidae). Molecular Biology and Evolution 

15:626–637. 

Koltzoff, N. K. 1906. Studien über die Gestalt der Zelle. 

I. Untersuchungen über dies Spermien der Decapoden, 

als Einleitung in das Problem der Zellengestalt. 

Archiv für Mikroskopische Anatomomie 67:364– 

572. 

Korn, O. M. 1995. Naupliar evidence for cirripede taxonomy 

and phylogeny. In Crustacean issues 10. New 

frontiers in barnacle evolution, ed. F. R. Schram and 

J. T. Høeg, 87–121. Rotterdam: A. A. Balkema 


Kornicker, L. S. 1986. Sarsiellidae of the western Atlantic 

and northern Gulf of Mexico, and revision of the 

Sarsiellinae (Ostracoda: Myodocopina). Smithsonian 

Contributions to Zoology 415:1–217. 

Kornicker, L. S., and I. G. Sohn. 1976. Phylogeny, ontogeny, 

and morphology of living and fossil Thaumatocypridacea 

(Myodocopa: Ostracoda). Smithsonian 


Kropp, R. K., and R. B. Manning. 1985. Cryptochiridae, 

the correct name for the family containing the gall 

crabs (Crustacea: Decapoda: Brachyura). Proceedings 

of the Biological Society of Washington 98:954– 

955. 

. 1987. The Atlantic gall crabs, family Cryptochiridae 

(Crustacea: Decapoda: Brachyura). Smithsonian 


Kussakin, O. G. 1988. Marine and brackish isopods (Isopoda) 

of cold and temperate waters of the northern 

hemisphere. Volume 3. Suborder Asellota. Part 1. 

Families Janiridae, Santiidae, Dendrotionidae, Munnidae, 

Paramunnidae, Haplomunnidae, Mesosignidae, 

Halponiscidae, Mictosomatidae, Ischnomesidae. 

Opredeliteli Faune SSR 152:1–500. 

Lake, J. A. 1990. Origin of the Metazoa. Proceedings of 

the National Academy of Sciences of the United 

States of America 87:763–766. 

Lamb, E. J., G. A. Boxshall, P. J. Mill, and J. Grahame. 

1996. Nucellicolidae: a new family of endoparasitic 

copepods (Poecilostomatoida) from the dog whelk 

Nucella lapillus (Gastropoda). Journal of Crustacean 

Biology 16:142–148. 

Land, M. F. 1996. Les yeux: structure et fonctionnement 

des mécanismes optiques. In Traité de Zoologie. Anatomie, 

Systématique, Biologie. Crustacés. Tome 



Lange, S., F. R. Schram, C. H. J. Hof, and F. A. Steeman. 

In press. A new genus and species of Thylacocephala, 

with observations that allow assignment of the 

group at last to the Crustacea. Palaeontology. 

Larsen, K., and G. D. F. Wilson. 1998. Tanaidomorphan 

systematics—is it obsolete? Journal of Crustacean 

Biology 18:346–362. 

Latreille, P. A. 1802. Histoire naturelle générale et particulière 

des Crustacés et des Insectes, vol. 3, 1–467. 

(For date of publication, see Dupuis, 1975). 

Laubier, L. 1988. Le genre Anomopsyllus Sars, 1921, co- 

pepode parasite d’annelides polychetes: A. pranizoides 

Sars, 1921 et A. abyssorum nov. sp. Crustaceana 

55:180–192. 

Laubitz, D. R. 1993. Caprellidae (Crustacea: Amphipoda): 

towards a new synthesis. Journal of Natural 

History 27:965–976. 

Leach, W. E. 1815. A tabular view of the external characters 

of four classes of animals, which Linn., arranged 

under Insecta; with the distribution of the 

genera composing three of these classes into orders, 

&c. and descriptions of several new genera and species. 

Transactions of the Linnean Society of London 

11:306–400. 

Lim, G. S. Y., and P. K. L. Ng. 1988. The first zoeal stage 

of Harrovia albolineata Adams and White, 1848 

(Crustacea: Brachyura: Pilumnidae), with a note on 

Eumedonine systematics. Journal of Natural History 

22:217–223. 

Lincoln, R. J., and G. A. Boxshall. 1983. Deep-sea asellote 

isopods of the north-east Atlantic: the family Dendrotionidae 

and some new ectoparasitic copepods. 


318. 

Lopretto, E. C., and J. J. Morrone. 1998. Anaspidacea, 

Bathynellacea (Crustacea, Syncarida), generalised 

tracks, and the biogeographical relationships of 

South America. Zoologica Scripta 27:311–318. 

Lowry, J. K., and A. A. Myers. 2000. A family level 

phylogeny of iphemedioid amphipods (Crustacea, 

Amphipoda). Abstracts of the 10th Colloqium 

on Amphipoda (http://www.odu.edu/%7Ejrh100f/ 

amphome): 14. 

Lowry, J. K., and H. E. Stoddart. 1983. The shallow-water 

gammaridean Amphipoda of the subantarctic islands 

of New Zealand and Australia: Lysianassoidea. Journal 

of the Royal Society of New Zealand 13:279– 

394. 

. 1990. The Wandinidae, a new Indo-Pacific family 

of lysianassoid Amphipoda (Crustacea). Records of 

the Australian Museum 42:159–171. 

. 1995. The Amphipoda (Crustacea) of Madang 

Lagoon: Lysianassidae, Opisidae, Uristidae, Wandinidae 

and Stegocephalidae. In Amphipoda (Crustacea) 

of the Madang Lagoon, Papua New Guinea, ed. 

J. K. Lowry. Records of the Australian Museum, 

22(supplement):97–174. 

. 1996. New lysianassoid amphipod species from 

Namibia and Madagascar (Lysianassidae Dana, 

1849, Podoprionidae Fam. Nov.). Bolletino di Museo 

Civico di Storia Naturale di Verona 20:225–247. 

. 1997. Amphipoda Crustacea IV. Families Aristiidae, 

Cyphocarididae, Endevouridae, Lysianassidae, 

Scopelocheiridae, Uristidae. Memoirs of the Hourglass 

Cruises (Florida Marine Research Institute, St. 

Petersburg, Florida) 10:1–148. 

Lützen, J., and T. Takahashi. 1996. Morphology and biology 

of Polysaccus japonicus (Crustacea, Rhizocephala, 

Akentrogonida, Polysaccidae, fam. N.), a parasite 

of the ghost-shrimp Callianassa japonica. Zoologica 

Scripta 25:171–181. 

Maddison, W. P. 1997. Gene trees in species trees. Systematic 

Biology 46:523–536. 

Magalhães, C., and M. Türkay. 1996a. Taxonomy of the 

Neotropical freshwater crab family Trichodactylidae. 

I. The generic system with description of some 

new genera (Crustacea: Decapoda: Brachyura). 

Senckenbergiana Biologica 75:63–95. 

. 1996b. Taxonomy of the Neotropical freshwater 


crab family Trichodactylidae. II. The genera Forsteria, 

Melocarcinus, Sylviocarcinus and Zilchiopsis 

(Crustacea: Decapoda: Brachyura). Senckenbergiana 

Biologica 75:97–130. 

. 1996c. Taxonomy of the Neotropical freshwater 

crab family Trichodactylidae. III. The genera Fredilocarcinus 

and Goyazana (Crustacea: Decapoda: 

Brachyura). Senckenbergiana Biologia 75:131–142. 

Maley, L. E., and C. R. Marshall. 1998. The coming of 

age of molecular systematics. Science 279:505–506. 

Manning, R. B. 1995. Stomatopod Crustacea of Vietnam: 

the legacy of Raoul Serene. The Carcinological Society 

of Japan. Crustacean Research, Special no. 4: 

viii � 339 pp. 

Manning, R. B., and A. J. Bruce. 1984. Erythrosquilla 

megalops, a remarkable new stomatopod from the 

western Indian Ocean. Journal of Crustacean Biology 

4:329–332. 

Manning, R. B., and D. K. Camp. 1993. Erythrosquilloidea, 

a new superfamily, and Tetrasquillidae, a new 

family of stomatopod crustaceans. Proceedings of 

the Biological Society of Washington 106:85–91. 

Manning, R. B., and D. L. Felder. 1991. Revision of the 

American Callianassidae (Crustacea: Decapoda: 

Thalassinidea). Proceedings of the Biological Society 


. 1996. Nannotheres moorei, a new genus and species 

of minute pinnotherid crab from Belize, Caribbean 

Sea (Crustacea: Decapoda: Pinnotheridae). 


109:311–317. 

Manning, R. B., and L. B. Holthuis. 1981. West African 

brachyuran crabs. Smithsonian Contributions to Zoology 

306:1–379. 

Marchenkov, A. V., and G. A. Boxshall. 1995. A new family 

of copepods associated with ascidiaceans in the 

White Sea, and an analysis of antennulary segmentation 

and setation patterns in the order Poecilostomatoida. 

Zoologisches Anzeiger 234:133–143. 

Margolis, L., T. E. McDonald, and E. L. Bousfield. 2000. 

The whale-lice (Amphipoda: Cyamidae) of the 

northeastern Pacific region. Amphipacifica 2:63– 

117. 

Marincek, M., and B. Petrov. 1991. A review of the Conchostraca 

(Crustacea) of Yugoslavia. Bulletin du 

Muséum d’Histoire Naturelle de Belgrade, series B, 

Sciences Biologiques 39:105–122. 

Martens, K. 1992. On Namibcypris costata n. gen., n. sp. 

(Crustacea, Ostracoda, Candoninae) from a spring 

in northern Namibia, with the description of a new 

tribe and a discussion on the classification of the Podocopina. 

Stygologia 7:27–42. 

Martens, K., D. J. Horne, and H. I. Griffiths. 1998. Age 

and diversity of non-marine ostracodes. In Sex and 

parthenogenesis: evolutionary ecology of reproductive 

modes in non-marine ostracods, ed. K. Martens, 

37–55. Leiden: Backhuys Publishers, 336 pp. 

Martin, J. W. 1984. Notes and bibliography on the larvae 

of xanthid crabs, with a key to the known zoeas of 

the western Atlantic and Gulf of Mexico. Bulletin of 

Marine Science 34:220–239.Martin, J. W. 1988. 

Phylogenetic significance of the brachyuran megalopa: 

evidence from the Xanthidae. Symposium of the 

Zoological Society of London 59:69–102. 

. 1991. Crabs of the family Homolodromiidae, III. 

First record of the larvae. Journal of Crustacean Biology 

11:156–161. 

. 1992. Branchiopoda. In Microscopic anatomy of 

invertebrates, vol. 9, Crustacea, ed. F. W. Harrison 

and A. G. Humes, 25–224. New York: Wiley-Liss. 

. 1995. Review of: Walossek, D. 1993. The Upper 

Cambrian Rehbachiella and the phylogeny of Branchiopoda 

and Crustacea. Earth-Science Reviews 38: 

72–75. 

. 1996. The Crustacean biodiversity survey. Program 

and Abstracts, The Crustacean Society 1996 

Summer Meeting, San Diego, California: 28. 

Martin, J. W., and L. G. Abele. 1986. Phylogenetic relationships 

of the genus Aegla (Decapoda: Anomura: 

Aeglidae), with comments on anomuran phylogeny. 


Martin, J. W., and D. Belk. 1988. Review of the clam 

shrimp family Lynceidae Stebbing, 1902 (Branchiopoda: 

Conchostraca), in the Americas. Journal of 


Martin, J. W., and C. Cash-Clark. 1995. The external 

morphology of the onychopod ‘cladoceran’ genus 

Bythotrephes (Crustacea, Branchiopoda, Onychopoda, 

Cercopagididae), with notes on the morphology 

and phylogeny of the order Onychopoda. Zoologica 

Scripta 24:61–90. 

Martin, J. W., and J. C. Christiansen. 1995a. A morphological 

comparison of the phyllopodous thoracic 

limbs of a leptostracan (Nebalia sp.) and a spinicaudate 

conchostracan (Leptestheria sp.), with comments 

on the use of Phyllopoda as a taxonomic category. 

Canadian Journal of Zoology 73:2283–2291. 

. 1995b. A new species of the shrimp genus Chorocaris 

Martin and Hessler, 1990 (Crustacea: Decapoda: 

Bresiliidae) from hydrothermal vent fields 

along the Mid-Atlantic Ridge. Proceedings of the Biological 


Martin, J. W., and S. Trautwein. In press. Fossil crabs 

(Crustacea, Decapoda, Brachyura) from Lothagam. 

In Lothagam: the dawn of humanity in eastern Africa, 

ed. M. G. Leakey and J. M. Harris. New York: 

Columbia University Press. 

Martin, J. W., F. M. Truesdale, and D. L. Felder. 1985. 

Larval development of Panopeus bermudensis Benedict 

and Rathbun, 1891 (Brachyura, Xanthidae) 

with notes on zoeal characters in xanthid crabs. 


Martin, J. W., E. W. Vetter, and C. E. Cash-Clark. 1996. 

Description, external morphology, and natural history 

observations of Nebalia hessleri, new species 

(Phyllocarida: Leptostraca), from southern California, 

with a key to the extant families and genera of 

the Leptostraca. Journal of Crustacean Biology 16: 

347–372. 

Mattern, D., and M. Schlegel. 2001. Molecular evolution 

of the small subunit ribosomal DNA in woodlice 

(Crustacea, Isopoda, Oniscidea) and implications for 

oniscidean phylogeny. Molecular Phylogenetics and 


Mayrat, A., and M. de Saint Laurent. 1996. Classe des 

Malacostracés (Malacostraca Latreille, 1802). Considerations 

sur la classe des Malacostracés. In Traité 

de Zoologie. Anatomie, Systématique, Biologie. 

Crustacés. Tome VII, Fascicule II. Généralités (suite) 

et Systématique, ed. J. Forest, 841–863. Paris: Masson, 

1002 pp. 

McHugh, D., and K. M. Halanych. 1998. Introduction to 

the symposium: evolutionary relationships of Metazoan 

phyla: advances, problems, and approaches. 

American Zoologist 38:813–817. 

McKenzie, K. G. 1991. Crustacean evolutionary sequenc- 


es and consequences. In Proceedings of the 1990 International 

Crustacean Conference, ed. P. J. F. Davie 

and R. H. Quinn. Memoirs of the Queensland Museum 

31:19–38. 

McKenzie, K. G., K. J. Müller, and M. N. Gramm. 1983. 

Phylogeny of Ostracoda. In Crustacean issues 1. 

Crustacean phylogeny, ed. F. R. Schram, 29–46. Rotterdam: 

A. A. Balkema Press, 372 pp. 

McKinnon, A. D. 1994. Australopsyllus fallaz gen. et sp. 

nov., the third known species of the family Thaumatopsyllidae 

(Copepoda: Cyclopoida). Sarsia 79: 

27–32. 

McLaughlin, P. A. 1983a. A review of the phylogenetic 

position of the Lomidae (Crustacea: Decapoda: Anomala). 


. 1983b. Hermit crabs—are they really polyphyletic? 


McLaughlin, P. A., and P. Hogarth. 1998. Hermit crabs 

(Decapoda: Anomura: Paguridea) from the Seychelles. 

Zoologisches Mededelingen 318:1–48. 

McLaughlin, P. A., and L. B. Holthuis. 1985. Anomura 

versus Anomala. Crustaceana 49:204–209. 

McLaughlin, P. A., and R. Lemaitre. 1997. Carcinization 

in the Anomura—fact or fiction? I. Evidence from 

adult morphology. Contributions in Zoology (Amsterdam) 

67:79–123. 

. 2000. Aspects of evolution in the anomuran superfamily 

Paguroidea: one larval prospective. Invertebrate 

Reproduction and Development 38:159– 

169. 

McLay, C. L. 1991. Crustacea Decapoda: the sponge 

crabs (Dromiidae) of New Caledonia and the Philippines 

with a review of the genera. In Résultats des 

Campagnes Musorstom, ed. A. Crosnier. Mémoires 

de Muséum National d’Histoire Naturelle, vol. 10, 

no. 156, 11–251. 

. 1999. Crustacea Decapoda: revision of the family 

Dynomenidae. In Résultats des Campagnes Musorstom, 

ed. A. Crosnier. Mémoires de Muséum National 

d’Histoire Naturelle, vol. 20, no. 180, 427– 

569. 

Meier, R., and S. Richter. 1992. Suggestions for a more 

precise usage of proper names of taxa. Zeitschrift für 

Zoologische Systematik und Evolutionsforschung 

30:81–88. 

Milius, S. 1999. Should we junk Linnaeus? Science News 

156:268–270. 

Min, G. S., S. H. Kim, and W. Kim. 1998. Molecular phylogeny 

of arthropods and their relatives: polyphyletic 

origin of arthropodization. Molecules and Cells 8: 

75–83. 

Mizrahi, L., Y. Achituv, D. J. Katcoff, and R. Perl-Treves. 

1998. Phylogenetic position of Ibla (Cirripedia: 

Thoracica) based on 18S rDNA sequence analysis. 


Monod, Th. 1984. La position systématique des Thermosbaenacea. 

Annales de Societe Royale de Zoologie 

Belgique 114(supplement 1):204–206. 

Monod, Th., and Ph. Cals. 1988. Systématique et évolution 

des Thermosbénacés (Arthropodes, Crustacés), 

d’après l’ordonnance des structures épidermiques superficielles. 

Comptes Rendus Hebdomadaires des Seances 

de l’Academie des Sciences, series 3, 306:99– 

108. 

. 1999. Ordre des Thermosbaenacés (Thermosbaenacea 

Monod, 1927). In Traité de Zoologie. Anatomie, 





Monod, Th., and J. Forest. 1996. Histoire de la classification 

des Crustacés. In Traité de Zoologie. Anatomie, 

Systématique, Biologie. Crustacés. Tome VII, 

Fascicule II. Généralités (suite) et Systématique, ed. 

J. Forest, 235–267. Paris: Masson, 1002 pp. 

Monod, Th., and L. Laubier. 1996. Les Crustacés dans la 

Biosphere. In Traité de Zoologie. Anatomie, Systématique, 


Généralités (suite) et Systématique, ed. J. Forest, 91– 

166. Paris: Masson, 1002 pp. 

Moore, R. C. 1969. Editorial preface. In Treatise on invertebrate 

paleontology, part R, Arthropoda 4, ed. 

R. C. Moore, xi–xxxvi. Lawrence, Kansas: Geological 


Press. 

Moore, R. C., and McCormick. 1969. General features of 

Crustacea. In Treatise on invertebrate paleontology, 

part R, Arthropoda 4, ed. R. C. Moore, R57—R120. 

Lawrence, Kansas: Geological Society of America 

and the University of Kansas Press. 

Moosa, M. K. 1991. The Stomatopoda of New Caledonia 

and Chesterfield Islands. In Le benthos des fonds 

meubles des lagons de Nouvelle-Calédonie, vol. 1, 

ed. B. Richer de Forges, 149–219. Paris: ORSTOM 

Editions. 

Morin, J. G., and A. C. Cohen. 1991. Bioluminescent displays, 

courtship, and reproduction in ostracodes. In 

Crustacean sexual biology, ed. R. T. Bauer and J. W. 

Martin, 1–16. New York: Columbia University 

Press. 

Morrison, C. L., and C. W. Cunningham. 1999. Dramatic 

mitochondrial DNA gene rearrangements clarify relationships 

among anomuran crustaceans. Program 

and Abstracts, The Crustacean Society 1999 Summer 


Moura, G., and M. L. Christoffersen. 1996. The system 

of the mandibulate arthropods: Tracheata and Remipedia 

as sister groups, ‘‘Crustacea’’ non-monophyletic. 

Journal of Comparative Biology 1:95–113. 

Müller, H.-G. 1994. World catalogue and bibliography of 

the Recent Stomatopoda. Berlin: Wissenschaftlicher 

Verlag, 228 pp. 

Müller, K. J. 1982. Hesslandona unisulcata sp. nov. (Ostracoda) 

with phosphatized appendages from Upper 

Cambrian ‘Orsten’ of Sweden. In A research manual 

of fossil and Recent ostracods, ed. R. H. Bate et al., 

276–307. Chichester, England: Ellis Horwood. 

. 1983. Crustaceans with preserved soft parts from 

the Upper Cambrian of Sweden. Lethaia 16:93–109. 

Müller, K. J., and D. Walossek. 1985a. A remarkable arthropod 

fauna from the Upper Cambrian ‘‘Orsten’’ 

of Sweden. Transactions of the Royal Society of Edinburgh, 

Earth Sciences 76:161–172. 

. 1985b. Skaracarida, a new order of Crustacea 

from the Upper Cambrian of Västergötland, Sweden. 

Fossils and Strata 17:1–65. 

. 1986a. Martinssonia elongata gen. et. sp. n., a 

crustacean-like euarthropod from the Upper Cambrian 

‘Orsten’ of Sweden. Zoologica Scripta 15:73– 

92. 

. 1986b. Arthropod larvae from the Upper Cambrian 

of Sweden. Transactions of the Royal Society 

of Edinburgh, Earth Sciences 77:157–179. 

. 1988. External morphology and larval development 

of the Upper Cambrian maxillopod Bredocaris 

admirabilis. Fossils and Strata 23:1–70. 


Negrea, S., N. Botnariuc, and H. J. Dumont. 1999. Phylogeny, 

evolution and classification of the Branchiopoda 

(Crustacea). Hydrobiologia 412:191–212. 

Newman, W. A. 1982. Evolution within the Crustacea, 

part 3: Cirripedia. In The biology of Crustacea, vol. 

1, Systematics, the fossil record, and biogeography, 

ed. L. G. Abele, 197–221. New York: Academic 

Press. 

. 1983. Origin of the Maxillopoda; urmalacostracan 

ontogeny and progenesis. In Crustacean issues 

1. Crustacean phylogeny, ed. F. R. Schram, 105–119. 


. 1987. Evolution of cirripedes and their major 

groups. In Crustacean issues 5. Barnacle biology, ed. 

A. J. Southward, 3–42. Rotterdam: A. A. Balkema 


. 1989. Juvenile ontogeny and metamorphosis in 

the most primitive living sessile barnacle, Neoverruca, 

from abyssal hydrothermal springs. Bulletin of 

Marine Science 45:467–477. 

. 1991. Cirripedia. In Encyclopedia Britannica, 

15th edition, vol. 16, 849–854, 859. Chicago: Encyclopedia 

Britannica. 

. 1992. Origin of Maxillopoda. Acta Zoologica 73: 

271–392. 

. 1993. Darwin and cirripedology. In Crustacean 

issues 8. History of carcinology, ed. F. M. Truesdale, 


. 1996. Sous-classe de Cirripedes (Cirripedia Burmeister, 

1834) super-ordre des Thoraciques et des 

Acrothoraciques (Thoracica Darwin, 1854—Acrothoracica 

Gruvel, 1905). In Traité de Zoologie. Anatomie, 




Newman, W. A., and R. R. Hessler. 1989. A new abyssal 

hydrothermal verrucomorphan (Cirripedia; Sessilia): 

the most primitive living sessile barnacle. Transactions 

of the San Diego Society of Natural History 

21:259–273. 

Newman, W. A., and T. Yamaguchi. 1995. A new sessile 

barnacle (Cirripedia: Balanomorpha) from the Lau 

Back-Arc Basin, Tonga: first record of a living representative 

since the Miocene. Bulletin du Museum 

National d’Histoire Naturelle (Paris) 17A:211–243. 

Ng, P. K. L. 1983. Aspects of the systematics of the family 

Pilumnidae Samouelle, 1819 (Crustacea, Decapoda, 

Brachyura) and a study of evolutionary trends within 

the superfamily Xanthoidea (sensu Guinot, 1978). 

Unpublished B. Sc. Honors Thesis, Department of 

Zoology, National University of Singapore, 251 pp, 

12 plates. 

. 1988. The freshwater crabs of peninsular Malaysia 

and Singapore. Singapore: Department of Zoology, 

University of Singapore, and Shing Lee Publishers, 

156 pp. 

. 1998. Marine crabs. In FAO species identification 

guide for fishery purposes. The living marine resources 

of the Western-Central Pacific, vol. 2. Cephalopods, 

crustaceans, holothurians and sharks, ed. 

K. E. Carpenter and V. H. Niem, 1045–1155. Rome: 

Food and Agriculture Organisation. 

Ng, P. K. L, and P. F. Clark. 1999. The systematic position 

of the Eumedonidae: evidence from adult and larval 

morphology. In General information and book of 

abstracts, 7th Colloquium Crustacea Decapoda 

Mediterranea, ed. J. Paula. Lisbon, Portugal: University 

of Lisbon, 179 pp. 

. 2000. The eumedonid file: a case study of systematic 

compatibility using larval and adult characters 

(Crustacea: Decapoda: Brachyura). Invertebrate Reproduction 


Ng, P. K. L., and G. Rodriguez. 1986. New records of 

Mimilambrus wileyi Williams, 1979 (Crustacea: Decapoda: 

Brachyura), with notes on the systematics of 

the Mimilambridae Williams, 1979 and Parthenopoidea 

MacLeay, 1838 sensu Guinot, 1978. Proceedings 

of the Biological Society of Washington 99: 

88–99. 

Ng, P. K. L., and B. Sket. 1996. The freshwater crab fauna 

(Crustacea: Brachyura) of the Philippines. IV. On a 

collection of Parathelphusidae from Bohol. Proceedings 

of the Biological Society of Washington 109: 

695–706. 

Ng, P. K. L., and M. Takeda. 1994. Skelosothelphusa 

(Crustacea, Decapoda, Brachyura), a new genus of 

potamonautid freshwater crab from Madagascar, 

with descriptions of two new species. Bulletin of the 

National Science Museum, Tokyo, series A (Zoology) 

20:161–172. 

Nicholls, G. E. 1943. The Phreatoicoidea: part I. The Amphisopidae. 

Papers and Proceedings of the Royal Society 

of Tasmania, 1942:1–145. 

. 1944. The Phreatoicoidea: part II. The Phreatocoidae. 

Papers and Proceedings of the Royal Society 

of Tasmania, 1943:1–156. 

Nicol, S., and Y. Endo. 1999. Krill fisheries: development, 

management and ecosystem implications. Aquatic 

Living Resources 12:105–120. 

Nielsen, C. 1995. Animal evolution: interrelationships of 

the living phyla. Oxford: Oxford University Press, 

467 pp. 

Nielsen, C., N. Scharff, and D. Eibye-Jacobsen. 1996. Cladistic 

analysis of the animal kingdom. Biological 


Nielsen, C., W. F. Walker, H. R. Bode, and R. E. Steele. 

1989. Phylogeny and molecular data (published as 

three separate comments on the paper by Field et al., 

1988). Science 243:548–551. 

Nixon, K., and J. Carpenter. 2000. On the other ‘‘phylogenetic 

systematics.’’ Cladistics 16:298–318. 

Nouvel, H., J.-P. Casanova, and J.-P. Lagardère. 1999. Ordre 

des Mysidacés (Mysidacea Boas 1883). In Traité 


Tome VII, Fascicule IIIA. Crustacés Péracarides, ed. 

J. Forest. Memoires de l’Institut Oceanographique 

Fondation Albert I er , Prince de Monaco, 19:39–86. 

Ohtsuka, S., H. S. J. Roe, and G. A. Boxshall. 1993. A 

new family of calanoid copepods, the Hyperbionychidae, 

collected from the deep-sea hyperbenthic 

community in the northeastern Atlantic. Sarsia 78: 

69–82. 

Olesen, J. 1996. External morphology and phylogenetic 

significance of the dorsal/neck organ in the Conchostraca 

and the head pores of the cladoceran family 

Chydoridae (Crustacea, Branchiopoda). Hydrobiologia 

330:213–226. 

. 1998. A phylogenetic analysis of the Conchostraca 

and Cladocera (Crustacea, Branchiopoda, Diplostraca). 

Zoological Journal of the Linnean Society 

122:491–536. 

. 1999a. Larval and post-larval development of the 

branchiopod clam shrimp Cyclestheria hislopi 

(Baird, 1859) (Crustacea, Branchiopoda, Conchostraca, 

Spinicaudata). Acta Zoologica 80:163–184. 

. 1999b. A new species of Nebalia (Crustacea, Lep- 


tostraca) form Unguja Island (Zanzibar), Tanzania, 

East Africa, with a phylogenetic analysis of leptostracan 

genera. Journal of Natural History 33:1789– 

1809. 

. 2000. An updated phylogeny of the Conchostraca–Cladocera 

clade (Branchiopoda, Diplostraca). 


. 2001. External morphology and larval development 

of Derocheilocaris remanei Delamare-Debouteville 

& Chapuis, 1951 (Crustacea, Mystacocarida), 

with a comparison of crustacean segmentation and 

tagmosis patterns. Biologiske Skrifter 53:1–59. 

Olesen, J., J. W. Martin, and E. Roessler. 1997. External 

morphology of the male of Cyclestheria hislopi 

(Baird, 1859) (Crustacea, Branchiopoda, Spinicaudata), 

with comparison of male claspers among the 

Conchostraca and Cladocera and its bearing on phylogeny 

of the ‘‘bivalved’’ Branchiopoda. Zoologica 

Scripta 25:291–316. 

Orr, P. J., and D. E. G. Briggs. 1999. Exceptionally preserved 

conchostracans and other crustaceans from 

the Upper Carboniferous of Ireland. Special Papers 

in Paleontology (The Paleontology Association of 

London) 62:1–68. 

Overstreet, R. M., I. Dyková, and W. E. Hawkins. 1992. 

Branchiura. In Microscopic anatomy of invertebrates, 

vol. 9, Crustacea, ed. F. W. Harrison and A. 

G. Humes, 385–413. New York: Wiley-Liss. 

Page, R. D. M., and M. A. Charleston. 1998. From gene 

to organismal phylogeny: reconciled trees and the 

gene tree/species tree problem. Molecular Phylogenetics 

and Evolution 7:231–240. 

Palmer, A. R. 1993. Whither Pentastomida? Nature 361: 

214. 

Panganiban, G., S. M. Irvine, C. Lowe, H. Roehl, L. S. 

Corley, B. Sherbon, J. K. Grenier, J. F. Fallon, J. Kimble, 

M. Walker, G. A. Wray, B. J. Swalla, M. Q. 

Martindale, and S. B. Carroll. 1997. The origin and 

evolution of animal appendages. Proceedings of the 

National Academy of Sciences 94:5162–5166. 

Panganiban, G., A. Sebring, L. Nagy, and S. Carroll. 1995. 

The development of crustacean limbs and the evolution 

of arthropods. Science 270:1363–1366. 

Park, T. 1986. Phylogeny of calanoid copepods. Syllogeus 

58:191–196. 

Paul, D. H. 1989. A neurophylogenists view of decapod 

Crustacea. Bulletin of Marine Science 45:487–504. 

Pennant, T. 1777. The British zoology, vol. 4, 4th edition, 

Crustacea, Mollusca, Testacea. London: B. White, 

136 pp. 

Pérez Farfante, I., and B. F. Kensley. 1997. Penaeoid and 

sergestoid shrimps and prawns of the world. Keys 

and diagnoses for the families and genera. Mémoires 

du Muséum National d’Histoire Naturelle, vol. 175, 

1–233. 

Perl-Treves, R., L. Mizrahi, D. J. Katcoff, and Y. Achituv. 

2000. Elucidation of the phylogenetic relationship of 

three thecostracans, Verruca, Paralepas, and Dendrogaster 

based on 18S rDNA sequence. Journal of 


Pinna, G., P. Arduini, C. Pesarini, and G. Teruzzi. 1982. 

Thylacocephala: una nuova classe di Crostacei fossili. 

Atti della Societá Italiano di Scienze Naturali e 

del Museo Civico di Storia Naturale Milano 123: 

469–482. 

. 1985. Some controversial aspects of the morphology 

and anatomy of Ostenocaris cypriformis 

(Crustacea, Thylacocephala). Transactions of the 

Royal Society of Edinburgh 76:373–379. 

Pires, A. M. S. 1987. Potiicoara brasiliensis: a new genus 

and species of Spelaeogriphacea (Crustacea: Peracarida) 

from Brazil with a phylogenetic analysis of the 

Peracarida. Journal of Natural History 21:225–238. 

Pirlot, J. M. 1934. Les amphipodes de l’expedition du Siboga. 

Deuxieme partie, II. Les amphipodes de lar 

mer profonde. 2. Hyperiopsidae, Jassidae. Siboga 

Expedite 33d:167–235. 

Pleijel, F., and G. W. Rouse. 2000. Least-inclusive taxonomic 

unit: a new taxonomic concept for biology. 

Proceedings of the Royal Society of London 267B: 

627–630. 

Pohle, G., and F. Marques. Larval stages of Paradasygyius 

depressus (Bell, 1835) (Crustacea: Decapoda: Brachyura: 

Majidae) and a phylogenetic analysis for 21 

genera of Majidae. Proceedings of the Biological Society 


Poore, G. C. B. 1987. Lynseiidae (Isopoda, Flabellifera), 

a new monotypic family from Australia. Journal of 


. 1994. A phylogeny of the families of Thalassinidea 

(Crustacea: Decapoda) with keys to families and 

genera. Memoirs of the Museum of Victoria 54:79– 

120. 

. 1997. A review of the thalassinidean families Callianideidae 

Kossman, Micheleidae Sakai, and Thomassiniidae 

de Saint Laurent (Crustacea, Decapoda) 

with descriptions of fifteen new species. Zoosystema 

19:345–420. 

. 1998. Phylogeny of the Anthuridea (Isopoda). 



. 2001a. Families and genera of Isopoda Anthuridea. 

In Crustacean issues 13. Isopod systematics and 

evolution, ed. B. Kensley and R. C. Brusca, 63–174. 


. 2001b. Isopoda Valvifera: diagnoses and relationships 

of the families. Journal of Crustacean Biology 

21:205–230. 

Poore, G. C. B., and T. M. Bardsley. 1992. Austrarcturellidae 

(Crustacea: Isopoda: Valvifera), a new family 

from Australia. Invertebrate Taxonomy 6:843–908. 

Poore, G. C. B., and W. F. Humphreys. 1998. First record 

of Spelaeogriphacea from Australasia: a new genus 

and species from an aquifer in the arid Pilbara of 

Western Australia. Crustaceana 71:721–742. 

Poore, G. C. B., and H. M. Lew Ton. 1988. Antheluridae, 

a new family of Crustacea (Isopoda: Anthuridea) 

with new species from Australia. Journal of Natural 

History 22:489–506. 

Popadić, A., D. Rusch, M. Peterson, B. T. Rogers, and T. 

C. Kaufman. 1996. Origin of the arthropod mandible. 

Nature 380:395. 

Por, F. D. 1984. Canuellidae Lang (Harpacticoida: Polyarthra) 

and the ancestry of the Copepoda. Crustaceana 

7(supplement):1–24. 

. 1986. A re-evaluation of the family Cleotodidae 

Sars, Lang (Copepoda, Harpacticoida). Syllogeus 58: 

420–425. 

Pretzmann, G. 1973. Grundlagen und Ergebnisse der Systematik 

der Pseudothelphusidae. Zeitschrift für 

Zoologische Systematik und Evolutionsforschung 

11:196–218. 

Preuss, G. 1951. Die Verwandtschaft der Anostraca und 

Phyllopoda. Zoologisches Anzeiger 147:50–63. 

Raff, R. A., C. R. Marshall, and J. M. Turbeville. 1994. 


Using DNA sequences to unravel the Cambrian radiation 

of the animal phyla. Annual Review of Ecology 

and Systematics 25:351–375. 

Rafinesque, C. S. 1815. Analyse de la nature ou tableau 

de l’univers et des corps organisés. Palermo: 

l’Imprimerie de Jean Barravecchia, 224 pp. 

Raibaut, A. 1996. Copépodes II. Les Copépodes parasites. 

In Traité de Zoologie. Anatomie, Systématique, Biologie. 




Razouls, C. 1996. Copépodes I. Les Copépodes libres. In 

Traité de Zoologie. Anatomie, Systématique, Biologie. 




Razouls, C., and A. Raibaut. 1996. Copépodes III. Phylogénie 

et classification. In Traité de Zoologie. Anatomie, 




Regier, J. C., and J. W. Shultz. 1997. Molecular phylogeny 

of the major arthropod groups indicates polyphyly 

of crustaceans and a new hypothesis for the origin 

of hexapods. Molecular Biology and Evolution 14: 

902–913. 

. 1998a. Resolving arthropod phylogeny using 

multiple nuclear genes. American Zoologist 37: 

102A. 

. 1998b. Molecular phylogeny of arthropods and 

the significance of the Cambrian ‘‘explosion’’ for molecular 

systematics. American Zoologist 38:918– 

928. 

Remigio, E. A., and P. D. Hebert. 2000. Affinities among 

anostracan (Crustacea: Branchiopoda) families inferred 

from phylogenetic analyses of multiple gene 

sequences. Molecular Phylogenetics and Evolution 

17:117–128. 

Ren, X.-Q. 1999. A new family of superfamily Haustorioidea 

(Crustacea: Amphipoda: Gammaridea) from 

the China sea. Chinese Journal of Oceanology and 

Limnology 17:344–349. 

Rice, A. L. 1980. Crab zoeal morphology and its bearing 

on the classification of the Brachyura. Transactions 

of the Zoological Society of London 35:271–424. 

. 1981. The megalopa stage in brachyuran crabs. 

The Podotremata Guinot. Journal of Natural History 

15:1003–1011. 

. 1983. Zoeal evidence for brachyuran phylogeny. 

In Crustacean issues 1. Crustacean phylogeny, ed. F. 

R. Schram, 313–329. Rotterdam: A. A. Balkema 


. 1988. The megalopa stage in majid crabs, with a 

review of spider crab relationships based on larval 

characters. Symposium of the Zoological Society of 

London 59:27–46. 

Richer de Forges, B., B. G. M. Jamieson, D. Guinot, and 

C. C. Tudge. 1997. Ultrastructure of the spermatozoan 

of Hymenosomatidae (Crustacea: Brachyura) 

and the relationships of the family. Marine Biology 

130:233–242. 

Richter, S. 1994. Monophylie der Lophogastrida und Phylogenetisches 

System der Peracarida (Crustacea). 

Verhandlungen der Deutschen Zoologischer Gesellschaft 

87:231. 

. 1999. The structure of the ommatidia of the Malacostraca 

(Crustacea)—a phylogenetic approach. 

Verhandlungen der Naturwissenschaftlichen Vereine 

in Hamburg 38:161–204. 

Richter, S., A. Braband, N. Aladin, and G. Scholtz. 2001. 

The phylogenetic relationships of ‘‘predatory waterfleas’’ 

(Cladocera: Onychopoda, Haplopoda) inferred 

from 12S rDNA. Molecular Phylogenetics and 


Richter, S., and G. Scholtz. 1994. Morphological evidence 

for a hermit crab ancestry of lithodids (Crustacea, 

Decapoda, Anomala, Paguroidea). Zoologische Anzeiger 

233:187–210. 

. In press. Phylogenetic analysis of the Malacostraca 

(Crustacea). Journal of Zoological Systematics 

and Evolutionary Research. 

Riley, J. 1986. The biology of pentastomids. In Advances 

in parasitology, vol. 25, 45–128. London and New 


Riley, J., A. A. Banaja, and J. L. James. 1978. The phylogenetic 

relationships of the Pentastomida: the case 

for their inclusion within the Crustacea. International 

Journal for Parasitology 8:245–254. 

Rodríguez, G. 1982. Les crabes d’eau douce d’Amerique. 

Familie des Pseudothelphusidae. Paris: ORSTOM, 

Faune Tropicale, vol. 22, 223 pp. 

. 1986. Centers of radiation of freshwater crabs in 

the Neotropics. In Crustacean issues 4. Crustacean 

biogeography, ed. R. H. Gore and K. L. Heck, 51– 

67. Rotterdam: A. A. Balkema Press, 292 pp. 

. 1992. The freshwater crabs of America. Family 

Trichodactylidae and a supplement to the family 

Pseudothelphusidae. Paris: ORSTOM, Faune Tropicale, 

vol. 31, 189 pp. 

Roessler, E. W. 1991. Estudios sobre Entomostraces de 

Colombia VI—Paraimnadiidae, una nueva familia 

de Crustacea—Conchostraca. Revista de la Academia 

Colombiana de Ciencias Exactas, Físicas y Naturales 

18:93–104. 

. 1995a. Review of Colombian conchostraca (Crustacea)—morphotaxonomic 

aspects. Hydrobiologia 

298 (Developmental Hydrobiology 103):253–262. 

. 1995b. Review of Colombian conchostraca 

(Crustacea)—ecological aspects and life cycles—families 

Lynceidae, Limndadiidae, Leptestheriidae, and 

Metalimnadiidae. Hydrobiologia 298 (Developmental 

Hydrobiology 103):113–124. 

Rolfe, W. D. I. 1981. Phyllocarida and the origin of the 

Malacostraca. Geobios 14:17–27. 

. 1985. Form and function in the Thylacocephala, 

Conchyliocarida and Concavicarida (?Crustacea): a 

problem of interpretation. Transactions of the Royal 

Society of Edinburgh, Earth Sciences, 76:391–399. 

. 1992. Not yet proven Crustacea: the Thylacocephala. 


Roman, M.-L., and H. Dalens. 1999. Ordre des Isopodes 

(Épicarides exclus) (Isopoda Latreille, 1817). In Traité 


Tome VII, Fascicule IIIA. Crustacés Péracarides, ed. 

J. Forest. Memoires de l’Institut Oceanographique 

Fondation Albert I er , Prince de Monaco, 19:177– 

278. 

Roush, W. 1995. Gene ties arthropods together. Science 

270:1297. 

Rudolphi, K. A. 1809. Entozoorum, sive Vermium Intestinalis 

Historia naturalis 2 (1). 

Ruppert, E. E., and R. D. Barnes. 1994. Invertebrate zoology, 

6th edition. Orlando, Florida: Saunders College 

Publishing and Harcourt Brace Jovanovich, 

1056 pp. 


Saint Laurent, M. de. 1979. Vers une nouvelle classification 

des Crustaces Decapodes Reptantia. Bulletin de 

l’Office National des Pêches République Tunisienne 

(Tunisia), Ministere de l’Agriculture 3:15–31. 

. 1980a. Sur classification et phylogenie des Crustaces 

Decapodes brachyoures. I. Podotremata Guinot, 

1977, et Eubrachyura sect. nov. Comptes Rendus 

Hebdomadaires des Seances de l’Academie des Sciences, 

series D, 290:1265–1268. 

. 1980b. Sur classification et phylogenie des Crustaces 

Decapodes brachyoures. II. Heterotremata et 

Thoracotremata Guinot, 1977. Comptes Rendus 


series D, 290:1317–1320. 

. 1985. Remarques sur la distribution des Crustacés 

Décapodes. In Peuplements profonds du Golfe de 

Gascogne, Campagnes Biogas, ed. L. Laubier and C. 

Monniot, 469–478. Ifremer: Institut Francaise de 

Rechereche pour l’Exploraion de la Mer. 

. 1988. Enoplometopoidea, nouvelle superfamille 

de Crustacés Décapodes Astacidea. Comptes Rendus 


series 3, 307:59–62. 

. 1989. La nouvelle superfamille des Retroplumoidea 

Gill, 1894 (Decapoda, Brachyura): systématique, 

affinités et évolution. In Résultats des Campagnes 

MUSORSTOM, vol. 5, ed. J. Forest. Mémoires 

du Muséum National d’Histoire Naturelle 

(Paris), sére A, vol. 144, 103–179. 

Sakai, K. 1992. The families Callianideidae and Thalassinidae, 

with the description of two new subfamilies, 

one new genus and two new species. Naturalists 4: 

1–33. 

. 1999. Synopsis of the family Callianassidae, with 

keys to subfamilies, genera and species, and the description 

of new taxa (Crustacea: Decapoda: Thalassinidea). 

Zoologische Verhandelingen (Leiden) 

326:1–152. 

Samouelle, G. 1819. The entomologist’s useful compendium; 

or an introduction to the knowledge of British 

insects, comprising the best means of obtaining and 

preserving them, and a description of the apparatus 

generally used; together with the genera of Linné, 

and the modern method of arranging the classes 

Crustacea, Myriapoda, Spiders, Mites and Insects, 

from their affinities and structure, according to the 

views of Dr. Leach, Also an explanation of the terms 

used in entomology; a calendar of the times of appearance 

and usual situations of near 3,000 species 

of British insects; with instructions for collecting and 

fitting up objects for the microscope. London: 

Thomas Boys, 496 pp. 

Sanders, H. L. 1963. Significance of the Cephalocarida. In 

Phylogeny and evolution of Crustacea, ed. H. B. 

Whittington and W. D. I Rolfe, 163–175. Cambridge, 

Massachusetts: Special Publication of the 

Museum of Comparative Zoology. 

Sanders, H. L., R. R. Hessler, and S. P. Garner. 1985. Hirsutia 

bathyalis, a new unusual deep-sea benthic peracaridan 

crustacean from the tropical Atlantic. Journal 


Sassaman, C. 1992. Description of the mature female and 

epicaridium lava of Cabiropsis montereyensis Sassaman 

from southern California (Crustacea: Isopoda: 

Cabiropidae). Proceedings of the Biological Society 


. 1995. Sex determination and evolution of unisex- 

uality in the Conchostraca. Hydrobiologia 298:45– 

65. 

Schmalfuss, H. 1989. Phylogenetics in Oniscidea. Monitore 

Zoologico Italiano, n. s., Monografia 4:3–27. 

Schmidt, M., and S. Harzsch. 1999. Comparative analysis 

of neurogenesis in the central olfactory pathway of 

adult decapod crustaceans by in vivo BrdU labeling. 

Biological Bulletin 196:127–136. 

Schmidt-Rhaesa, A., U. Ehlers, T. Bartolomaeus, C. Lemburg, 

and J. R. Garey. 1998. The phylogenetic position 

of the Arthropoda. Journal of Morphology 

238:263–285. 

Schminke, H. K. 1973. Evolution, System und Verbreitungsgeschichte 

der Familie Parabathynellidae (Bathynellacea, 

Malacostraca). Mikrofauna des Meeresbodens 

24:1–192. 

Schmitt, W. L. 1965. Crustaceans. Ann Arbor, Michigan: 

The University of Michigan Press, 204 pp. 

Scholtz, G. 1995. Expression of the engrailed gene reveals 

nine putative segment-anlagen in the embryonic 

pleon of the freshwater crayfish Cherax destructor 

(Crustacea, Malacostraca, Decapoda). Biological 

Bulletin 188:157–165. 

. 1998. Von Zellen und Kontinenten—die Evolution 

der Flußkrebse (Decapoda, Astacida). Stapfia 

58:205–212. 

. 1999. Freshwater crayfish evolution. Freshwater 

crayfish 12 (Proceedings of the Twelfth Symposium 

of the International Association of Astacology): 36– 

48. 

Scholtz, G., and S. Richter. 1995. Phylogenetic systematics 

of the reptantian Decapoda (Crustacea, Malacostraca). 

Zoological Journal of the Linnean Society 113: 

289–328. 

Schram, F. R. 1971. Polyphyly in the Eumalacostraca? 


. 1981. On the classification of Eumalacostraca. 


. (editor). 1983a. Crustacean issues 1. Crustacean 

phylogeny. Rotterdam: A. A. Balkema Press, 372 pp. 

. 1983b. Remipedia and crustacean phylogeny. In 

Crustacean issues 1. Crustacean phylogeny, ed. F. R. 

Schram, 23–28. Rotterdam: A. A. Balkema Press, 

372 pp. 

. 1984a. Relationships within eumalacostracan 

Crustacea. Transactions of the San Diego Society of 

Natural History 20:301–312. 

. 1984b. Fossil Syncarida. Transactions of the San 

Diego Society of Natural History 20:189–246. 

. 1986. Crustacea. New York: Oxford University 


. 1991. Arthropod pattern theory: a new approach 

to arthropod phylogeny. In Proceedings of the 1990 

International Crustacean Conference, ed. P. J. F. Davie 

and R. H. Quinn. Memoirs of the Queensland 

Museum 31:1–18. 

Schram, F. R., and J. T. Høeg. 1995. New frontiers in 

barnacle evolution. In Crustacean issues 10. New 

frontiers in barnacle evolution, ed. F. R. Schram and 

J. T. Høeg, 297–312. Rotterdam: A. A. Balkema 

Press. 

Schram, F. R., and C. H. J. Hof. 1998. Fossils and the 

interrelationships of major crustacean groups. In Arthropod 

fossils and phylogeny, ed. G. D. Edgecombe, 

233–302. New York: Columbia University Press. 

Schram, F. R., C. H. J. Hof, and F. A. Steeman. 1999. 

Thylacocephala (Arthropoda: Crustacea?) from the 


Cretaceous of Lebanon and implications for thylacocephalan 

systematics. Palaeontology 42:769–797. 

Schram, F. R., and C. Lewis. 1989. Functional morphology 

of feeding in the Nectiopoda. In Crustacean issues 

6. Functional morphology of feeding and 

grooming in Crustacea, ed. B. E. Felgenhauer, 

L. Watling, and A. B. Thistle, 115–122. Rotterdam: 

A. A. Balkema Press. 

Schram, F. R., J. Sieg, and E. Malzahn. 1986. Fossil Tanaidacea. 

Transactions of the San Diego Society of 

Natural History 21:127–144. 

Schram, F. R., S. Yanbin, R. Vonk, and R. S. Taylor. 2000. 

The first fossil stenopodidean. Crustaceana 73:235– 

242. 

Schram, F. R., R. Vonk, and C. H. J. Hof. 1997. Mazon 

Creek Cycloidea. Journal of Paleontology 71:261– 

284. 

Schram, F. R., and J. C. von Vaupel Klein (editors). 1999. 

Crustaceans and the biodiversity crisis. Proceedings 

of the 4th International Crustacean Congress, Amsterdam. 

Leiden: Brill, 1021 pp. 

Schram, F. R., J. Yager, and M. J. Emerson. 1986. Remipedia, 

part 1, systematics. San Diego Society of Natural 

History Memoir 15:1–60. 

Schubart, C. D., J. A. Cuesta, R. Diesel, and D. L. Felder. 

2000a. Molecular phylogeny, taxonomy, and evolution 

of nonmarine lineages within the American 

grapsoid crabs (Crustacea: Brachyura). Molecular 

Phylogenetics and Evolution 15:179–190. 

Schubart, C. D., J. A. Cuesta, and D. L. Felder. In press. 

Glyptograpsidae, a new brachyuran family from 

Central America: larval and adult morphology and 

a molecular phylogeny of the Grapsoidea. Journal of 

Crustacean Biology. 

Schubart, C. D., J. A. Cuesta, and A. Rodríguez. In press. 

Molecular phylogeny of the crab genus Brachynotus 

(Brachyura: Varunidae) based on the 16S rRNA 

gene. Hydrobiologia. 

Schubart, C. D., J. E. Neigel, and D. L. Felder. 1998. The 

use of the mitochondrial 16S gene for the study of 

crustacean phylogenies and biogeography. Proceedings 

and abstracts of the 4th International Crustacean 


. 2000b. Molecular phylogeny of mud crabs 

(Brachyura: Panopeidae) from the northwestern Atlantic 

and the role of morphological stasis and convergence. 


. 2000c. Use of the mitochondrial 16S rRNA gene 

for phylogenetic and population studies of Crustacea. 

In Crustacean issues 12. The biodiversity crisis 

and Crustacea, ed. J. C. von Vaupel Klein and F. R. 

Schram, 817–830. Rotterdam: A. A. Balkema Press. 

Schultz, G. A. 1995. Terrestrial isopod crustaceans (Oniscidea) 

from Paraguay with definition of a new family. 

Revue Suisse de Zoologie 102:387–424. 

Schweitzer, C. E., and R. M. Feldmann. 2000. New species 

of calappid crabs from western North America 

and reconsideration of the Calappidae sensu lato. 

Journal of Paleontology 74:230–246. 

Schweitzer, C. E., and E. W. Salva. 2000. First recognition 

of the Cheiragonidae (Decapoda) in the fossil record 

and comparison of the family with the Atelecyclidae. 


Schwenk, K., A. Sand, M. Boersma, M. Brehem, E. Mader, 

D. Offerhaus, and P. Spaak. 1998. Genetic markers, 

genealogies and biogeographic patterns in the Cladocera. 

Aquatic Ecology 32:37–51. 

Scott, A. 1909. The Copepoda of the Siboga Expedition. 

Part I. Free-swimming, littoral and semi-parasitic 

Copepoda. Siboga Expeditie Monograph 29a:1–323, 

plates 1–69. 

Secretan, S. 1985. Conchyliocarida, a class of fossil crustaceans: 

relationships to Malacostraca and postulated 

behaviour. Transactions of the Royal Society of 

Edinburgh, Earth Sciences 76:381–389. 

. 1998. The sella turcica of crabs and the endophragmal 

system of decapods. Journal of Natural 

History 32:2753–1767. 

Segonzac, M., M. de Saint Laurent, and B. Casanova. 

1993. L’enigma du comportement trophique des 

crevettes Alvinocarididae des sites hydrothermaux de 

la dorsale medio-atlantique. Cahiers de Biologie Marine 

34:535–571. 

Self, J. T. 1969. Biological relationships of the Pentastomida; 

a bibliography on the Pentastomida. Experimental 

Parasitology 24:63–119. 

Serejo, C. S. 2000. A preliminary cladistic analysis of the 

superfamily Talitroidea (Amphipoda, Gammaridea). 

Abstracts of the 10th Colloqium on Amphipoda 

(http://www.odu.edu/%7Ejrh 100f/amphome): 6–7. 

Serène, R. 1984. Crustacés Décapodes Brachyoures de 

l’Ocean Indien Occidental et de la Mer Rouge, Xanthoidea: 

Xanthidae et Trapeziidae. Avec un addendum 

par Crosnier, A.: Carpiliidae et Menippidae. 

Faune Tropicale 24:1–400. 

Seridji, R. 1993. Descriptions of some planktonic larvae 

of the Calappidae (Crustacea: Decapoda: Brachyura). 

Journal of Plankton Research 15:437–453. 

Serov, P. A., and G. D. F. Wilson. 1999. A revision of the 

Pseudojaniridae Wilson, with a description of a new 

genus of Stenetriidae Hansen (Crustacea: Isopoda: 

Asellota). Invertebrate Taxonomy 13:67–116. 

Shank, T. M., M. B. Black, K. M. Halanych, R. A. Lutz, 

and R. C. Vrijenhoek. 1999. Miocene radiation of 

deep-sea hydrothermal vent shrimp (Caridea: Bresiliidae): 

evidence from mitochondrial cytochrome oxidase 

subunit I. Molecular Phylogenetics and Evolution 

13:244–254.Shen, Y. 1984. Occurrence of 

Permian leaid conchostracans in China and its palaeogeographical 

significance. Acta Paleontologica 

Sinica 29:505–513. 

Shen, Y. 1990. A new conchostracan genus from Lower 

Permian Tungziyan Formation, Fujian. Acta Paleontologica 

Sinica 29:309–314. 

Shen, Y., R. S. Taylor, and F. R. Schram. 1998. A new 

spelaeogriphacean (Crustacea: Peracarida) from the 

Upper Jurassic of China. Contributions to Zoology 

(Amsterdam) 68:19–35. 

Shubin, N., C. Tabin, and S. Carroll. 1997. Fossils, genes 

and the evolution of animal limbs. Nature 388:639– 

648. 

Shultz, J. W., and J. C. Regier. 2000. Phylogenetic analysis 

of arthropods using two nuclear protein-encoding 

genes supports a crustacean � hexapod clade. Proceedings 

of the Royal Society of London 267B: 

1011–1019. 

Sieg, J. 1973. Ein Beitrag zum Natürlichen System der 

Dikonophora Lang. (Textteil, Tafelteil). Ph.D. Dissertation, 

University of Kiel, 298 pp. 

. 1976. Zum Natürlichen System der Tanaidacea 

Lang (Crustacea, Tanaidacea). Zeitschrift für Zoologischer 

Systematik und Evolutionsforschung 14: 

177–198. 

. 1982. Uber ein ‘‘connecting link’’ in der Phylogenie 

der Tanaidomorpha (Tanaidacea). Crustaceana 

43:65–77. 


. 1983a. Evolution of Tanaidacea. In Crustacean 

issues 1. Crustacean phylogeny, ed. F. R. Schram, 


. 1983b. Tanaidacea. In Crustaceorum catalogus 

pars 6, ed. H. E. Gruner and L. B. Holthuis, 1–552. 

The Hague: W. Junk. 

. 1984. Neuere Erkenntnisse zum Natürliche System 

der Tanaidacea. Eine phylogenetische Studie. 

Zoologica (Stuttgart) 136:1–132. 

. 1986a. Crustacea Tanaidacea of the Antarctic and 

subanctarctic. 1. On material collected at Tierra del 

Fuego, Isla de los Estados, and the west coast of the 

Antarctic peninsula. Biology of the Antarctic Seas 

18:1–180. 

. 1986b. Tanaidacea (Crustacea) von der Antarktis 

und Subantarktis. II. Tanaidacea gesammelt von Dr. 

J. W. Wägele während der Deutschen Antarktis Expedition 

1983. Mitteilungen aus dem Zoologischen 

Museum der Universität Keil 2:1–80. 

Siveter, D. J., and M. Williams. 1997. Cambrian bradoriid 

and phosphatocopid arthropods of North America. 

Special Papers in Paleontology (The Paleontological 

Association of London) 57:1–69. 

Smirnov, N. N. 1992. The Macrothricidae of the world. 

Guides to the Identification of Microinvertebrates of 

the Continental Waters of the World 1:1–143. 

Soh, H. Y., S. Ohtsuka, H. Imabayashi, and H.-L. Suh. 

1999. A new deep-water calanoid copepod and the 

phylogeny of the genus Nullosetigera nom. nov. in 

the Nullosetigeridae nom. nov. (pro Phyllopus: Phyllopodidae) 

from Japanese waters. Journal of Natural 

History 33:1581–1602. 

Sohn, I. G. 1988. Darwinulocopina (Crustacea, Podocopa), 

a new suborder proposed for nonmarine Palaeozoic 

to Holocene Ostracoda. Proceedings of the 


Souza-Kury, L. A. 1998. Malacostraca—Peracarida. Isopoda. 

Oniscidea. In Catalogue of Crustacea of Brazil, 

ed. P. S. Young, 653–674. Rio de Janeiro: Museu 

Nacional. 

Spears, T., and L. G. Abele. 1988. Molecular phylogeny 

of brachyuran crustaceans based on 18S rRNA nucleotide 

sequences. American Zoologist 28:2A. 

. 1997. Crustacean phylogeny inferred from 18S 

rDNA. In Arthropod relationships, ed. R. A. Fortey 

and R. H. Thomas, 169–187. Systematics Association 

Special Volume series 55. London: Chapman 

and Hall. 

. 1998. The role of 18S rDNA hypervariable regions 

in crustacean phylogeny. Proceedings and Abstracts 

of the 4th International Crustacean Congress, 

Amsterdam: 92 (abstract 171). 

. 1999a. Crustacean molecular systematics and 

phylogeny: the 18S rDNA chronicles. Program and 

Abstracts, The Crustacean Society 1999 Summer 


. 1999b. The phylogenetic relationships of crustaceans 

with foliaceous limbs: an 18S rDNA study of 

Branchiopoda, Cephalocarida, and Phyllocarida. 


. 2000. Branchiopod monophyly and interordinal 

phylogeny inferred from 18S ribosomal DNA. Journal 


Spears, T., L. G. Abele, and M. A. Applegate. 1994. Phylogenetic 

study of cirripedes and selected relatives 

(Thecostraca) based on 18S rDNA sequence analysis. 


Spears, T., L. G. Abele, and W. Kim. 1992. The mono- 

phyly of brachyuran crabs: a phylogenetic study 

based on 18S rRNA. Systematic Biology 41:446– 

461. 

Spears, T., N. Cumberlidge, and L. G. Abele. 2000. A 

molecular-based phylogeny of the freshwater crabs. 


2000 Summer Meeting, Puerto Vallarta, Mexico: 46. 

Spivak, E. D., and J. A. Cuesta. 2000. Larval development 

of Cyrtograpsus affinis (Dana) (Decapoda, Brachyura, 

Varunidae) from Rio de la Plata estuary, reared 

in the laboratory. Scientia Marina 64:29–47. 

Starobogatov, Y. I. 1986. Systematics of Crustacea. Zoologicheskiy 

Zhurnal 65:1769–1781. [In Russian, with 

English summary] 

. 1988. Systematics of Crustacea. Journal of Crustacean 

Biology 8:300–311. (English translation by 

M. J. Grygier of Starobogatov, 1986, above). 

Stebbing, T. R. R. 1899. Revision of Amphipoda (continued). 

Annals and Magazine of Natural History, series 

7, 4:205–211. 

Sternberg, R. von. 1996. Carcinization as an underlying 

synapomorphy for the decapod crustacean taxon 

Meiura. Evolutionary Theory 11:153–162. 

. 1997. Cladistics of the freshwater crab family Trichodactylidae 

(Crustacea: Decapoda): appraising the 

reappraisal. Journal of Comparative Biology 2:49– 

62. 

Sternberg, R. von, and N. Cumberlidge. 1999. A cladistic 

analysis of Platythelphusa A. Milne Edwards, 1887, 

from lake Tanganyika, East Africa (Decapoda: Potamoidea: 

Platythelphusidae) with comments on the 

phylogenetic position of the group. Journal of Natural 

History 33:493–511. 

. 2000a. Cladistic relationships of the true freshwater 

crabs (Decapoda: Eubrachyura: Gecarcinucicoidea, 

Potamoidea, Pseudothelphusoidea, and Trichodactylidae). 

Program and Abstracts, The Crustacean 

Society 2000 Summer Meeting, Puerto Vallarta, 

Mexico: 46. 

. 2000b. Taxic relationships within the Grapsidae 

MacLeay, 1838 (Crustacea: Decapoda: Brachyura). 

Journal of Comparative Biology 3:115–136. 

. 2001. On the heterotreme-thoracotreme distinction 

in the Eubrachyura de Saint-Laurent, 1980 (Decapoda, 

Brachyura). Crustaceana 74:321–338. 

. In press. Notes on the position of the true freshwater 

crabs within the brachyrhynchan Eubrachyura 

(Crustacea: Decapoda: Brachyura). Hydrobiologia. 

Sternberg, R. von, N. Cumberlidge, and G. Rodríguez. 

1999. On the marine sister groups of the freshwater 

crabs (Crustacea: Decapoda: Brachyura). Journal of 

Zoological Systematics and Evolutionary Research 

37:19–38. 

Sˇtevčić, Z. 1973. The systematic position of the family 

Raninidae. Systematic Zoology 22:625–632. 

. 1983. Revision of the Calappidae. Australian Museum 

Memoir 18:165–171. 

. 1988. The status of the family Cheiragonidae Ortmann, 

1893. International Journal of Marine Biology 

and Oceanography (Taranto, Italy) 14:1–14. 

. 1994. Contribution to the re-classification of the 

family Majidae. Periodicum Biologorum 96:419– 

420. 

. 1995. Brachyuran systematics and the position of 

the family Raninidae reconsidered. Arthropoda Selecta 

4:27–36. 

. 1998. Evolutionary arrangement of the brachy- 


uran families together with a checklist. Periodicum 

Biologorum 100:101–104. 

Sˇtevčić, Z., P. Castro, and R. H. Gore. 1988. Re-establishment 

of the family Eumedonidae Dana, 1853 

(Crustacea: Decapoda: Brachyura). Journal of Natural 

History 22:1301–1324. 

Sˇtevčić, Z., and R. H. Gore. 1982. Are the Oxyrhyncha a 

natural group? Thalassia Jugoslavica 17:1–16. 

Stock, J. H. 1968. Vectoriella marinovi, un copépode nouveau, 

parasite d’une annélide polychète pontique. 

Crustaceana 1(supplement 10):186–192. 

Storch, V. 1984. Pentastomida. In Biology of the integument, 

ed. J. Bereiter-Hahn, A. G. Matoltsky, and K. 

S. Richards, 709–713. Berlin: Springer Verlag. 

Storch, V., and B. G. M. Jamieson. 1992. Further spermatological 

evidence for including the Pentastomida 

(tongue worms) in the Crustacea. International Journal 

of Parasitology 22:95–108. 

Strausfeld, N. J. 1998. Crustacean–insect relationships: 

the use of brain characters to derive phylogeny 

among segmented invertebrates. Brain, Behavior, 


Suarez-Morales, E., and T. M. Iliffe. 1996. New superfamily 

of Calanoida (Copepoda) from an anchialine cave 

in the Bahamas. Journal of Crustacean Biology 16: 

754–762. 

Suzuki, H., and C. L. McLay. 1998. Gill-cleaning mechanisms 

of Paratya curvirostris (Caridea, Atyidae) 

and comparisons with seven species of Japanese 

atyid shrimps. Journal of Crustacean Biology 18: 

253–270. 

Svavarsson, J. 1984. Description of the male of Pseudomesus 

brevicornis Hansen, 1916 (Isopoda, Asellota, 

Desmosomatidae) and rejection of the family Pseudomesidae. 

Sarsia 69:37–44. 

. 1987. Reevaluation of Katianira in Arctic waters 

and erection of a new family, Katianiridae (Isopoda: 

Asellota). Journal of Crustacean Biology 7:704–720. 

Swanson, K. M. 1989a. Ostracod phylogeny and evolution—a 

manawan perspective. Courier Forschungsinstitut 

Senckenberg 113:11–20. 

. 1989b. Manawa staceyi n. sp. (Punciidae, Ostracoda): 

soft anatomy and ontongeny. Courier Forschungsinstitut 


. 1990. The punciid ostracod—a new crustacean 

evolutionary window. Courier Forschungsinstitut 


. 1991. Distribution, affinities and origin of the 

Punciidae (Crustacea: Ostracoda). In Proceedings of 

the 1990 International Crustacean Conference, ed. 

P. J. F. Davie and R. H. Quinn. Memoirs of the 

Queensland Museum 31:77–92. 

Tabacaru, I., and D. L. Danielopol. 1996a. Phylogénie des 

Isopodes terrestres. Comptes Rendus Hebdomadaires 

des Seances de l’Academie des Sciences 319:71– 

80. 

. 1996b. Phylogenèse et convergence chez les Isopodes 

terrestres. Vie et Milieu 46:171–181. 

Tabuki, R., and T. Hanai. 1999. A new sigillid ostracod 

from submarine caves of the Ryukyu Islands, Japan. 

Paleontology 42:569–593. 

Takeuchi, I. 1993. Is the Caprellidea a monophyletic 

group? Journal of Natural History 27:947–964. 

Tasch, P. 1969. Branchiopoda. In Treatise on invertebrate 

paleontology, part R, Arthropoda 4, ed. R. C. 

Moore, R128–R191. Lawrence, Kansas: The Geological 


Press. 

Tavares, M. S. 1991. Espèces nouvelles de Cyclodorippoidea 

Ortmann et remarques sur les genres Tymolus 

Stimpson et Cyclodorippe A. Milne Edwards 

(Crustacea, Decapoda, Brachyura). Bulletin de Museum 

National d’Histore Naturelle, series 4, 12:623– 

648. 

. 1993. Crustacea Decapoda: Les Cyclodorippidae 

et Cymonomidae de l’Indo-Ouest-Pacifique à 

l’exclusion du genre Cymonomus. In Résultats des 

Campagnes MUSORSTOM, vol. 10, ed. A. Crosnier. 


vol. 156, 253–313. 

. 1998. Phyllotymolinidae, nouvelle famille de 

Brachyoures Podotremata (Crustacea, Decapoda). 

Zoosystema 20:109–122. 

Taylor, D. J., T. J. Crease, and W. M. Brown. 1999. Phylogenetic 

evidence for a single long-lived clade of 

crustacean cyclic parthenogens and its implications 

for the evolution of sex. Proceedings of the Royal 

Society of London 266B:791–797. 

Taylor, R. S., and F. R. Schram. 1999. Meiura (anomalan 

and brachyuran crabs). In Functional morphology of 

the invertebrate skeleton, ed. E. Savazzi, 517–528. 

Chichester: John Wiley. 

Taylor, R. S., F. R. Schram, and Y. Shen. 1999. A new 

crayfish family (Decapoda: Astacida) from the Upper 

Jurassic of China, with a reinterpretation of other 

Chinese crayfish taxa. Paleontological Research 3: 

121–136. 

Taylor, R. S., Y. Shen, and F. R. Schram. 1998. New pygocephalomorph 

crustaceans from the Permian of 

China and their phylogenetic relationships. Paleontology 

41:815–834. 

Tchindonova, J. G. 1981. New data on the systematic 

position of some deep-sea mysids (Mysidacea, Crustacea) 

and their distribution in the world ocean. In 

Biology of the Pacific Ocean depths, ed. N. G. Vinogradova, 

24–33. Vladivostok: Academy of Sciences 

of the USSR. [In Russian] 

Telford, M. J., and R. H. Thomas. 1995. Demise of the 

Atelocerata? Nature 376:123–124. 

Thatcher, V. E. 1986. The parasitic crustaceans of fishes 

from the Brazilian Amazon, 16, Amazonicopeus 

elongatus gen. et sp. nov. (Copepoda: Poecilostomatoida) 

with the proposal of Amazonicopeidae 

fam. nov. and remarks on its pathogenicity. Amazoniana 

10:49–56. 

Thatcher, V. E., and B. A. Robertson. 1984. The parasitic 

crustaceans of fishes from the Brazilian Amazon. 11. 

Vaigamidae fam. nov. (Copepoda: Poecilostomatoida) 

with males and females of Vaigamus retrobarbatus 

gen. et sp. nov. and V. spinicephalus sp. nov. 

from plankton. Canadian Journal of Zoology 62: 

716–729. 

Thiéry, A. 1996. Branchiopodes I. Ordres des Anostracés, 

Notostracés, Spinicaudata et Laevicaudata (Anostraca 

Sars, 1867—Notostraca Sars, 1867—Spinicaudata 

Linder, 1945—Laevicaudata Linder, 1945). In 

Traité de Zoologie. Anatomie, Systematique, Biologie. 

Tome VII, Fascicule II. Généralités (suite) et Systématique, 

ed. J. Forest, 287–351. Paris: Masson, 

1002 pp. 

Thurston, M. H. 1982. Cheus annae, new genus, new species 

(Cheidae, new family), a fossorial amphipod 

from the Falkland Islands. Journal of Crustacean Biology 

2:410–419. 

. 1989. A new species of Valettia (Crustacea: Amphipoda) 

and the relationship of the Valettidae to the 


Lysianassoidea. Journal of Natural History 23: 

1093–1107. 

Trilles, J.-P. 1999. Ordre des Isopodes sous-ordre des Épicarides 

(Epicaridea Latreille, 1825). In Traité de 

Zoologie. Anatomie, Systématique, Biologie. Tome 

VII, Fascicule IIIA. Crustacés Péracarides, ed. J. Forest. 

Memoires de l’Institut Oceanographique Fondation 

Albert I er , Prince de Monaco, No. 19, 279– 

352. 

Tshudy, D., and L. E. Babcock. 1997. Morphology-based 

phylogenetic analysis of the clawed lobsters (family 

Nephropidae and the new family Chilenophoberidae). 


Tucker, A. B. 1998. Systematics of the Raninidae (Crustacea: 

Decapoda: Brachyura) with accounts of three 

new genera and two new species. Proceedings of the 


Tudge, C. C. 1991. Spermatophore diversity within and 

among the hermit crab families, Coenobitidae, Diogenidae 

and Paguridae (Paguroidea, Anomura, Decapoda). 

Biological Bulletin 181:238–247. 

. 1992. Comparative ultrastructure of hermit crab 

spermatozoa (Paguroidea, Anomura, Decapoda). 


. 1995. Ultrastructure and phylogeny of the spermatozoa 

of the infraorders Thalassinidea and Anomura 

(Decapoda, Crustacea). In Advances in spermatozoal 

phylogeny and taxonomy, ed. B. G. M. Jamieson, 

J. Ausio, and J.-L. Justine. Mémoires du Muséum 

National d’Histoire Naturelle, vol. 166, 251– 

263. 

. 1997a. Spermatological evidence supports the 

taxonomic placement of the Australian endemic 

hairy stone crab, Lomis hirta (Decapoda, Anomura, 

Lomidae). Memoirs of the Museum of Victoria 56: 

235–244. 

. 1997b. Phylogeny of the Anomura (Decapoda, 

Crustacea): spermatozoan and spermatophore morphological 

evidence. Contributions to Zoology 67: 

125–141. 

. 1999a. Ultrastructure of the spermatophore lateral 

ridge in hermit crabs (Decapoda, Anomura, Paguroidea). 


. 1999b. Spermatophore morphology in the hermit 

crab families Paguridae and Parapaguridae (Paguroidea, 

Anomura, Decapoda). Invertebrate Reproduction 


Tudge, C. C., B. G. M. Jamieson, L. Sandberg, and C. 

Erseus. 1998a. Ultrastructure of the mature spermatozoon 

of the king crab Lithodes maja (Lithodidae, 

Anomura, Decapoda): further confirmation of a 

lithodid-pagurid relationship. Invertebrate Biology 

117:57–66. 

Tudge, C. C., B. G. M. Jamieson, M. Segonzac, and D. 

Guinot. 1998b. Spermatozoal ultrastructure in three 

species of hydrothermal vent crab, in the genera Bythograea, 

Austinograea and Segonzacia (Decapoda, 

Brachyura, Bythograeidae). Invertebrate Reproduction 


Tudge, C. C., G. C. B. Poore, and R. Lemaitre. 2000. 

Preliminary phylogenetic analysis of generic relationships 

within the Callianassidae and Ctenochelidae 

(Decapoda: Thalassinidea: Callianassoidea). Journal 

of Crustacean Biology 20(special no. 2):129–149. 

Tudge, C. C., D. M. Scheltinga, and B. G. M. Jamieson. 

1999. Spermatozoal ultrastructure in the Hippoidea 

(Anomura, Decapoda). Journal of Submicroscopic 

Cytology and Pathology 31:1–13. 

Turbeville, J. M., D. M. Pfeiffer, K. G. Field, and R. A. 

Raff. 1991. The phylogenetic status of arthropods, 

as inferred from 18S rRNA sequences. Molecular Biology 


Vader, W. 1972. A list of amphipod genera and species 

described by W. Lilljeborg. Amphipod Newsletter 2: 

13–15. 

Vader, W., A. Baldinger, and T. Krapp-Schickel. 1998. 

IXth International Amphipod Meeting (Kronenberg, 

Germany). Ecdysiast (Newsletter of The Crustacean 

Society,) vol. 17, no. 2:6–7. 

Van Dover, C. L., R. H. Gore, and P. Castro. 1986. Echinoecus 

pentagonus (A. Milne Edwards, 1879): larval 

development and systematic position (Crustacea: 

Brachyura: Xanthoidea nec Parthenopoidea). Journal 


Van Lieshout, S. E. N. 1983. Amsterdam expedition to 

the West Indian islands, report 27. Calabazoidea, 

new suborder of stygobiont Isopoda, discovered in 

Venezuela. Bijdragen tot de Dierkunde 53:165–177. 

Vannier, J., and K. Abe. 1995. Size, body plan, and respiration 

in the Ostracoda. Paleontology 38:843– 

873. 

Vannier, J., M. Williams, and D. J. Siveter. 1997. The 

Cambrian origin of the circulatory system of crustaceans. 

Lethaia 30:169–184. 

Vereshchaka, A. L. 1996. A new genus and species of caridean 

shrimp (Crustacea: Decapoda: Alvinocarididae) 

from North Atlantic hydrothermal vents. Journal 

of the Marine Biological Association of the United 

Kingdom 76:951–961. 

. 1997a. A new family for a deep-sea caridean 

shrimp from North Atlantic hydrothermal vents. 

Journal of the Marine Biological Association of the 

United Kingdom 77:425–438. 

. 1997b. A new family and superfamily for a deepsea 

caridean shrimp from the Galathea collections. 


Vinogradov, M. E., A. F. Volkov, and T. N. Semenova. 

1982 (1996). Hyperiid amphipods (Amphipoda, Hyperiidea) 

of the world oceans. Editor of 1996 English 

version: D. Siegel-Causey, translated under agreement 

with the Smithsonian Institution Libraries, 

Washington, D.C., by Amerind Publishing Co. Pvt. 

Ltd., New Delhi. [English translation of Amfipodygiperiidy 

(Amphipoda, Hyperiidea) mirovogo 

okeana, Leningrad: Nauka Publishers.] 

Vonk, R., and F. R. Schram. 1998. On the distribution 

and phylogeny of ingolfiellid amphipods. Proceedings 

and Abstracts of the 4th International Crustacean 


Wägele, J.-W. 1983. On the origin of the Microcerberidae 

(Crustacea: Isopoda). Zeitschrift für Zoologische 

Systematik und Evolutionsforschung 21:249–262. 

. 1989. Evolution und phylogenetisches System der 

Isopoda. Stand der Forschung und neue Erkenntnisse. 

Zoologica (Stuttgart) 140:1–262. 

. 1994. Review of methodological problems of 

‘computer cladistics’ exemplified with a case study 

on isopod phylogeny (Crustacea: Isopoda). Zeitschift 

für Zoologische Systematik und Evolutionsforschung 

32:81–107. 

. 1996. The theory and methodology of phylogenetic 

systematics is still evolving: a reply to Wilson. 

Vie et Milieu 46:183–184. 

Wägele, J.-W., and A. Brandt. 1988. Protognathia n. gen. 

bathypelagica (Schultz, 1977) rediscovered in the 


Weddell Sea: a missing link between the Gnathiidae 

and the Cirolanidae. Polar Biology 8:359–365. 

Wägele, J.-W., and G. Stanjek. 1995. Arthropod phylogeny 

inferred from partial 12S rRNA revisited: monophyly 

of the Tracheata depends on sequence alignment. 

Journal of Zoological Systematics and Evolutionary 

Research 33:75–80. 

Wagner, H. P. 1994. A monographic review of the Thermosbaenacea. 

Zoologische Verhandelingen 291:1– 

338. 

Walker-Smith, G. K. 2000. Levinebalia maria, a new genus 

and species of Leptostraca (Crustacea) from 

Australia. Memoirs of Museum Victoria 58:137– 

148. 

Walker-Smith, G. K., and G. C. B. Poore. 2001. A phylogeny 

of the Leptostraca (Crustacea) with keys to 

families and genera. Memoirs of Museum Victoria 

58:383–410. 

Walossek, D. 1993. The Upper Cambrian Rehbachiella 

kinnekullensis and the phylogeny of Branchiopoda 

and Crustacea. Fossils and Strata 32:1–202. 

. 1995. The Upper Cambrian Rehbachiella, its larval 

development, morphology, and significance for 

the phylogeny of Branchiopoda and Crustacea. Hydrobiologia 

298:1–13. 

. 1999. On the Cambrian diversity of Crustacea. In 

Crustaceans and the biodiversity crisis. Proceedings 

of the 4th International Crustacean Congress, Amsterdam, 

vol. 1, ed. F. R. Schram and J. C. von Vaupel 

Klein, 3–27. Leiden: Brill. 

Walossek, D., and K. J. Müller. 1990. Stem-lineage crustaceans 

from the Upper Cambrian of Sweden and 

their bearing upon the position of Agnostus. Lethaia 

23:409–427. 

. 1992. The ‘Alum Shale Window’—contribution 

of ‘Orsten’ arthropods to the phylogeny of Crustacea. 


. 1994. Pentastomid parasites from the Lower Paleozoic 

of Sweden. Transactions of the Royal Society 

of Edinburgh, Earth Sciences 85:1–37. 

. 1997. Cambrian ‘‘Orsten’’-type arthropods and 

the phylogeny of Crustacea. In Arthropod relationships, 

ed. R. A. Fortey, 139–153. Systematics Association 

Special Volume Series 55. London: Chapman 

and Hall. 

. 1998. Early arthropod phylogeny in light of the 

Cambrian ‘‘Orsten’’ fossils. In Arthropod fossils and 

phylogeny, ed. G. D. Edgecombe, 185–231. New 

York: Columbia University Press. 

Walossek, D., J. E. Repetski, and K. J. Müller. 1994. An 

exceptionally preserved parasitic arthropod, Heymoniscambria 

taylori n. sp. (Arthropoda incertae 

sedis: Pentastomida), from Cambrian-Ordovician 

boundary beds of Newfoundland, Canada. Canadian 

Journal of Earth Sciences 31:1664–1671. 

Walossek, D., and H. Szaniawski. 1991. Cambrocaris baltica, 

n. gen. n. sp., a possible stem-lineage crustacean 

from the Upper Cambrian of Poland. Lethaia 24: 

363–378. 

Watling, L. 1981. An alternative phylogeny of peracarid 

crustaceans. Journal of Crustacean Biology 1:201– 

210. 

. 1983. Peracaridan disunity and its bearing on Eumalacostracan 

phylogeny with a redefinition of Eumalacostracan 

superorders. In Crustacean issues 1. 

Crustacean phylogeny, ed. F. R. Schram, 213–228. 


. 1998. On peracarid unity: who goes, who stays? 



. 1999a. The place of the Hoplocarida in the malacostracan 

pantheon. Program and Abstracts, The 

Crustacean Society 1999 Summer Meeting, Lafayette, 

Louisiana: 47. 

. 1999b. Toward understanding the relationships of 

the peracaridan orders: the necessity of determining 

exact homologies. In Crustaceans and the biodiversity 

crisis. Proceedings of the 4th International Crustacean 

Congress, Amsterdam, vol. 1, ed. F. R. 

Schram and J. C. von Vaupel Klein, 73–89. Leiden: 

Brill. 

Watling, L., C. H. J. Hof, and F. R. Schram. 2000. The 

place of the Hoplocarida in the malacostracan pantheon. 

Journal of Crustacean Biology 20(special 

number 2):1–11. 

Watling, L., and I. Kornfield. 1996. Cumacean web page, 

URL http://nature.umesci.maine.edu/pub/cumacea. 

html. 

Whatley, R. C., D. J. Siveter, and I. D. Boomer. 1993. 

Arthropoda (Crustacea: Ostracoda). In The fossil record 

2, ed. M. J. Benton, 343–356. London: Chapman 

and Hall. 

Wheeler, W. C. 1997. Sampling, groundplans, total evidence 

and the systematics of arthropods. In Arthropod 

relationships, ed. R. A. Fortey and R. H. Thomas, 

87–96. Systematics Association Special Volume 

Series 55. London: Chapman and Hall. 

. 1998. Molecular systematics and arthropods. In 

Arthropod fossils and phylogeny, ed. G. D. Edgecombe, 

9–32. New York: Columbia University Press. 

Wheeler, W. C., P. Cartwright, and C. Y. Hayashi. 1993. 

Arthropod phylogeny: a combined approach. Cladistics 

9:1–39. 

Willen, E. 1999. Preliminary revision of the Pseudotachidiidae 

Lang, 1936 (Copepoda, Harpacticoida). Courier 

Forschungsinstitut 215:221–225. 

Williams, A. B. 1979. A new crab family from shallow 

waters of the West Indies (Crustacea: Decapoda: 

Brachyura). Proceedings of the Biological Society of 


. 1980. A new crab family from the vicinity of submarine 

thermal vents on the Galapagos Rift (Crustacea: 

Decapoda: Brachyura). Proceedings of the Biological 


. 1988. Lobsters of the world—an illustrated guide. 

Huntington, New York: Osprey Books, 186 pp. 

Williams, T. A., and L. M. Nagy. 1995. Brine shrimp add 

salt to the stew. Current Biology 5:1330–1333. 

Williams, W. D., and J. L. Barnard. 1988. The taxonomy 

of crangonyctoid Amphipoda (Crustacea) from Australian 

fresh waters: foundation studies. Records of 

the Australian Museum, 10(supplement):1–180. 

Williamson, D. I. 1976. Larval characters and the origin 

of crabs (Crustacea, Decapoda, Brachyura). Thalassia 

Jugoslavica 10:401–414. 

. 1982. Larval morphology and diversity. In The 

biology of Crustacea, vol. 2. Embryology, morphology, 

and genetics, ed. L. G. Abele, 43–110. New 


. 1988a. Evolutionary trends in larval form. In Aspects 

of decapod crustacean biology, ed. A. A. Fincham 

and P. Rainbow, 11–25. London: Symposium 

of the Zoological Society of London 59, 375 pp. 

. 1988b. Incongruous larvae and the origin of some 

invertebrate life histories. Progress in Oceanography 

19:87–116. 


Wills, M. A. 1997. A phylogeny of recent and fossil Crustacea 

derived from morphological characters. In Arthropod 

relationships, ed. R. A. Fortey and R. H. 

Thomas, 189–209. Systematics Association Special 

Volume Series 55. London: Chapman and Hall. 

. 1998. Crustacean disparity through the Phanerozoic: 

comparing morphological and stratigraphic 

data. Biological Journal of the Linnean Society 65: 

455–500. 

Wills, M. A, D. E. Briggs, R. A. Fortey, and M. Wilkinson. 

1995. The significance of fossils in understanding arthropod 

evolution. Verhandlungen der Deutschen 

Zoologischen Gesellschaft 88:203–215. 

Wills, M. A., D. E. G. Briggs, R. A. Fortey, M. Wilkinson, 

and P. H. A. Sneath. 1998. An arthropod phylogeny 

based on fossil and Recent taxa. In Arthropod fossils 

and phylogeny, ed. G. D. Edgecombe, 33–105. New 

York: Columbia University Press. 

Wilson, G. D. F. 1986. Pseudojaniridae (Crustacea: Isopoda), 

a new family for Pseudojanira stenetrioides 

Barnard, 1925, a species intermediate between the 

asellote superfamilies Stenetrioidea and Janiroidea. 


99:350–358. 

. 1987. The road to the Janiroidea: comparative 

morphology and evolution of the asellote isopod 

crustaceans. Zeitschrift für Zoologische Systematik 

und Evolutionsforschung 25:257–280. 

. 1989. A systematic revision of the deep-sea subfamily 

Lipomerinae, of the isopod crustacean family 

Munnopsidae. Bulletin of the Scripps Institution of 

Oceanography (University of California, San Diego) 

27:1–138. 

. 1992. Computerized analysis of crustacean relationships. 


. 1994. A phylogenetic analysis of the isopod family 

Janiridae (Crustacea). Invertebrate Taxonomy 8: 

749–766. 

. 1996. Of uropods and isopod crustacean trees: a 

comparison of ‘‘groundpattern’’ and cladistic methods. 

Viet et Milieu 46:139–153. 

Wilson, G. D. F., and R. T. Johnson. 1999. Ancient endemism 

among freshwater isopods (Crustacea, 

Phreatoicidea). In The other 99%. The conservation 

and biodiversity of invertebrates, ed. W. Ponder and 

D. Lunney, 264–268. Mosman, New South Wales: 

Transactions of the Royal Zoological Society of New 

South Wales. 

Wilson, G. D. F., and S. J. Keable. 1999. A new genus of 

phreatoicidean isopod (Crustacea) from the north 

Kimberly region, western Australia. Zoological Journal 

of the Linnean Society 126:51–79. 

. 2001. Systematics of the Phreatoicidea. In Crustacean 

Issues 13. Isopod systematics and evolution, 

ed. B. Kensley and R. C. Brusca, 175–194. Rotterdam: 

A. A. Balkema Press, 357 pp. 

Wilson, K., V. Cahill, E. Ballment, and J. Benzie. 2000. 

The complete sequence of the mitochondrial genome 

of the crustacean Penaeus monodon: are malacostracan 

crustaceans more closely related to insects than 

to branchiopods? Molecular Biology and Evolution 

17:863–874. 

Wingstrand, K. G. 1972. Comparative spermatology of a 

pentastomid Raeillietiella hemidactyli and a branchiuran 

crustacean Argulus foliaceus with a discus- 

sion of pentastomid relationships. Kongelige Danske 

Videnskabarens Selskab Biologiske Skrifter 19:1–72. 

. 1978. Comparative spermatology of the Crustacea 

Entomostraca. 1. Subclass Branchiopoda. Kongelige 

Danske Videnskabarens Selskab Biologiske 

Skrifter 22:1–66. 

. 1988. Comparative spermatology of the Crustacea 

Entomostraca. 2. Subclass Ostracoda. Kongelige 

Danske Videnskabarens Selskab Biologiske Skrifter 

32:1–149. 

Winnepenninckx, B. M. H., T. Backeljau, and R. M. Kristensen. 

1998. Relations of the new phylum Cycliophora. 

Nature 393:636–638. 

Wolff, T. 1989. The genera of Santiidae Kussakin, 1988, 

with the description of a new genus and species 

(Crustacea, Isopoda, Asellota). Steenstrupia 15:177– 

191. 

Wray, G. A., J. S. Levinton, and L. H. Shapiro. 1996. 

Molecular evidence for deep Precambrian divergences 

among metazoan phyla. Science 274:568–573. 

Yager, J. 1981. Remipedia, a new class of crustaceans 

from a marine cave in the Bahamas. Journal of Crustacean 

Biology 1:328–333. 

. 1989a. The male reproductive system, sperm, and 

spermatophores of the primitive hermaphroditic, remipede 

crustacean Speleonectes benjamini. Invertebrate 

Reproduction and Development 15:75–81. 

. 1989b. Pleomothra apletocheles and Godzilliognomus 

frondosus, two new genera and species of remipede 

crustaceans (Godzilliidae) from anchialine 

caves in the Bahamas. Bulletin of Marine Science 44: 

1195–1206. 

. 1991. The Remipedia (Crustacea): recent investigations 

of their biology and phylogeny. Verhandlungen 

der Deutschen Zoologischen Gesellschaft, Stuttgart 

84:261–269. 

Yager, J., and J. H. Carpenter. 1999. Speleonectes epilimnius 

new species (Remipedia, Speleonectidae) from 

surface water of an anchialine cave on San Salvador 

Island, Bahamas. Crustaceana 72:965–977. 

Yager, J., and W. F. Humphreys. 1996. Lasionectes exleyi, 

sp. nov., the first remipede crustacean recorded from 

Australia and the Indian Ocean, with a key to the 

world species. Invertebrate Taxonomy 10:171–187. 

Yager, J., and F. R. Schram. 1986. Lasionectes entrichoma, 

new genus, new species (Crustacea: Remipedia) from 

anchialine caves in the Turks and Caicos, British 

West Indies. Proceedings of the Biological Society of 


Yamaguchi, T., and W. A. Newman. 1990. A new and 

primitive barnacle (Cirripedia: Balanomorpha) from 

the North Fiji Basin abyssal hydrothermal field, and 

its evolutionary implications. Pacific Science 44:135– 

155. 

Yamaguti, S. 1963. Parasitic Copepoda and Branchiura of 

fishes. New York: Interscience Publishers (John Wiley 

& Sons), i–vii � 1104 pp. 

Young, P. S. (editor). 1998. Catalogue of Crustacea of 

Brazil. Rio de Janeiro: Museo Nacional, Série Livros 

6, i–xvii � 717 pp. 

Zhang, W., Y. Shen, and N. Shaowu. 1990. Discovery of 

Jurassic conchostracans with well preserved soft 

parts and notes on its biological significance. Palaeontologia 

Cathayana 5:311–352. 


APPENDIX I. COMMENTS AND OPINIONS 

The following comments and opinions were provided 

by colleagues (all of whom are listed in Appendix 

II) after seeing the penultimate draft of the 

classification. The authors wish to gratefully acknowledge 

them for allowing us to reproduce their 

remarks. References are listed after each comment 

only if those references are not already listed in our 

Literature Cited section. Some authors did not supply 

full references; consequently, references may be 

missing for some papers cited below. 

CRUSTACEA (GENERAL) 

The authors choose to treat the Crustacea as a 

monophyletic group and thus find it justifiable to 

produce an updated classification for organizing 

museum collections and helping students of crustaceans 

to search unfamiliar taxa. It should thus 

become a useful taxonomic tool. I find much merit 

in (1) the exposition of reasons for preferred arrangements 

and (2) the attempt to introduce readers 

to alternative opinions. The permanent drawback 

of this compilation (considered by the authors) 

is that taxa are not justified by diagnostic 

characters. 

As a means of reflecting some current phylogenetic 

ideas on crustaceans, however, the present attempt 

will be considered obsolete almost immediately 

by some workers. The monophyly of the 

Crustacea is far from settled. In fact, in my opinion, 

it is very unlikely. The mandibulate arthropods are 

traditionally divided into two grades (crustaceans 

and tracheates), and it is obvious that the closest 

relatives of the terrestrial tracheates should be 

sought among aquatic crustaceans. If this scenario 

is reasonable, the Crustacea become, in principle, a 

nonmonophyletic grade-group. The Remipedia and 

Malacostraca have been pinpointed as two successive 

outgroups of the Tracheata (Moura and Christoffersen, 

1996). If there is merit in such a proposal, 

an incorrect assumption of monophyly could immediately 

account for many discrepancies noted 

among cladistic papers establishing the position 

and internal relationships of the Crustacea. Researchers 

striving for a phylogenetic arrangement 

of the crustaceans should not exclude the terrestrial 

descendants of crustaceans from their system. For 

these reasons, rather than a practical, largely consensual, 

and authority-based classification of the 

Recent Crustacea, we need to reconstruct the system 

of the Mandibulata (apparently the smallest 

clade that includes all the so-called crustaceans, as 

well as their myriapod and hexapod descendants). 

Furthermore, apomorphic characters need to be 

provided to distinguish acceptable monophyletic 

taxa from unstudied, unknown, or unresolved traditional 

taxa. Let me suggest that this become another 

demanding, but long overdue, story. 

Submitted by Martin L. Christoffersen, 

Federal University of Paraíba, Brazil 

BRANCHIOPODA AS PRIMITIVE 

In regards to your first argument here, there are 

three different sets of authors who cannot confirm 

a branchiopod affinity for this taxon [Rehbachiella] 

and consequently there in fact may be no Cambrian 

branchiopods. The second part of your argument, 

that there are neither Cambrian cephalocarids, nor 

remipedes, is a non-sequitor. The late Ralph Gordon 

Johnson used to say about the apparent age of 

fossils ‘‘Things are always older than you think they 

are.’’ An example of which relates to those Carboniferous 

remipedes; there is in fact something in 

the Silurian of Wisconsin, yet undescribed, that 

may be a remipede. So, your first argument is weak. 

Your second argument, derived from apomorphic 

development, would seem to be valid, at least 

under traditional assumptions. However, two 

points might be mentioned in this regard. The 

weakest point relates to the basic assumption of 

anamorphy � primitive. Certain aspects emerging 

from developmental genetics might suggest an alternative; 

however, this needs to be developed and 

published (something I have not had time to do as 

yet). Nevertheless, if we consider the matter in 

strictly cladistic terms, if as you correctly state that 

anamorphy is unique to branchiopods, within 

Crustacea sensu stricto the issue of plesiomorphy is 

not resolved—branchiopods have it, but non-branchiopods 

(apparently) don’t. If you add outgroups 

from the ‘‘other Mandibulata,’’ in an attempt to polarize 

patterns of development, then if insects are 

in fact a sister group to crustaceans, epimorphy 

could be argued as plesiomorphic. 

Third, the molecular data cited here is not being 

employed properly by you. The distinctness of 

branchiopods here in the papers you cite is stronger 

than you indicate. For example, Spears and Abele 

(1997) under certain assumptions actually pull 

branchiopods into the hexapods, which possibly indicates 

crustacean polyphyly. Of course you say 

that branchiopods are (might be) closer to other 

groups of arthropods—a fair judgment. If true, that 

would indicate that the position of branchiopods 

far exceeds that of a potential ‘‘basal group’’ of 

crustaceans. Primitiveness under those circumstances 

has nothing to do with it. 

In short, you are wise not to create any additional 

taxonomic categories. Moreover, your threepronged 

argument would appear to be not clearly 

drawn at all. 

On the ancestral crustacean . . . you remark that 

Schram and Hof (1998) obtain a clade Phyllopoda. 

First, if you look at the paper carefully, we sometimes 

get a phyllopodan clade, and sometimes 

not—depending on the assumptions and inclusiveness 

of the database employed. Contrary to Schram 

(1986), I think Hof and I would state that the issue 

of whether or not there is a monophyletic clade 

Phyllopoda is indeed an open one—which is not 

102 � Contributions in Science, Number 39 Appendix I: Comments and Opinions

what your sentence says. Second, just because one 

clade in a comprehensive analysis does not find 

wide favor does not necessarily call other aspects 

of the analysis into question. [Editors’note: In our 

penultimate draft, we criticized the recognition of 

the Phyllopoda by Schram and Hof, and then used 

that criticism to cast doubt on other of their findings 

in that paper; this unfair criticism has since 

been removed.] What the main conclusion of 

Schram and Hof indicated was that the issue of 

crustacean phylogenetic relationships has more 

mileage in it before we hope to approach a solution. 

That ought to be conveyed in your text at this 

point. 

Submitted by Frederick R. Schram, 

Zoölogisches Museum, Amsterdam 

BRANCHIOPODA AS PRIMITIVE 

You state several places that you place the Branchiopoda 

as the sister group to the remaining crustaceans. 

This may be correct, but you mention no 

arguments. The only possible arguments could be 

characters shared by the remaining crustaceans that 

would set the Branchiopoda aside. It is not enough 

to state that they [look] very primitive and that 

some of them look like some of the ‘Orsten’ fossils. 

I agree, of course, that the branchiopods ARE indeed 

some of the most primitive Recent Crustacea 

we have, but this doesn’t automatically give them 

sister group position to the rest (only synapomorphies 

for the remaining ...,asmentioned above). 

It is NOT difficult to imagine the branchiopods (or 

the cephalocarids) placed a little bit up in the system. 

It would only require that the primitive features 

that they have are retained a couple of nodes, 

and that those that actually are branched off first 

(Malacostraca, Remipedia, whatever) have attained 

their special modifications independently from other 

Crustacea. 

So, to summarize, the discussion of which Crustacea 

is the most primitive to look at, and which is 

the sister group to the rest, is a mixture of two 

discussions which actually should be separate. The 

two discussions have been treated as one when certain 

other authors have been discussing the same 

for cephalocarids and remipedes, I know, but it 

does not make the discussion more sensible. I believe 

plenty of examples could be mentioned where 

the sister group to a larger group is far from being 

the best candidate as the most primitive one. To 

take an example from animals we are both interested 

in: If for example notostracans are the sister 

group to all the ‘bivalved’ branchiopods, it doesn’t 

follow that they also are the most primitive. This 

is the same story for the possible sister group to the 

Crustacea. We should not exclude any of the derived 

forms from having that honorary position. 

Only synapomorphies uniting the rest can place a 

taxon in this position. 

My advice would be to skip the idea of branchiopod 

as sister group to the rest, unless you pro- 

vide arguments. But of course, you should retain 

the point of branchiopods being quite primitive 

(based on similarities to certain ‘Orsten’ fossils), 

but I think it is impossible and subjective to distinguish 

between the branchiopods and the cephalocarids 

in this respect. [Both] look like certain ‘Orsten’ 

fossils, and not least the cephalocarids. There 

is not [an] objective way to say which is most primitive, 

because it depends on the feature you focus 

on. So, perhaps you should mention both taxa as 

the best candidates to being ‘primitive’. 

Also, I simply don’t understand how you can say 

that we ‘are treating the class Branchiopoda as the 

most primitive of the Crustacea’ when this is not 

included in your classification. It sounds like you 

don’t believe it enough to actually include it (by 

finding a name for the rest). In my opinion, it contains 

no information about primitivity to mention 

it as the first of the classes in your classification. 

Submitted by Jørgen Olesen, 

University of Copenhagen, Denmark 

BRANCHIOPODA 

Elucidation of the relationships of the ‘‘cladoceran’’ 

and ‘‘conchostraca’’ branchiopods appears to have 

reached what is doubtless a temporary impasse. 

Morphology seems to be saying one thing, some 

molecular evidence another. In morphology, the 

‘‘cladoceran’’ orders differ much from each other, 

and attempts to unite them are unsatisfactory. On 

his own estimation, Olesen (1998), who would do 

so, feels that the monophyly of the ‘‘Cladocera’’ 

‘‘may not seem well supported’’ by his cladistic 

analysis. In fact, of five characters used in support, 

three are wrong, one is of no significance, and the 

other is but a small, to be expected, adaptive 

change that could have happened more than once. 

The four constituent groups, which merit ordinal 

rank, differ from each other more than do the various 

orders of the Copepoda. Although some copepods 

are modified for parasitic habits, some representatives 

of all orders retain various fundamental 

similarities. 

Olesen himself says that his analysis does not 

support the ‘‘Conchostraca,’’ nor, incidentally, the 

Spinicaudata, a well-defined component of that 

group, especially if the divergent Cyclestheria is 

segregated from it. Nevertheless, he unites the morphologically 

diverse ‘‘cladoceran’’ orders with the 

unsupported, and very different, ‘‘Conchostraca’’ as 

the ‘‘Diplostraca,’’ which compounds the difficulties. 

All the alleged synapomorphies of the ‘‘Diplostraca’’ 

are incorrect (Fryer, 1999b). Walossek’s 

(1993, 1995) less detailed attempt to demonstrate 

the same relationship fails for similar reasons. 

The Spinicaudata was fully differentiated at least 

as long ago as the early Devonian. Ephippia of even 

extant genera of the ‘‘cladoceran’’ order Anomopoda 

are known from the Lower Cretaceous, and 

molecular evidence suggests that Daphnia originated 

more than 200 My ago (Colbourne and Hebert, 

Contributions in Science, Number 39 Appendix I: Comments and Opinions � 103

1996). The order must be extremely ancient. If the 

‘‘cladoceran’’ orders prove to be monophyletic, they 

must be of extremely ancient origin. The most convincing 

molecular evidence of affinity of the ‘‘cladoceran’’ 

orders is that in all four the V4 and V7 

regions of the small subunit ribosomal RNA possesses 

four helices, three of which are present in 

Cyclestheria but are otherwise so far unique 

(Crease and Taylor, 1998). Cyclestheria, long regarded 

as a somewhat recalcitrant spinicaudatan, 

has often been cast in the role of ancestor of the 

‘‘Cladocera’’—without however demonstrating 

how such different orders as the Anomopoda and 

Haplopoda could have been derived from it. Although 

the helices are very different in length and 

primary sequences of their distal ends in the different 

orders, their locations, secondary structures, 

and primary sequences at their proximal ends are 

conserved, which suggests homology. None of these 

peculiarities is shared with the Spinicaudata, within 

which order Cyclestheria was long included and to 

which it is vastly more similar in morphology than 

it is to any ‘‘cladoceran’’ order! According to some 

investigators, evidence deduced from 18S ribosomal 

DNA supports these relationships (Spears and 

Abele, 2000). However, according to Dumont 

(2000), ‘‘ongoing molecular work using the full sequence 

of the 18S rDNA nuclear gene’’ not only 

confirms the distinction of that order ‘‘but also suggests 

that the Onychopoda might even be more 

closely related to the Anostraca than with the cladoceran 

orders Ctenopoda and Anomopoda.’’ 

Note, also, that the widely accepted 18S rRNA 

phylogenetic tree of the Protozoa has now been seriously 

questioned, and is probably unreliable (Phillippe 

and Adoutte, 1998)! 

With qualifications, some molecular evidence is 

seductive and welcome, but is contradicted by other 

molecular findings, and cannot gainsay either the 

great morphological differences between the groups 

concerned, or the failure to justify either the ‘‘Cladocera,’’ 

‘‘Conchostraca,’’ or ‘‘Diplostraca’’ by cladistic 

analyses. To change the classification of these 

animals on the basis of still-contentious molecular 

evidence while ignoring the larger corpus of information 

now accumulated, not only on morphology 

but on morphology whose functional significance is 

sometimes understood, and on life histories, would 

merely upset what may indeed eventually prove to 

be only an interim scheme, but one which for the 

time being is perfectly serviceable. As Avise (1994) 

notes, morphological and molecular evolution may 

proceed at different rates, and the overall magnitude 

of genetic distance between taxa is not necessarily 

the only, or the best, guide to phylogenetic 

relationships within groups. 

The subclasses Sarsostraca and Phyllopoda seem 

to be unnecessary. The latter name has also already 

been a source of much confusion. A case can be 

made for the Notostraca as being as distinctive as 

the Anostraca, which alone renders grouping into 

subclasses untenable. 

Additional References 

Avise, J. C. 1994. Molecular markers, natural history and 

evolution. New York: Chapman and Hall. 

Colbourne, J. K., and P. D. N. Hebert. 1996. The systematics 

of the North American Daphnia (Crustacea: 

Anomopoda): a molecular phylogenetic approach. 

Philosophical Transactions of the Royal Society of 

London 351B:349–360. 

Dumont, H. J. 2000. Endemism in the Ponto-Caspian fauna, 

with special emphasis on the Onychopoda (Crustacea). 

Advances in Ecological Research 31:181– 

196. 

Phillippe, H., and A. Adoutte. 1998. The molecular phylogeny 

of Eukaryota: solid facts and uncertainties. In 

Evolutionary relationships among Protozoa, eds. G. 

H. Coombs et al., 25–56. London: Chapman and 

Hall. 

Submitted by Geoffrey Fryer, 

University of Lancaster, United Kingdom 

BRANCHIOPODA 

I am not sure that you should not include the Ilyocryptidae 

in your classification. After all, it is a 

quite serious action not to follow the advice of the 

most important Recent taxonomist working in the 

Cladocera that we have (N. N. Smirnov). Especially 

since you follow so many other taxonomists in their 

suggestions. You present no arguments for not doing 

so. One could argue that an eventual splitting 

of the Macrothricidae should await a phylogenetic 

revision, but such a revision is likely not to appear 

in due time. It is true that the change suggested by 

Smirnov may not be based on phylogenetic criteria 

(and the remaining macrothricids may still be paraphyletic), 

but the same could be said about so 

much of your classification anyway, as you mention 

a couple of times. 

I think when it comes to the lower level classification, 

I believe it would be wise to follow the advice 

of the people actually working on the taxa, 

unless you have personal, strong arguments no to 

do so. The case of the ‘Moinidae’ is different because 

Fryer convincingly argues for their unity with 

the rest of the Daphniidae. You could also cite his 

1991 monograph on Daphniidae adaptive radiation 

here. 

The step you take concerning Cyclestheria is OK, 

I think. It is understandable that you choose something 

between the two alternatives. If we one day 

decide to take the full step of the possible sister 

group relation to the Cladocera, then a name is already 

available by Ax (1999). He suggests the term 

‘Cladoceromorpha.’ There are also a couple of new 

molecular papers out on the issue that seem to support 

Cyclestheria in the mentioned sister group position. 



BRANCHIOPODA 

The quotation from Fryer really encapsulates what 

is wrong with the old ideas about crustacean phy- 


logeny and taxonomy. This focus on ‘‘. . . animals 

that work . . .’’ is directly lifted from the later writings 

of Sidnie Manton. Schram (1993, The British 

School: Calman, Canon, and Manton and their effect 

on carcinology in the English speaking world; 

Crustacean Issues 8:321–348) outlined the roots of 

Mantonian reasoning in an idealist philosophical 

tradition that passed on through Thompson and his 

treatise On Growth and Form. This is essentially a 

Platonic view of comparative biology, and stands 

essentially at odds with the current emphasis, either 

a priori or a posteriori, on elucidating ground 

plans. You are of course free to quote Fryer, but 

you ought to give fair play to alternative philosophical 

and conceptual foundations for systematics. 



BRANCHIOPODA: ANOSTRACA 

Weekers et al. (in press) examined small subunit 

ribosomal DNA of anostracans from 23 genera belonging 

to eight of the nine families recognized by 

Brtek (1997). Their results do not support the family 

Linderiellidae or Polyartemiidae. Instead, they 

group Linderiella with Polyartemia and Polyartemiella 

as a subfamily of the family Chirocephalidae. 

Morphological considerations support this arrangement 

in that the three genera share rigid antennal 

appendages on otherwise simple antennae and double 

pre-epipodites. Unfortunately, these workers 

were not able to obtain usable Artemiopsis. Thus, 

the validity of Artemiopsidae remains untested by 

molecular methods; however, I continue to consider 

that the morphology of the penes places Artemiopsis 

in the family Chirocephalidae. 


Weekers, P. H. H., G. Murugan, J. R. Vanfleteren, and H. 

J. Dumont. In press. Phylogenetic analysis of anostracans 

(Branchiopoda: Anostraca) inferred from 

SSU rDNA sequences. Molecular Phylogenetics and 

Evolution. 

Submitted by Denton Belk, 

Our Lady of the Lake University, 

San Antonio, Texas 

REMIPEDIA 

See comments from G. Boxshall under Maxillopoda 

and from M. Christoffersen under Crustacea. 

REMIPEDIA 

In the section about the Remipedia, you mention 

that the similarities between the Maxillopoda and 

the Remipedia are symplesiomorphies. But what 

are these? The only similarities I can think of, I 

would not consider as symplesiomorphies, but perhaps 

as convergences. Perhaps it is unwise to mention 

something like this without also mentioning 

the characters. The first question people will raise 

is what these characters are. In the same section 

you use the term ‘basal’ about branchiopods, but 

what does that actually mean? There are two possibilities, 

either early off split (e.g., sister group) or 

primitive (or at least with many primitive features), 

but these are two different things, as addressed earlier. 



CEPHALOCARIDA 

In the section about the Cephalocarida, you say 

that the sequence of the classes reflects something 

(it doesn’t matter exactly what in this context). My 

problem here is that I don’t think that the sequence 

of taxa of equal rank in a classification reflects anything. 

If a classification shall reflect anything concerning 

relationship, it has to be put into the hierachical 

categories (like you have done for the classification 

within the Branchiopoda, for example). I 

think this is an old way of thinking with no meaning 

today. 



MAXILLOPODA 

The status of the Maxillopoda remains uncertain. I 

consider that there is a group of related taxa which 

form the core of a Maxillopoda: these are the Copepoda, 

Thecostraca, Tantulocarida and Ostracoda 

(excluding the Phosphatocopines which are not ostracods 

and do not even belong to the crown group 

of the Crustacea). The Mystacocarida and Branchiura 

may also belong to this group but the available 

supporting evidence is weaker. I also consider 

that the Remipedia is related to the maxillopodan 

lineage. Remipedes share several derived features of 

the thoracopods, maxillules and maxillae with other 

maxillopodans as indicated in my paper on comparative 

musculature (Boxshall, 1997). 


Boxshall, G. A. 1997. Comparative limb morphology in 

major arthropod groups: the coxa-basis joint in postmandibular 

limbs. In Arthropod relationships, eds. 

R. A. Fortey and R. H. Thomas, 155–167. London: 

Chapman and Hall. 

Submitted by Geoff Boxshall, 

Natural History Museum, London 

MAXILLOPODA 

I really understand your difficulties here. To cut the 

message short, I think you should have chosen to 

include the component taxa of the Maxillopoda as 

classes and then skip the ‘Maxillopoda’ (as you also 

almost decided to, I can see from your writing). 

I know you [are trying] to be conservative by 

following Bowman and Abele here, but actually, to 

be real conservative you should skip that level. This 


would be a choice of the future for the reasons 

mentioned below. 

I think it is better to have your higher level classification 

to include only what is quite certain. The 

highest categories (classes) should then be something 

like the following: Malacostraca, Branchiopoda, 

Remipedia, Copepoda, Mystacocarida, Branchiura, 

Thecostraca, Cephalocarida, Ostracoda, 

Tantulocarida, (Pentastomida). 

These are with the highest certainty all monophyletic 

(not considering that insects may go in 

somewhere). As for the grouping of these taxa, we 

appear to know too little yet. As you know, this is 

reflected in the high number of different schemes 

put forward that all differ from each other. Perhaps 

it will take 50–100 years before we get the full story, 

if ever. The great advantage of having such a flat 

structure is that it would tell people what the crustacean 

community thinks is certain, but it would 

also point at what is unknown by not having any 

of these weakly supported higher level taxa included 

(like Maxillopoda, Entomostraca, Thoracopoda, 

and the one you now suggest being comprised of 

all non-branchiopod Crustacea). This will be a logical 

starting point for any students of the Crustacea 

that want to address the higher level phylogeny. If 

a taxon like Maxillopoda is included, for example, 

then the starting point is most likely already polluted. 



MAXILLOPODA: RHIZOCEPHALA 

Boschma (1928) is without any doubt the author 

of the family Lernaeodiscidae, but both the families 

Peltogastridae and Sacculinidae must be ascribed to 

Lilljeborg (1860). This has been duly checked. 

Boschma lived 1893–1976, and cannot possibly be 

the author of these two families. Holthuis and I 

consulted Lilljeborg’s (1860) publication, a copy of 

which is in our library; there is not a shadow of a 

doubt concerning his authorship! 

Submitted by W. Vervoort, 

Rijksmuseum van Natuurlijke Historie, 

Leiden, The Netherlands 

MAXILLOPODA: COPEPODA 

I suggest you strictly adhere to what is already published. 

Names should in my view not be introduced 

unofficially but through full and reviewed papers. 

Two PhD theses have just been completed here with 

phylogenetic revisions of the Cyclopoida and one 

branch of Harpacticoida. I could tell you all the 

changes they entail but that would alter your list 

quite visibly. The Poecilostomatoida, e.g., are not a 

separate order but a specialised branch within Cyclopoida. 

There are many new families and others 

had to be synonymized. So, please, stick to published 

and avoid cryptic information (� pers. 

comm.). 

Submitted by H. Kurt Schminke, 

Universität Oldenburg, Germany 

MAXILLOPODA: PENTASTOMIDA 

First, on a separate subclass Pentastomida—what 

can I say. You cite all the relevant papers that argue 

and provide evidence that these are Branchiura, and 

yet you reject these and separate them. This is one 

of the few places where we have good apomorphies 

to unite the groups involved. If you accept Thecostraca, 

then why not accept a single subclass Branchiura 

with two orders: Arguloida and Cephalobaenida? 

Concerning the Walossek arguments in the second 

paragraph: All this Cambrian apparent pentastomid 

says is that Pentastomida are older than we 

thought they were. It does not argue against anything. 

You rightly point out that the fossils might 

not even be pentastomids. As to whether or not the 

hosts ‘‘were on the scene,’’ you must be careful. Recent 

issues of Science and Nature have featured a 

stunningly preserved early chordate that to all intents 

and purposes looks like it was drawn by old 

Al Romer himself when figuring a vertebrate ancestor. 

This Chengjiang fossil in fact trumps Brusca’s 

suggestion, which is true by the way, that the 

conodont animal is a chordate. 



OSTRACODA 

I am sure the classification and appended rationale 

will be useful and will advance the study of crustaceans. 

I am still of the opinion that the suborders 

of the order Podocopida are unnecessary and 

should be deleted, especially as each contains only 

one superfamily except for the Cypridoidea, all superfamilies 

of which are monotypic. 

It is likely that the paleontologists will follow the 

classification that is published in the revised Treatise, 

and that classification will be determined by 

Professor Whatley and his team of specialists, 

which includes Dr. Martens. 

As for -acea v. -oidea, you must of course be consistent 

throughout your classification. Some volumes 

of the Treatise (most notably the revision of 

the brachiopods) have now begun to follow the recommendation 

of the ICZN, but you should realize 

that these are only recommendations, not rules; and 

they may sometimes lead to the curious duplications 

of names among superfamilies and genera. 

Good luck with the classification. I look forward 

to seeing the final version. 


Submitted by Roger L. Kaesler, 

Paleontological Institute, 

The University of Kansas 

OSTRACODA 

Spelling of Suborder Halocyprina Dana, 1853. 

Dana (1853: 1281) based his subfamily Halocyprinae 

and family Halocypridae on his new genus Halocypris. 

Therefore, at least according to present 

rules, the subfamily should be Halocypridinae and 

the family Halocyprididae. Dana did not use the 

names Halocyprina or Halocyprida. If you are basing 

your Halocyprina and Halocyprida on the family 

name Halocyprididae, it seems to me that, to be 

consistent, the suborder should be Halocypridina 

and the order should be Halocypridida. If you are 

basing your Halocyprina on the commonly used 

name for the order, Halocyprida, then I think you 

are correct in using Halocyprina. Possibly, you 

should explain your reasoning for using Halocyprina, 

because I think that you are creating a new 

spelling for the suborder. [Editors’note: we retained 

the spelling Halocyprida for the order, as listed in 

Bowman and Abele (1982: 13), and Halocyprina 

for the suborder based on the order name.] 

Submitted by Louis Kornicker, 

Smithsonian Institution, 

National Museum of Natural History 

STOMATOPODA 

I am leery of following suggestions made in abstracts 

concerning higher taxonomy. Cappola has 

never published her Pseudosquilloidea (which I see 

you accept) with documented reasons for her decision. 

In fact, some of the new analyses of Ahyong 

and Hof (not yet published) would not entirely support 

such an arrangement. 

Thus, while we are at it, you need to turn to 

[page 86 in original draft]. I suggest for now you 

simply leave all the ‘‘gonodactyloid’’ families in one 

superfamily Gonodactyloidea. When we can identify 

clear clades and suggest valid groupings, you 

can change it; or when people actually publish revisions 

in a refereed journal. 



AMPHIPODA 

Although I agree in general with the thrust of your 

arguments, you fail to recognise the complexity of 

amphipod morphology and the lack of family level 

revisions, which makes the development of an acceptable 

classification extremely difficult. Suborder 

and families were established long ago and for the 

most part have never been revised. Superfamilies 

were to a certain extent based on gestalt, which 

worked well for some groups like corophioids, lysianssoids 

and haustorioids, but failed for families 

which didn’t show clear body-plan relationships. 

Even groups as seemingly distinctive as the Lysianassoidea 

are very difficult to define morphologically 

when all genera are considered. When Barnard 

and Karaman (1991) collapsed the majority 

of corophioid families, they did it because these traditional 

families (although workable when they 

were originally established) were no longer definable 

and could no longer be supported. Genera described 

over the years had been pigeon-holed into 

one family or another until any characters which 

might define them had become totally diluted. It 

will take a large effort using modern phylogenetic 

techniques to develop an acceptable classification. 

The results of these works have to be published in 

reputable journals after careful peer-group review. 

Attempts to revise classifications are underway. For 

instance, Lowry and Myers are currently revising 

the iphimedioid group and Myers and Lowry are 

revising the corophioid group. The website 

www.crustacea.net has recently been established to 

publish information and retrieval systems (electronic 

monographs) for all crustaceans. For instance, 

Watling and his students are currently preparing 

cumacean data bases and Lowry and his students 

are working on amphipod data bases for the website. 

It is unfortunate that the use of poorly refereed 

journals and pseudophylogenetic methodologies 

have been used in some cases to produce untestable 

and, in some cases, unacceptable classification systems. 

Because of these problems, we currently list our 

taxa alphabetically in the Amphipoda. I do not see 

the problem. All classifications are hypotheses 

which change as new hypotheses are produced. In 

a large monograph, it is fine to discuss and list the 

phylogenetic classification, but probably the taxonomic 

section should be alphabetical. Trying to find 

families or genera listed phylogenetically in a large 

monograph can be a nightmare for those not in the 

know (basically everyone but experts). It is relatively 

easy, for example, to find a family level taxon 

in Barnard and Karaman (1991). One does not 

have to continually consult the index. 

Submitted by Jim Lowry, 

Australian Museum, Sydney 

AMPHIPODA: GAMMARIDEA 

As Ed Bousfield was not present at the amphipod 

conference in Amsterdam to defend the value of 

phyletic vs. alphabetical classification of the Gammaridea, 

several points raised in the Vader-Baldinger-K-S-Watling 

report seem largely matters of mechanics 

rather than matters of phyletic substance. 

Some points of your recent ‘‘critique’’ summary 

may require modification, viz: (1) ‘‘the schedules of 

Jerry Barnard and Ed Bousfield (are) often not very 

compatible’’ and (2) ‘‘. . . not espousing one worker’s 

view over another.’’ With all due respect to Jerry’s 

enormous contribution to gammaridean taxonomy, 

his formal ‘‘track record’’ in gammaridean 

phylogeny was actually quite modest in scope. 


Thus, he did recognize (temporarily, at various 

times) Talitroidea Bulycheva, 1957, Corophioidea 

Barnard, 1973, and Haustorioidea Barnard and 

Drummond, 1982. Several of Jerry’s informal ‘‘anglicized’’ 

groupings of freshwater families (e.g., 

‘‘gammarida,’’ ‘‘crangonyctoids,’’ ‘‘hadzioid 

group,’’ etc., in Barnard and Barnard, 1983; Williams 

and Barnard, 1988) rather closely resemble 

some of the superfamilies (and families) formally 

named and fully defined previously (1973, 1977, 

1979, 1982) by Bousfield and co-workers (e.g., 

about 75% compatibility with Gammaroidea, Hadzioidea, 

Crangonyctoidea, Melphidippoidea, etc.). 

However, he did not attempt formal phyletic groupings 

of most marine gammaridean families, nor formal 

integration with other amphipod suborders. 

Unlike Sars (1895), Stebbing (1906), and other 

‘‘turn-of-the-century’’ workers, Jerry apparently did 

not recognize the significance of reproductive form 

and behaviour in amphipod phylogeny. Jerry’s final 

major work (with Gordan Karaman, 1991, p. 7) 

disavowed the significance or use of the formal superfamily 

concept, and listed families alphabetically 

rather than phyletically or semi-phyletically (as in 

Sars and Stebbing, above). Some classifications are 

based on carefully defined characters and character 

states that have required (and will continue to require) 

modification according to features found in 

subsequently discovered species and genera, and 

are consistent at proper classificatory levels. The 

cladistic arrangement by Kim and Kim (1993), reviewed 

rather unfavourably by Schram (1994), underscores 

the unreliability of cladistic analysis when 

care is not taken in the appropriate selection and 

accurate definition of characters and character 

states. 

[Concerning your statement about Bousfield and 

Shih], Bousfield and Shih (1994) represents an updating 

and refinement of previous �20 years of 

study and publication on gammaridean phylogeny. 

[Concerning your statement about Reptantia], 

‘‘Natantia’’ and ‘‘Reptantia’’ are terms (names) 

pragmatically defined, but not incorporated formally 

by Bousfield and Shih (1994). The terms are 

analogous to former groupings of families and superfamilies, 

etc., within the Order Decapoda. 

[Concerning your statement about names and 

dates], omission of author names and dates in tabular 

listing of families and superfamilies is modeled 

after similar ‘‘heading’’ omissions in Barnard’s 

‘‘Families and Genera . . .‘‘ (1969) and earlier ‘‘Index 

. . .’’ (1958). Obviously, these names are fully 

treated in the major references (e.g., Stebbing, 

1906; Gurjanova, 1951; Bousfield 1979, 1982, 

l983; Schram, 1986). Readers are expected to provide 

something of substance to the discussion, such 

as commentary on the paper’s extensive analysis of 

‘‘across-the-phyletic-board’’ variability of major 

characters and character states (antennae to telson) 

that would be of prime significance in a cladistic 

treatment. 

[Concerning your statement to the effect that we 

presented different phylogenetic hypotheses in our 

1994 paper], Bousfield and Shih acknowledge 

(problems in resolution) that they do not have a 

‘‘final answer’’ to the probably correct evolutionary 

history of the Amphipoda (only one answer can be 

correct!). Their ‘‘semi-phyletic’’ methodology modifies 

the strictly phenetic format of Sneath and Sokal 

(1973) by careful ordering of character states 

to arrive at a ‘‘plesio-apo-morphic index’’ of probably 

correct phyletic relativity for each taxon. This 

approach tends to minimize the negative effects of 

homoplasious convergence in many of these character 

states (analyzed above). Mike Ghiselin (1984) 

correctly points out, rigid and uncritical application 

of cladistic methodology alone quite frequently 

leads the user to a less-than-credible phylogenetic 

result. Thus, use of the ‘‘Wagner 78’’ cladistic program 

often provides multiple ‘‘trees’’ from the same 

data base, each one different, each one tending to 

invalidate the other, and none probably correct! 

[Concerning your statement about cladistic analyses 

having high priority], to my knowledge, cladistic 

‘‘purists’’ have not yet actually demonstrated 

a cladistically derived treatment of all 118 gammaridean 

families of your list. Chances of doing so 

would appear ‘‘slim-to-non-existent.’’ Instead, advocacy 

of rDNA methodology would probably result 

much sooner in a most-probably-correct answer! 

[Concerning your statement that most workers 

would prefer to see the families listed alphabetically 

rather than by superfamily], how surprising that 

such an unsupported statement should come from 

Les Watling, a confirmed crustacean phylogenist! 

On more serious reflection, Les may find that quite 

a few current workers (e.g., Mike Thurston, John 

Holsinger) do not ‘‘give up’’ so easily on the full 

solution of this difficult problem. 

[Concerning your use of the word hypotheses], 

do you mean ‘‘concepts’’? All family and superfamily 

names represent ‘‘concepts’’ of presumed natural 

groupings of species. Some are better defined (in 

terms of careful definition of character states) and 

longer time-tested than others. Most superfamily 

names in Bousfield and Shih (1994) have been carefully 

and fully (multiple-character) defined, their 

component families named, and time-tested (by 

other workers as well) over a 15� year period. 

Since superfamily taxonomic stability (75%) would 

appear at least equal to that of the component family-level 

names of the current Martin–Davis list, although 

both lists are ‘‘conceptual,’’ neither can realistically 

be termed ‘‘hypothetical.’’ 


[Note: Dr. Bousfield did not supply references to all papers 

mentioned above.] 

Bousfield, E. L. 1995. A contribution to the natural classification 

of Lower and Middle Cambrian arthropods: 

food gathering and feeding mechanisms. Amphipacifica 

II(1):3–34. 


Bousfield, E. L. 1996. A contribution to the reclassification 

of neotropical freshwater hyalellid amphipods 

(Crustacea: Gammaridea: Talitroidea). Bull. Mus. 

civ. St. nat. Verona 20[1993 (1996)]:175–224. 

Submitted by Ed Bousfield, 

Ottawa, Canada 

AMPHIPODA: GAMMARIDEA 

There would seem to be a second main reason why 

you might regret not employing a natural (superfamily) 

classification of the Gammaridea. Not only 

the Lysianassoidea, Talitroidea and Corophioidea, 

but about 75% of superfamilies of the Bousfield– 

Schram phyletic classification (including Jerry Barnard’s 

anglicized versions) are variously utilized by 

major workers today—if only because they make 

pragmatic (workable) sense. 

Interestingly, and to my knowledge, none of 

those who apparently condemn the present superfamily 

categories because they ‘‘have not been derived 

cladistically’’ has attempted a natural treatment 

of all 113 families (embracing �5000� species!) 

of your list, based on cladistics alone. 

Why?—not only is the task extremely difficult and 

time-consuming, but the feasibility of obtaining a 

single, credible, ‘‘all-inclusive’’ answer with that 

methodology alone is highly improbable, and I 

think they know it! On the other hand, rDNA studies 

seem virtually unaffected by homoplasious convergence 

of morphological character states ‘‘across 

the board’’ and are quite promising—if only someone 

would get started! 

The second, and perhaps more important, essentially 

scientific reason is that gammarideans, virtually 

alone among crustacean higher taxa (including 

the 3 other amphipod suborders!) would remain 

unclassified phyletically. Such an anomalous situation 

will be corrected inevitably—hopefully sooner 

than later—providing the principal reason for phyletic 

classification in the forthcoming CNAI lists 

and Pacific amphipod guide. Sars, Stebbing, and 

other perceptive ‘‘turn-of-the-century’’ amphipodologists 

might then cease ‘‘rolling over in their 

graves’’! 

Submitted by Ed Bousfield, 

Ottawa, Canada 

ISOPODA 

In the near future, we must abandon the use of 

Linnean categories, because we are currently identifying 

many more encaptic levels of monophyletic 

groups than there are hierarchical levels in the Linnean 

system. The Paranthuridae, for example, are 

definitely a monophyletic group that contains further 

subgroups. To erect new families for these subgroups 

means to give up a categorical rank for the 

taxon Paranthuridae. 

The same problem exists for the Epicaridea. New 

molecular evidence (Ph.D. thesis of H. Dreyer) 

proves that these parasites of crustaceans are de- 

rived from a common ancestor shared with the Cymothoidae 

(fish parasites). Thus, the suborder Epicaridea 

is placed within the suborder ‘‘Flabellifera’’ 

or, more precisely, within the suborder Cymothoidea 

sensu Wägele (1989), the sister group of the 

suborder being a taxon classified as a family. 

Concerning the hypothesis that the Sphaeromatidae, 

Serolidae, and other groups are derived from 

a disc-shaped ancestor (the ancestor of the Sphaeromatidea 

sensu Wägele, 1989), new evidence was 

discovered with the fossil Schweglerella stroebli 

(Polz, H. 1998. Archaeopteryx 16:19–28). This animal 

shows neither the apomorphies of the Serolidae 

nor of the Sphaeromatidae or other related extant 

taxa, but shows those characters identified as 

apomorphies of the suborder Sphaeromatidea (e.g., 

disc-shaped body, head immersed in first pereonite, 

dorsal eyes). 

The subdivision of the Oniscidea into Tylomorpha 

and Ligiamorpha does not reflect the phylogeny 

of terrestrial isopods, as shown by Erhard 

(1996, 1998). Detailed phylogenetic analyses based 

on morphological characters will be published soon 

(Ph.D. theses of C. Schmidt and of A. Leistikow). 

Submitted by J. W. Wägele, 

Ruhr-Universität Bochum, Germany 

SYNCARIDA 

The author of both the Bathynellidae and Bathynellacea 

is Chappuis, 1915. I have copied the paper 

by Chappuis (1915) for you. I am a bit surprised 

that you cite Lopretto and Morrone (1998) who 

have added nothing new to our understanding of 

Syncarida. You should quote those who have. 

Submitted by H. Kurt Schminke, 

Universität Oldenburg, Germany 

DECAPODA: CARIDEA 

I am puzzled to find the family Barbouridae among 

the superfamily Bresilioidea. Chace (1997) put 

them among the hippolytids. Christoffersen (1987, 

1990) put them in the superfamily Crangonoidea. 

Who put them among the bresilioideans, and why? 

This is not stated clearly in your section on the superfamily 

Bresilioidea on p. 61. [Editor’s note: the 

family Barbouriidae Christoffersen was mistakenly 

placed by us in the Bresilioidea; this has since been 

corrected and they are now listed among the Alpheoidea.] 

Otherwise, the classification contains the usual 

fights between lumpers and splitters. I think that 

Christoffersen’s classification may fall apart in the 

future because much of it is based on descriptions 

from the literature and not on examination of actual 

specimens. Some of the descriptions are inaccurate 

or do not contain pertinent information 

needed in classification today. 

Submitted by Mary K. Wicksten, 

Texas A&M University 


DECAPODA: CARIDEA 

I of course must strongly disagree with the proposed 

arrangement of the caridean families into superfamilies, 

because I see this as a retrocess from 

taxa sustained by apomorphic characters (Christoffersen, 

1990) back to groupings based on overall 

resemblance, authority (Chace, 1992; Holthuis, 

1993), or arbitrary usage. It is true that my proposals 

have had little following in the carcinological 

community, and that some of my employed 

characters may be questionable. But it is also true 

that my efforts remain the first attempt to produce 

a phylogenetic system of the Caridea. Because my 

system differs substantially from the traditional arrangements, 

my suggestions have usually been dismissed 

as totally heretical, without any serious attempt 

to argue alternative possibilities sustained by 

better uniquely shared characters. It is rather depressing 

to note that the present authors follow this 

same tactic. They do not accept a single superfamily 

as synthesized in Christoffersen (1990). More 

explicitly, but without justification, they reject my 

proposal to combine alpheoids, crangonoids and 

pandaloids into a monophyletic taxon. This is surprising 

to me, because these superfamilies, as redefined 

in my cited works, share a remarkable synapomorphy, 

the multiarticulated carpus of the second 

pereiopod, which is a unique adaptation within 

the carideans for body cleaning. For this transformation 

series, there is even a transitional stage represented 

by the nematocarcinoids, in which the carpus 

of the second pereiopods is longer than in the 

preceding carideans, before being subdivided in the 

sister group represented by pandaloids, crangonoids, 

and alpheoids. At a still higher level of generality, 

this transformation series is congruent with 

the presence of a well developed incisor process on 

the mandible of palaemonoids and all the previously 

mentioned superfamilies. Going to a lower 

hierarchical level, there is further congruence with 

the uniquely expanded first cheliped in crangonoids 

and alpheoids. My rearrangements of the traditional 

families into superfamilies eliminate all the paraphyletic 

family-level taxa, including the notably 

unsatisfactory Hippolytidae. Finally, just to mention 

one remarkable autapomorphy justifying one 

of my new superfamilies, only palaemonids and 

rhynchocinetids share a second distolateral tooth 

on the basal segment of the antennule, in addition 

to the usual stylocerite. Some researchers complain 

that I presented few characters for each node, but 

this is because my approach is qualitative and I selected 

the best possible evidence from detailed studies 

of the total morphological and species diversity 

of the Caridea. To refute the phylogenetic system, 

it is necessary that researchers argue for alternative 

replacement characters where they believe I have 

failed. Simply ignoring the system does not justify 

the usual assumption that my arrangements are totally 

wrong! 

Submitted by Martin L. Christoffersen, 

Federal University of Paraíba, Brazil 

DECAPODA: REPTANTIA 

I really do not understand why you do not use a 

separate category for Reptantia. It is one of the 

clearest, most universally accepted groups (taxonomically 

or cladistically) among the decapods that 

we have. 



DECAPODA: ASTACIDEA 

You really do get yourselves into deep water when 

you try to offer editorial comments on cladistic 

analyses. Here you hit another one. Where do you 

get the idea of ‘‘extremely primitive Neoglyphea’’ 

from? Forest and de St. Laurent (1989, Nouvelle 

contribution a la connaissance de Neoglyphea inopinata 

a propos de la description de la femelle adulte, 

Res. Camp. Musorstom 5, Memoirs Mus. Nat. 

His. Nat., series A, 144:75–92) made [a] good argument 

for allying glypheoids with astacids—not a 

particularly primitive alliance. My own preliminary 

examination of decapod phylogeny (submitted, Hydrobiologia) 

not only fairly well confirms the 

Scholtz and Richter scheme, but also squarely places 

Neoglyphea within the Fractosternalia. 

As I say, my own examination of the subject in 

connection with an assignment to address decapod 

phylogeny in connection with the beginning revision 

of the decapod section of the Treatise on Invertebrate 

Paleontology has, to my surprise, uncovered 

the basic robustness of the Scholtz and Richter 

analysis. I think you would do well to leave yourself 

an opening here. 

Of course, I see why you are keen to downplay 

Scholtz and Richter because here you adapt a very 

conservative combination of ‘‘clawed lobsters.’’ I 

can accept this for now. However, I think it would 

only be fair for you to point out that Scholtz and 

Richter would segregate the ‘‘clawed (true) lobsters’’ 

as Homarida from the crayfish as Astacida. 

My own on-going, recent work indicates that at 

least the genus Neoglyphea is a fractosternalian in 

some kind of proximity to the Astacida, and that 

Enoplometopus may even be a separate clade from 

the Nephropoidea. That this paraphyly should 

emerge among ‘‘lobsters’’ is not too surprising, 

since we discover again and again that supposedly 

robust, traditional groups bearing a lot [of] plesiomorphies 

emerge on closer examination as paraphyletic 

taxa. Why should macrurous lobsters be 

any different? 



DECAPODA: ANOMURA 

I fully agree with the different parts of my specialty 

(Anomura). I agree with the changes included in 

this new version. As you mention . . . we need more 

studies (especially molecular) to improve our 


knowledge on the phylogeny and the classification 

of Crustacea, and obviously new, and perhaps 

strong, changes will come in the near future. However, 

we need to put in order our present knowledge 

of the group. 

Submitted by Enrique Macpherson, 

Centre D’Estudios Avancats de Blanes, Spain 

DECAPODA: ANOMURA 

I can only address your classification of the Anomura. 

Forest (1987a, b), while concurring with 

McLaughlin’s (1983) argument that the Paguridea 

represented a monophyletic taxon, did not agree 

with her elimination of the Coenobitoidea as a superfamily. 

Consequently he elevated the Paguridea 

to rank of Section and reinstated the superfamily 

Coenobitoidea to include the families Pylochelidae, 

Diogenidae and Coenobitidae. He did concur with 

McLaughlin’s removal of the Lomidae and its elevation 

to superfamily. He did not address the hierarchical 

ranking of the other Anomuran superfamilies. 

McLaughlin and Lemaitre (1997) acknowledged 

Forest’s sectional ranking for the Paguridea, 

but continued to refer to all of the 

anomuran major taxa as superfamilies. However, 

Forest et al. (2000), Forest and McLaughlin (2000), 

and de Saint Laurent and McLaughlin (2000) all 

refer to the superfamilies Coenobitoidea and Paguroidea, 

under the Section Paguridea. 

I personally still believe that the Paguridea represent 

a monophyletic taxon; however, I also believe 

that Forest’s argument for reinstatement of the 

Coenobitoidea is valid. For hierarchical balance 

within the Anomura, perhaps the other superfamilies 

should similarly be elevated to Section rank in 

your classification. 


Forest, J. 1987a. Les Pylochelidae ou ‘‘Pagures symetriques’’ 

(Crustacea Coeno-bitoidea). In Résultats des 

campagnes MUSORSTOM. Mémoires du Muséum 

National d’Histoire Naturelle, série A, Zoologie, vol. 

137, 1–254, figs. 1–82, plates 1–9. 

. 1987b. Ethology and distribution of Pylochelidae 

(Crustacea Decapoda Coenobitoidea). Bulletin of 

Marine Science 41(2):309–321. 

Forest, J., M. de Saint Laurent, P. A. McLaughlin, and R. 

Lemaitre. 2000. The marine fauna of New Zealand: 

Paguridea (Decapoda: Anomura) exclusive of the 

Lithodidae. NIWA Biodiversity Memoir 114 (in 

press). 

Forest, J., and P. A. McLaughlin, 2000. Superfamily Coenobitoidea. 

In The marine fauna of New Zealand: 

Paguridea (Decapoda: Anomura) exclusive of the 

Lithodidae, eds. J. Forest, M. de Saint Laurent, P. A. 

McLaughlin, and R. Lemaitre. NIWA Biodiversity 

Memoir 114. 

McLaughlin, P. A. and R. Lemaitre. 1997. Carcinization 

in the Anomura—fact or fiction? I. Evidence from 

adult morphology. Contributions to Zoology, Amsterdam 

67(2):79–123, figs. 1–13. 

Saint Laurent, M. de, and P. A. McLaughlin, 2000. Superfamily 

Paguroidea, Family Paguridae. In The ma- 

rine fauna of New Zealand: Paguridea (Decapoda: 

Anomura) exclusive of the Lithodidae, eds. J. Forest, 

M. de Saint Laurent, P. A. McLaughlin, and R. Lemaitre. 

NIWA Biodiversity Memoir 114. 

Submitted by Patsy McLaughlin, 

Shannon Point Marine Center, 

Anacortes, Washington 

DECAPODA: BRACHYURA 

As before, I think that the Oregoninae of Garth 

should be elevated to a family. I contacted Michel 

Hendrickx about the classification. He in turn 

quoted a paper that provided larval evidence for 

the distinction of the group as a family, and said 

that he will treat the group as such in his forthcoming 

work on crabs. Please contact Michel for further 

information. If you cannot contact him, let me 

know and I’ll find that larval paper for you. My 

own suspicion is that the oregoniids are not covered 

in most monographs because they are a cirumArctic 

and boreal northern hemisphere group that does 

not range at all into tropical waters, where most 

researchers work! 

Submitted by Mary K. Wicksten, 

Texas A&M University 


I strongly believe that the Pinnotheridae are not 

monophyletic. So if I argued that this family 

‘‘should remain in the Thoracotremata based on evidence 

from DNA sequencing’’ [as cited in your 

classification], I should add that this might only be 

true for some of its constituent subfamilies or genera. 

My statement was made based on the phylogenetic 

position of Pinnixa in molecular analyses 

that showed a strikingly close relationship to the 

Ocypodinae (Schubart et al., 2000a). 

I also think that the Ocypodidae in the traditional 

sense as well as the Ocypodoidea as defined in 

the latest draft of your classification might not be 

monophyletic. Molecular as well as larval morphological 

data suggest a close relationship between the 

Varunidae (Grapsoidea) and the Macropthalminae 

(Schubart et al., 2000a; Schubart and Cuesta, unpublished). 

I think that this possible phylogenetic 

link would be another reason to elevate ocypodid 

subfamilies to family level as already considered in 

your draft and suggested for the Grapsidae (Schubart 

et al., 2000b). This would certainly make justice 

to ocypodoid morphological diversity and allow 

a more objective comparison with other thoracotremes 

in the future. 

I disagree on the use of the superfamily name 

‘‘Grapsidoidea.’’ Since the stem of the name is 

Graps- (based on Cancer grapsus Linnaeus, see also 

family name Grapsidae) and the ending for superfamilies 

is -oidea, the superfamily should be called 

Grapsoidea (and not Grapsidoidea). The fact that 

the term Grapsoidea has been used in the past for 

a much wider systematic grouping of eubrachyuran 


crabs and is now restricted to the families Grapsidae, 

Gecarcinidae, Plagusiidae, Searmidae, and Varunidae 

should not influence the nomenclature. 


Schubart, C. D., J. A. Cuesta, R. Diesel, and D. L. Felder. 

2000b. Molecular phylogeny, taxonomy, and evolution 

of non-marine lineages within the American 

Grapsoidea (Crustacea: Brachyura). Molecular Phylogenetics 

and Evolution (in press). 

Schubart, C. D., J. E. Neigel, and D. L. Felder. 2000a. The 

use of the mitochondrial 16S rRNA gene for phylogenetic 

and population studies of Crustacea. Crustacean 

Issues 12 (in press). 

Submitted by Christoph Schubart, 

Universität Regensburg, Germany 


Although recently I published my arrangement of 

the brachyuran families, I have some new discoveries 

in the brachyuran classification, but it is not 

finished and it will be published next year. I was 

able to classify all dromiacean families into superfamilies, 

but not the eubrachyuran ones, because 

there are many families with obscure systematic position: 

Orithyiidae, Calappidae, Matutidae, Astenognathidae, 

Hexapodidae, Palicidae, Dairodidae 

and many up to now undescribed families (Acidopidae, 

Melybiidae, Speocarcinidae, etc.). Here are 

some of my remarks. 

(1) Dynomenidae are the most primitive Dromioidea, 

because only the last pair of legs is aberrant. 

(2) Among Homoloidea, the Poupinidae are the 

most primitive because the last pair of legs are of 

‘‘normal’’ structure but are partly subdorsal in position. 

(3) Raninidae are ‘‘Podotremata’’ (i.e. Dromiacea) 

because their sexual openings in both sexes 

are on the coxae of the legs (hence the name Podotremata). 

(4) The most primitive eubrachyuran 

family is the Atelecyclidae, because they have the 

antennules and antennae longitudinally directed, a 

narrow thoracic sternum, thoracic sternites 4/5–7/8 

continuous (entire), and sternites nearly regularly 

metamerized. (5) The Dorippidae are highly derived 

and aberrant: the dorsal position of the posterior 

pair of legs, the sternite 8 facing dorsally, and 

the narrowed buccal cavern all are secondarily attained. 

The similarity with the Dromiacea is thus 

superficial. (6) The same could be said for the Leucosiidae: 

highly derived crabs and consequently 

should be placed at the end of the classificatory 

scheme of the Heterotremata. (7) The Majidae are 

only one family with many subfamilies. The arrangement 

is enclosed [Sˇtevčić, Z. 1994. Contribution 

to the re-classification of the family Majidae. 

Periodicum Biologorum 96:419–420]. (8) The 

Parthenopidae are more primitive than Majidae, 

and therefore should be ahead of the Majidae. (9) 

The Retroplumidae are a very derived brachyuran 

family. (10) Geryonidae have a similar organization 

to the Goneplacidae s.s. (11) Your Xanthoidea is a 

highly polyphyletic group. (a) The most primitive 

‘‘xanthoids’’ are the Eriphiidae, not Menippidae! 

The most primitive Eriphiidae have sternites 4/5– 

7/8 entire, abdominal segments freely articulated in 

both sexes, and the second gonopod longer than the 

first. They are probably related to Trapeziidae. In 

the same assemblage with the Eriphiidae are the 

Pilumnoididae Guinot and Macpherson, 1987. (b) 

Xanthidae s.s. have [some] primitive representatives 

(Krausinae, with sternal sutures 4/5–7/8 entire), 

but abdominal segments 3–5 in the male are 

fused, and the second gonopod is short. They are 

related to the Panopeidae/Panopeinae and the Pseudorhombilidae. 

(c) Pilumnidae have a primitive abdomen 

(all segments freely articulated in both sexes) 

but specific first and second gonopods, the latter 

short. They are related to the Eumedonidae (in fact 

the Eumedoninae). (d) Goneplacidae s.s. are in fact 

a very small taxon, without any close relationships 

with the Xanthidae. They are probably close to the 

Geryonidae and Euryplacidae/Euryplacinae. (12) 

The Potamidae are in fact a very difficult problem, 

however the gaps among subfamilies are not quite 

distinct. The gaps are not always [clear] and therefore 

the separation of the freshwater crabs into 

families remains uncertain. (13) I think that between 

Ocypodidae and Mictyridae and between 

Grapsidae and Gecarcinidae the gaps are not decisive 

and only Ocypodidae and Grapsidae are true 

families (this will be published later). (14) Finally, 

I think that the Cancroidea are not a taxon, they 

are only a grade, not a clade (taxon i.e., monophyletic 

group). (15) Hepatinae are a subfamily of the 

family Aethridae. (16) Palicidae belong to the Heterotremata, 

with no close affinity with the Ocypodidae. 

Submitted by Zdravko Sˇtevčić, 

Rudjer Boskovic Institute, Croatia 


Concerning my special knowledge, the Brachyura, 

I do not agree with all decisions (see my responses), 

but I respect them. May I add my feeling, however. 

Concerning the Podotremata, the molecular data 

seem to outweigh all other considerations, despite 

the fact that the first results (Spears and Abele, 

1988; Spears, Abele, and Kim, 1992) were fragmentary, 

based only on very few taxa (only two 

Dromiidae were studied; and the conclusion was 

made without any Dynomenidae, Homolodromiidae, 

Homolidae, Latreilliidae, Cyclodorippidae, 

Cymonomidae, nor Phyllotymolinidae) and that the 

new results are not yet published. I am happy to 

see that Spears now returns to the opinion that the 

Dromiidae are true Brachyura, but we wait her paper 

where the new demonstration is given. 

Concerning your Section Raninoida, you write 

(p. 66, 69) that there is ‘‘possibly a mistake.’’ I recognize 

that the problem of the placement of on the 

one hand Cyclodorippidae, Cymonomidae, and 

Phyllotymolinidae, and on the other hand the Ran- 


inoidea is difficult, because they do not clearly enter 

in a major group. You write that, for Spears herself 

(p. 69), ‘‘molecular data seem to indicate a placement 

[of Cyclodorippoidea] somewhere between 

the raninids and the higher eubrachyurans.’’ So, the 

molecular data exactly give the same results that 

the morphological and ontogenetic ones. The two 

groups Raninoidea and Cyclodorippoidea (the last 

name is used by convenience, but perhaps they 

form three distinct families, see Tavares) seem 

apart, but where is the best way? 

Submitted by Danièle Guinot, 

Muséum National d’Histoire Naturelle, Paris 


Evidence from morphology and larval development 

points to the polyphyletic nature of the Trapeziidae. 

There are three separate groups: one comprises Trapezia, 

Quadrella, Hexagonalia, Calocarcinus, Philippicarcinus 

and Sphenomerides, a second Tetralia 

and Tetraloides, and a third Domecia, Jonesius, 

Palmyria and Maldivia. 

Submitted by Peter Castro, 

California State Polytechnic University, Pomona 


I disagree that all Brachyura with female gonopores 

on P3 coxa and with spermathecae at the extremities 

of thoracic sutures 7/8 are separated in two 

different major sections, Dromiacea and Eubrachyura, 

with the Raninoidea and Cyclodorippoidea 

distributed in a basal group inside the Eubrachyura. 

In that case, how to make a definition of both 

Dromiacea and Eubrachyura as a whole? The Podotremata 

may receive all Brachyura with female 

gonopores on P3 coxa and with spermathecae at 

the extremities of thoracic sutures 7/8, i.e., two different 

apertures. The Eubrachyura may receive all 

Brachyura with a sternal location of female gonopores 

(vulvae on the thoracic sternum, sternite 6); 

there is now a sole female orifice for reproduction 

(egg laying, intromission of male pleopod, and storage 

of the spermatozoas). Another synapomorphy 

(among others) of the assemblage Heterotremata- 

Thoracotremata is the morphology of the first male 

pleopod, which is completely closed and provided 

with two distinct basal foramina (instead of only 

one in the Podotremata). To concile the evident 

apart position of the Raninoidea and Cyclodorippoidea 

(but, perhaps consider three distinct families: 

Cyclodorippidae, Cymonomidae, Phyllotymolinidae), 

I suggest to range them among the Podotremata 

in Archaeobrachyura Guinot, 1977 emend. 

(i.e. with the exclusion of the Homoloidea). 



DECAPODA: BRACHYURA: DROMIACEA 

I disagree that the section Dromiacea contains the 

Homoloidea. The Dromiacea and Homoloidea are 

two different lineages. I suggest to consider a Section 

Podotremata, with three subsections: Subsection 

Dromiacea, containing two superfamilies 

Homolodromioidea (Homolodromiidae) and 

Dromioidea (Dromiidae, Dynomenidae); Subsection 

Homoloidea (Homolidae, Latreilliidae, Poupiniidae); 

Subsection Archaeobrachyura (Cyclodorippidae, 

Cymonomidae, Phyllotymolinidae, and 

Raninidae). The monophyly of the Dromiacea is 

well supported by many features; the same for 

Homoloidea. I recognize that the monophyly of the 

Archaeobrachyura emend. (without the Homoloidea) 

is not so well supported and that these crabs 

show puzzling features, but they are all very specialized 

and modified by the burrowing life. Their 

attribution to the Podotremata is, at least for the 

moment, supported by the appendicular location of 

female gonopores (on P3 coxa) and the spermathecae 

at the extremities of thoracic sutures 7/8, the 

features of the sternal plate, the arthrodial cavities 

of the pereiopods, and others characters. If we include 

the Cyclodorippidae, Cymonomidae, Phyllotymolinidae, 

and the Raninidae in the Eubrachyura, 

which becomes the diagnosis of the Eubrachyura? 



DECAPODA: BRACHYURA: 

HETEROTREMATA, THORACOTREMATA 

It is important to recall the original definition of 

the taxa given by Guinot (1977, 1978). 

The section Hererotremata contains the Brachyuran 

families, ALL THE MEMBERS of which 

are sternitreme for the female gonopores, and 

ONLY some members, at least, are podotreme 

for the male gonopores. 

The section Thoracotremata contains the Brachyuran 

families, all the members of which are sternitreme 

for the female and male gonopores. It 

means that, for the Heterotremata, in the Leucosiidae 

or Leucosioidea by example it exists members 

with male gonopores on the P5 coxa and other 

members with sternal male apertures. But, in the 

last case, it is only a coxo-sternal location of the 

penis. The same is true for the Dorippidae, where 

some members show a coxo-sternal location of the 

penis. 




Abele, Lawrence G. 

Baba, Keiji 

*Belk, Denton 

Bousfield, Ed 

Boxshall, Geoff 

Brandt, Angelika 

Brendonck, Luc 

Briggs, Derek 

Brusca, Gary 

Brusca, Richard 

Cadien, Don 

Camp, David 

Castro, Peter 

Causey, Douglas 

Chace, Fenner 

Christoffersen, Martin 

Clark, Paul 

Cohen, Anne 

Crandall, Keith 

Crosnier, Alain 

Cumberlidge, Neil 

*Dahl, Erik 

Dahms, Hans-Uwe 

Davie, Peter 

Elofsson, Rolfe 

Felder, Darryl L. 

Feldmann, Rodney 

Felgenhauer, Bruce 

Fryer, Geoffrey 

Galil, Bella 

Grygier, Mark 

Guinot, Danièle 

Haney, Todd 

Harvey, Alan 

APPENDIX II. LIST OF CONTRIBUTORS 

The following are colleagues who graciously gave of their time to review various 

drafts of the Classification of Recent Crustacea. 

* Denotes researchers recently deceased. 

Hayashi, Ken-Ichi 

Heard, Richard 

Hendrickx, Michel 

Hessler, Robert 

Ho, Ju-Shey 

Høeg, Jens 

Hof, Cees 

Holsinger, John 

Holthuis, Lipke B. 

*Humes, Arthur 

Huys, Rony 

Jamieson, Barry 

Jones, Diana S. 

Kaesler, Roger 

Kensley, Brian 

Kornicker, Lou 

Larsen, Kim 

LeCroy, Sara 

Lemaitre, Rafael 

Lowry, Jim 

MacPherson, Enrique 

Maddocks, Rosalie 

*Manning, Ray 

Markham, John 

McLaughlin, Pat 

Mickevich, Mary 

Modlin, Richard 

Morgan, Gary 

Myers, Alan 

Newman, William 

Ng, Peter 

Olesen, Jørgen 

Poore, Gary 

Regier, Jerome C. 

Rice, Tony 

Richer de Forges, Bertrand 

Richter, Stefan 

Riley, John 

Sakai, Katsushi 

St. Laurent, Michele de 

Schminke, Horst 

Scholtz, Gerhard 

Schram, Frederick 

Secretan, Sylvie 

Schubart, Christoph 

Sorbe, Jean Claude 

Spears, Trisha 

Sˇtevčić, Zdravko 

Takeuchi, Ichiro 

Tavares, Marcos 

Thomas, James D. 

Tudge, Christopher 

Vereshchaka, Alexander 

Vervoort, W. 

Wägele, Wolfgang 

Wallis, Elycia 

Walossek, Dieter 

Watling, Les 

Whatley, Robin 

Wicksten, Mary K. 

*Williams, Austin 

Wilson, Buz 

Yager, Jill 

Young, Paulo 

Zimmerman, Todd L. 

114 � Contributions in Science, Number 39 Appendix II: List of Contributors

APPENDIX III. OTHER CRUSTACEAN RESOURCES 

This appendix is subdivided into four sections. Section 

III-A contains a list of journals and newsletters 

and their current editors and addresses. Section 

III-B is an alphabetical list of currently active web 

sites and their URLs, followed by a short selection 

of ‘‘personal pages’’ of some workers with crustacean 

information on their web sites. Section III-C 

is a list of crustacean-related listservers. Section 

III-D is a list of natural history museums with significant 

crustacean holdings, some of which have 

searchable crustacean databases. 

III-A. JOURNALS AND NEWSLETTERS 

1. JOURNALS 

Journals that publish only crustacean-specific articles 

are rather few and currently include only the 

following (listed alphabetically). 

Crustaceana 

Description: ‘‘International Journal of Crustacean 

Research’’ publishing ‘‘papers dealing with Crustacea, 

from all branches of Zoology.’’ Issued eight 

times per year (January, February, March, April, 

June, July, September, October, November, and December). 

Current editor and address: J. C. Von Vaupel 

Klein, Crustaceana, Editorial Board Administrative 

Office, Beetslaan 32, NL-3723 DX, Bilthoven, The 

Netherlands. 

Publisher: Brill Academic Publishers, Inc., Leiden, 

The Netherlands. 

Crustacean Issues 

Description: An irregular series of collections of papers 

on Crustacea, each published as a hardbound 

volume and covering a discrete crustacean topic. 

Current editor and address: General editor, Frederick 

R. Schram, Zoological Museum, University of 

Amsterdam. 

Publisher: A. A. Balkema, Rotterdam, The Netherlands. 

Crustacean Research (formerly Researches on 

Crustacea) 

Description: A publication of the Carcinological 

Society of Japan, publishing papers dealing with 

‘‘any aspect of the biology of Crustacea.’’ Issued 

quarterly. 

Current editor: Keiji Baba, Crustacea Research, 

Faculty of Education, Kumamoto University, 860– 

8555, Japan. 

Publisher: Carcinological Society of Japan and 

Shimoto Printing, Kumamoto. 

Journal of Crustacean Biology 

Description: The official journal of The Crustacean 

Society, ‘‘for the publication of research on any aspect 

of the biology of Crustacea.’’ Issued quarterly. 

Current editor: David K. Camp, Journal of Crustacean 

Biology, P.O. Box 4430 Seminole, Florida 

33775–4430, USA. 

Publisher: The Crustacean Society and Allen 

Press, Lawrence, Kansas. 

Nauplius (Revista da Sociedade Brasiliera de 

Carcinologia) 

Description: The journal of the Sociedade Brasileira 

de Carcinologia, publishing ‘‘original papers based 

on research in any aspect of crustacean biology, including 

taxonomy, phylogeny, morphology, development, 

physiology, ecology, biogeography, bioenergetics, 

aquaculture and fisheries biology.’’ Issued 

quarterly. 

Current editor: Mónica A. Montú, Nauplius, Laboratorio 

de Carcinologia, Departamento de 

Oceanografia—FURG, Caixa Postal 474, CEP 

96201–900, Rio Grande, RS, Brazil. 

Publisher: Sociedade Brasileira de Carcinologia. 

There are of course many more journals that 

publish taxonomic/systematic/phylogenetic studies 

of crustaceans along with papers on other invertebrate 

groups. We conducted an informal survey of 

the subscribers to the crustacean listserver CRUST- 

L in March of 2000 and asked members to name 

the journals they consult on a regular basis for new 

information on crustacean relationships. The following 

journals, arranged alphabetically, were all 

mentioned more than once in that survey: Acta 

Zoologica, Arthropoda Selecta, Biological Bulletin, 

Bulletin of Marine Science, Canadian Journal of 

Zoology, Comptes Rendus de l’Academie des Sciences, 

Contributions to Zoology (University of Amsterdam), 

Deep-Sea Research, Evolution, Fishery 

Bulletin (US), Fossils and Strata, Gulf and Caribbean 

Research (formerly Gulf Research Reports), 

Hydrobiologia, Invertebrate Biology, Invertebrate 

Reproduction and Development, Invertebrate Taxonomy, 

Journal of Experimental Marine Biology 

and Ecology, Journal of the Marine Biological Association 

of the United Kingdom, Journal of Natural 

History, Journal of Plankton Research, Marine 

Biology, Marine Ecology Progress Series, Memoirs 

du Museum National d’Histoire Naturelle (Paris), 

Memoirs of the Museum of Victoria, Proceedings 

of the Biological Society of Washington, Proceedings 

of the Linnean Society of New South Wales, 

Proceedings of the Royal Society of London (series 

B), Raffles Bulletin of Zoology, Revista di Biologia 

Tropical, Sarsia, Smithsonian Contributions to Zoology, 

Zoologica Scripta, Zoological Journal of the 

Linnean Society, Zoologischer Anzeiger, Zoosystema. 

Contributions in Science, Number 39 Appendix III: Other Crustacean Resources � 115

2. NEWSLETTERS 

Included here are some of the more taxonomically 

or systematically oriented crustacean newsletters of 

which we are aware. We have purposely avoided 

listing newsletters that primarily target aspects of 

crustacean farming, aquaculture, and the aquarium 

trade. 

Amphipod Newsletter (see also the Amphipod 

Homepage) 

Editors as of March 2001: Jim Lowry and Wim 

Vader 

Address: Sydney, Australia (Jim Lowry); Tromso, 

Norway (Wim Vader) 

Homepage: http://web.odu.edu/sci/biology/ 

amphome/ 

Anostracan News (Newsletter of the IUCN/SSC 

Inland Water Crustacean Specialist Group) 

Editor as of March 2001: Denton Belk 

Address: 840 E. Mulberry Avenue, San Antonio, 

Texas 78212–3194, USA 

Homepage: none to our knowledge 

Boletin de la Association Latinoamericana de 

Carcinologia 

Editor as of March 2001: Guido Pereira (gpereira@ 

strix.ciens.ucv.ve) 

Address: Instituto de Zoologia Tropical, Universidad 

Central de Venezuela, Caracas, Venezuela 

Homepage: http://tierradelfuego.org.ar/alca/ 

Coral Reef Newsletter 

Editors as of March 2001: C. E. Birkland and L. 

G. Eldredge 

Address: Pacific Science Association, P.O. Box 

17801, Honolulu, Hawaii 96817, USA 


Crayfish News (Official Newsletter of the 

International Association of Astacology) 

Editor as of March 2001: Glen Whisson 

(twhisson@alpha2.curtin.edu.au) 

Address: IAA Secretariat, P.O. Box 44650, University 

of Louisiana at Lafayette, Lafayette, Louisiana 

70504, USA (jhuner@usl.edu) 

Homepage: http://www.uku.fi/english/ 

organizations/IAA/ 

Cumacean Newsletter 

Editors as of March 2001: Daniel Roccatagliata 

(rocca@bg.fcen.uba.ar), Richard W. Heard, Magdalena 

Blazewicz, and Ute Mühlenhardt-Siegel 

Address: (for Roccatagliata) Departamento de 

Biologia, Universidad de Buenos Aires, Ciudad Universitaria-Nunex, 

1428 Buenos Aires, Argentina 

Homepage: http://www.ims.usm.edu/cumacean/ 

index.html 

Cypris (Newsletter for Ostracodologists) 

(formerly The Ostracodologist: Newsletter for Ostracod 

Workers) 

Editor as of March 2001: Elisabeth M. Brouwers 

Address: See home page for regional representative 

Homepage: http://www.uh.edu/�rmaddock/ 

IRGO/cypris.html 

Ecdysiast (Official Newsletter of The Crustacean 

Society) 

Editor as of March 2001: Tim Stebbins (TDS@ 

sdcity.sannet.gov) 

Address: City of San Diego Marine Biology Laboratory, 

4918 N. Harbor Dr., Suite, 101, San Diego, 

California 92106, USA 

Homepage: http://www.lam.mus.ca.us/�tcs/ 

ecdysiast.htm 

(The) Isopod Newsletter 

Editor as of March 2001: Brian Kensley (kensley. 

brian@nmnh.si.edu) 

Address: Department of Invertebrate Zoology, 

NHB-163, Smithsonian Institution, Washington, 

D.C. 20560–0163, USA 


(The) Lobster Newsletter 

Editor as of March 2001: Mark Butler 

Address: Department of Biological Sciences, Old 

Dominion University, Norfolk, Virginia 23529– 

0266, USA 

Homepage: none 

Monoculus (Copepod Newsletter) 

Editors as of March 2001: Hans-U. Dahms and 

H. Kurt Schminke 

Address: Fachbereich 7 (Biologie), Universitat 

Oldenburg, D-26111, Oldenburg, Germany 

Homepage: http:www.hrz.uni-oldenburg.de/ 

monoculus 

Plankton Newsletter 

Editors as of March 2001: P. H. Schalk (peter@ 

eti.bio.uva.nl) and S. van der Spoel 

Address: P.O. Box 16915, 1001 RK 

Amsterdam, The Netherlands 


SCAMIT Newsletter (Southern California 

Association of Marine Invertebrate Taxonomists) 

Editor as of March 2001: Don Cadien (dcadien@ 

lacsd.org) 

Address: Marine Biology Laboratory, County 

Sanitation Districts of Los Angeles County, 24501 

South Figueroa Street, Carson, California 90745, 

USA 

Homepage: http://www.scamit.org/index.htm 

116 � Contributions in Science, Number 39 Appendix III: Other Crustacean Resources

(The) Stomatopod Newsletter 

Editors as of March 2001: Tatsuo Hamano 

(hamanot@fish-u.ac.jp) and Chris Norman 

(norman@snf.affrc.go.jp) 

Address: National Fisheries University, P.O. Box 

3, Yoshimi, Japan 

Homepage: None 

(The) Tanaidacea Newsletter 

Editors as of March 2001: Richard W. Heard 

(richard.heard@usm.htm) and Gary Anderson 

Address: Institute of Marine Sciences, The University 

of Southern Mississippi, P.O. Box 7000, 

Ocean Springs, Mississippi 39566–7000, USA 

Homepage: http://tidepool.st.usm.edu/tanaids/ 

newsletter98.htm 

Zoea (Larval development newsletter for 

carcinologists) 

Editors as of March 2001: Klaus Anger, José A. 

Cuesta, and Pablo J. López-González 

Address: Departamento de Ecologia, Facultad de 

Biologia, Apdo 1095, E-41080 Sevilla, Spain 

Homepage: http://members.es.tripod.de/ 

Megalopa/index.htm 

III-B. WEB SITES 

Knowing that any such list will become obsolete 

even before it is published because of the rapid 

growth of web sites in various areas of invertebrate 

biodiversity, we nevertheless offer here some of the 

more useful crustacean-related web sites of which 

we were aware at the time of printing. Although 

some of the sites were useful in constructing the 

current classification, listing below does not necessarily 

indicate our endorsement nor does it necessarily 

indicate that the authors of any of these sites 

are in agreement with the currently proposed classification. 

This list is far from exhaustive. It is meant to 

provide an introduction to the large and ever-growing 

number of web sites that may be of interest to 

students of carcinology. Additionally, the list excludes 

a number of ‘‘personal’’ sites (such as those 

of Colin MacLay, Jeff Shields, Dieter Walossek, and 

others), some of which are quite interesting and 

contain a lot of information on crustaceans as well. 

A brief selection of these personal sites is given after 

the alphabetized web page list. 

About Phreatoicidean Isopods in Australia 

http://www-personal.usyd.edu.au/�buz/ 

popular.html 

A site devoted to these fascinating crustaceans, 

maintained by George (Buz) Wilson, Australian 

Museum. 

(The) Amphipod Homepage 

http://www.odu.edu/�jrh100f/amphome/ 

Maintained by Stefan Koenemann at Old Dominion 

University, Norfolk, Virginia. Nice introductory 

page leading to the ‘‘Amphipod Newsletter,’’ 

web sites related to amphipods, pictures of 

amphipods, and various sites about crustacean biology. 

Animal Diversity Web 

http://www.oit.itd.umich.edu/bio108/Arthropoda/ 

Crustacea.shtml 

This address takes you to the Crustacea pages of 

the University of Michigan’s Animal Diversity web 

site. Provides general information on several classes, 

primarily geared to the nonspecialist. 

Animal Evolutionary Pattern Analysis Home Page 

http://www.bio.uva.nl/onderzoek/cepa/ 

Default.html#LL 

Presents the research activities of a group of scientists 

allied to the Institute for Systematics and 

Population Biology, a research institute within the 

Faculty of Biology of the University of Amsterdam, 

with links to their ongoing arthropod and crustacean 

projects. 

Animals4ever 

http://www.animals4ever.com/ 

A searchable and interactive listing, with figures 

and references, maintained in Belgium, with the 

goal of eventually grouping ‘‘all animals on the web 

in one place.’’ 

Ant’phipoda, The Antarctic Marine Biodiversity 

Reference Center Devoted to Amphipod 

Crustaceans 

http://www.naturalsciences.be/amphi/ 

Managed by the Laboratory of Carcinology at 

the Royal Belgian Institute of Natural Sciences, 

with links to the checklist of amphipods of the 

Southern Ocean, amphipodologists involved with 

Antarctic fauna, research activities, pictures, and 

numerous amphipod sites. 

(The) Appalachian Man’s Crayfish Photo Gallery 

http://webby.cc.denison.edu/�stocker/ 

cfgallery.html 

Many color crayfish photographs, plus links to 

other crayfish sites. Maintained by Whitney Stocker 

of Gunison University, Ohio, USA. 

Biographical Etymology of Marine Organism 

Names (BEMON) 

http://www.tmbl.gu.se/libdb/taxon/personetymol/ 

index.htm 


An interesting site attempting to track the history 

of taxonomic names of marine species, including 

crustaceans. Maintained by Hans G. Hansson. 

Biology of Copepods 

http://www.uni-oldenburg.de/zoomorphology/ 

Biology.html#biotable 

A page maintained by Thorsten D. Künnemann, 

with an introduction to the biology of copepods, 

scanning electron micrographs, copepod systematics, 

and anatomy of copepods (in preparation). 

Biomedia Home Page 

http://www.gla.ac.uk/Acad/IBLS/DEEB/biomedia/ 

home/home.htm 

Very general information on crustaceans (and 

other taxa) for the nonspecialist. 

BIOSIS—Internet Resource Guide for Zoology 

(Crustacea) 

(see Zoological Record) 

Biospeleology Home Page: The Biology of Caves, 

Karst, and Groundwater 

http://www.utexas.edu/depts/tnhc/.www/ 

biospeleology/ 

Provides information on some cave crustaceans. 

This site is maintained by the Texas Memorial Museum 

in Austin. 

(The) Blue Crab Home Page 

http://www.blue-crab.net/ 

A useful and large resource page, with connections 

to literature, other sites about blue crabs, and 

other researchers interested in nearly all aspects of 

the blue crab, Callinectes sapidus. Maintained by 

Vince Guillory. 

British Marine Life Study Society 

http://cbr.nc.us.mensa.org/homepages/BMLSS 

Brief reports of British marine life, with occasional 

reports of crustaceans and links to other marine 

life sites. 

Canadian Museum of Nature’s Database of 

Canadian Arthropod (excl. Insects) Systematists 

http://www.nature.ca/english/arthro.htm 

A database of Canadian systematists, with scientists 

organized by area of expertise in arthropods 

(excluding insects). 

Central Terminal for Crustacean Neuroscience 

http://wwwzoo.kfunigraz.ac.at/crusties.html 

A valuable site for everything related to crusta- 

cean neurology, with many interesting links to related 

sites. 

Cercopagis pengoi Page (Cladoceran) 

http://www.ku.lt/nemo/cercopag.htm 

A reference page for this cladoceran species; part 

of the Baltic Research Network on Ecology and 

Marine Invasions and Introductions, Estonian Marine 

Institute, Tallinn, Estonia. Contains taxonomic 

information, diagnosis, line drawings and color 

photographs, information on population dynamics, 

and references. 

Cladocera 

http://www.cladocera.uogluelph.ca/ 

This site, maintained by Paul Hebert, provides a 

variety of information useful for cladoceran researchers 

and others interested in the Cladocera. 

Includes pages on taxonomy, references, researchers, 

specimen wish lists, tools, and meetings. 

Copepods and Groundwater Biology 

http://www.uni-oldenburg.de/zoomorphology/ 

Groundwater.html 

Maintained by the Zoomorphology Section at 

the University of Oldenburg, this is an overview 

page with links to Giuseppe Pesce’s various groundwater 

biology sites. 

Crabs Found in Belgium Waters 

http://uc2.unicall.be/RVZ/CrabBook.html 

A clever, useful sight for learning about crabs in 

this part of the world. Click on any crab for further 

information. 

(The) Crayfish (T. H. Huxley, 1879, 1880) 

http://www.biology.ualberta.ca/palmer.hp/thh/ 

crayfish/htm 

T. H. Huxley’s classic paper on crayfish in its entirety, 

including all of the original woodcut illustrations, 

available online courtesy of Eric Eldred and 

the University of Alberta, Canada. 

(The) Crayfish Corner 

http://www.mackers.com/crayfish 

A lay person site with general information about 

crayfish, their appearance, behavior, internal anatomy, 

pictures, and more. 

Crayfish Home Page 

http://bioag.byu.edu/mlbean/CRAYFISH/ 

crayhome.htm 

Keith Crandall’s website highlighting lab personnel, 

publications and data, computer programs, lab 

links, lab tour, extensive crayfish photo gallery, and 


links to crustacean societies, conservation, and 

more. 

Crustacea Gopher (U.S. National Museum, 

Smithsonian) 

gopher://nmnhgoph.si.edu:70/11/.invertebrate/. 

crustaceans 

The gopher menu allows access to ‘‘Crayfish,’’ 

‘‘Isopods,’’ and the ‘‘CRUST-L Discussion Group 

Digests.’’ The ‘‘Crayfish’’ contains 13,000 searchable 

references. For isopods, see listing under 

‘‘World List of Marine, Freshwater, and Terrestrial 

Isopod Crustaceans.’’ 

Crustacea of Lake Biwa 

http://www.hirano-es.otsu.shiga.jp:80/fish-e.html 

Images and Japanese names of freshwater crustaceans 

in Lake Biwa. 

Crustacea Net 

http://www.crustacea.net 

Hosted on the Australian Museum website maintained 

by Jim Lowry, the DELTA (DEscription Language 

for TAxonomy) taxonomic computer program 

provides illustrated and interactive keys to 

identify higher Crustacea taxa, with keys to crustacean 

families. 

Crustacea Node of the Tree of Life Project 

http://phylogeny.arizona.edu/tree/eukaryotes/ 

animals/arthropoda/crustacea/crustacea.html 

This will take you directly to the Crustacea part 

of David and Wayne Maddison’s Tree of Life project. 

The crustacean section currently is based on 

Brusca and Brusca (1990). 

Crustacean Disease Information 

http://www.geocities.com/CapeCanaveral/Lab/ 

7490/index.html#crustdis 

Part of the Aquaculture Health Page, maintained 

by Bill Lussier. 

(The) Crustacean Biodiversity Survey 

http://www.nhm.org/cbs/ 

A site of general interest that includes a searchable, 

additive, database. 

(The) Crustacean Society 

http://www.vims.edu/tcs 

The Crustacean Society Home Page, maintained 

by Jeff Shields and hosted by the Virginia Institute 

of Marine Science. 

Crustacean Specimens of the Marine Biological 

Laboratory 

http://database.mbl.edu/SPECIMENS/phylum. 

taf?function�search&find 

�Arthropoda 

Crustacean specimens in the collections of the 

Marine Biological Laboratory, Woods Hole, Massachusetts. 

Crustacés Polynésiens 

http://biomar.free.fr/ 

Provides a list of species and authorships of Indo- 

Pacific taxa; many entries are represented with photographs. 

Maintained by J. Poupin. 

Cryptofauna of Empty Barnacle Shells and Lego 

Plastic Blocks 

http://www.ex.ac.uk/biology/adrianc.html 

Strange but true, an interesting site on an obscure 

topic, maintained by Adrian Clayton. 

Cumacean Home Page 

http://nature.umesci.maine.edu/cumacea.html 

A product of a PEET grant from the U.S. National 

Science Foundation, this site is maintained 

by Les Watling and Irv Kornfield (and students) at 

the University of Maine. 

Directory of Copepodologists 

http://www.univaq.it/�sc�amb/wac.html 

Self explanatory; this is a subpage of the Monoculus 

site. 

Diversity and Geographical Distribution of Pelagic 

Copepoda 

http://www.obs-banyuls.fr/RAZOULS/WEBCD/ 

accueil.htm 

A pelagic copepod site maintained by Claude Razouls 

and Francis de Bovée at the Observatoire 

Océanologique de Banyuls, France. 

European Register of Marine Species 

http://www.erms.biol.soton.ac.uk/ 

A register of marine species in Europe established 

to facilitate marine biodiversity research and management. 

Contains checklists of European species, 

including most of the major groups of crustaceans. 

Ellis and Messina Catalogue of Ostracoda 

http://www.micropress.org 

An electronic version of the former looseleaf catalogue 

from the American Museum of Natural History 

(Micropaleontology Press). Visitors must go to 

the catalogues section of the site. 


Epicaridea Page 

http://www.vims.edu/�jeff/isopod.htm#Epicaridea 

A thorough page devoted to parasitic isopods, 

maintained by Jeff Shields, Virginia Institute of Marine 

Science. 

(The) Expert Center for Taxonomic Identification 

(ETI) 

http://wwweti.eti.bio.uva.nl/ 

A nongovernmental organization working with 

UNESCO and sponsored by the Netherlands Organization 

for Scientific Research (NWO), the University 

of Amsterdam, and UNESCO. Includes the 

World Biodiversity Database (under construction), 

World Taxonomists Database, and UNESCO-IOC 

Register of Marine Organisms. 

Fiddler Crabs 

http://www.public.asu.edu/�mrosenb/Uca/ 

A fiddler crab web site maintained by Mike Rosenberg, 

with 1,700 references, color photographs, 

and systematic information, mostly from his recent 

(2000) dissertation. 

Génétique et Biologie des Populations de 

Crustacés 

http://labo.univ-poitiers.fr/umr6556/ 

A research program in genetics and population 

biology of crustaceans organized through the Université 

de Poitiers, France. 

Glossary of Morphological Terms 

http://www.nhm.org/lacmnh/departments/research/ 

invertebrates/crustacea/cbs/Glossary�of� 

Morphological�Terms/index.shtml 

A page of the Crustacean Biodiversity Survey, 

this will eventually be the largest existing glossary 

of crustacean terminology. Contains multiple definitions 

put forth by various authors. 

Groundwater Biology 

http://www.geocities.com/�mediaq/fauna.html 

Contains many links to groundwater crustacean 

sites including amphipods, isopods, copepods, remipedes, 

mysids, spelaeogriphaceans, syncarids, 

mictaceans, and others. Some links go to specialists’ 

home pages, others contain lists of taxa, still others 

are in the process of being developed. Maintained 

by Giuseppe L. Pesce. 

(The) International Association of 

Meiobenthologists 

http://www.mtsu.edu/�kwalt/meio/ 

A society representing meiobenthologists in all 

aquatic disciplines, producing a quarterly newslet- 

ter entitled Psammonalia. The site includes several 

photos of live copepods and links to researchers 

(including some with expertise in Crustacea). 

International Research Group on Ostracoda 

http://www.uh.edu/�maddock/IRGO/irgohome. 

html 

Includes links to many useful sites of interest to 

ostracod workers. Maintained by Rosalie Maddocks. 

International Web Site on Terrestrial Isopods 

http://mother.biolan.uni-koeln.de/institute/zoologie/ 

zoo3/terra/homepage.html 

This site was still being constructed as of our last 

check. 

(A) Key to Cladocerans (Crustacea) of British 

Columbia 

http://www.for.gov.bc.ca/ric/Pubs/Aquatic/ 

crustacea/ 

Provides keys to the families Holopedidae, Sididae, 

Daphniidae, Bosminidae, Leptodoridae, and 

Polyphemidae occurring in British Columbia (approximately 

45 species). Published by the Resources 

Inventory Committee of British Columbia. 

Keys to Marine Invertebrates of the Woods Hole 

Region 

http://www.mbl.edu/html/BB/KEYS/KEYScontents. 

html 

Chapters 11, 12, and 13 of this series deal with 

‘‘Lower Crustacea and Cirripedia,’’ ‘‘Pericaridan 

[sic] Crustaceans,’’ and ‘‘Decapod and Stomatopod 

Crustaceans,’’ respectively. 

Laboratory of Aquaculture and Artemia Reference 

Center 

http://allserv.rug.ac.be/�jdhont/index.htm 

The Artemia Reference Center at the University 

of Ghent, Belgium. 

Large Branchiopod Home Page 

http://mailbox.univie.ac.at/Erich.Eder/UZK/ 

Eric Eder’s site for ‘‘everything you ever wanted 

to know about large branchiopods.’’ 

Leptostraca 

http://www.nhm.org/�peet/ 

A comprehensive site on leptostracans maintained 

by Todd Haney (toddhaney@crustacea.net) 

as part of a PEET project funded by the U.S. National 

Science Foundation. 


(The) Lurker’s Guide to Stomatopods 

http://www.blueboard.com/mantis/welcome.htm 

Alan San Juan’s stomatopod site at Seton Hall, 

described by him as ‘‘an additional information resource 

for those people interested in the study and 

care of stomatopods (mantis shrimps).’’ 

Marine Crustaceans of Southern Australia 

http://www.mov.vic.gov.au/crust/page1a.html 

This excellent guide has been assembled by Gary 

Poore (Museum of Victoria, Melbourne) as a reference 

for the identification of a few (about 100) 

of the numerous species of marine crustaceans 

known to exist in southern Australia. Richly illustrated 

with excellent photographs and accompanied 

by background information on the biology, 

distinguishing characters, habitat, and distribution 

of the species illustrated. 

Monoculus—Copepod Newsletter 

http://www.uni-oldenburg.de/monoculus/ 

The home page of the copepodologist’s newsletter, 

edited by Hans-Uwe Dahms. 

National Center for Biotechnology Information 

Taxonomy Browser 

http://www3.ncbi.nlm.nih.gov/htbin-post/ 

Taxonomy/wgetorg?id�6681&lvl�10 

For locating DNA/RNA sequences of a variety of 

crustaceans. 

National Shellfisheries Association 

http://www.shellfish.org/ 

The home page of this association, with links to 

journals and other activities. 

‘‘Non-Cladoceran’’ Branchiopod Shrimp of Ohio 

http://www-obs.biosci.ohio-state.edu/f-shrimp.htm 

Contains information on anostracans, notostracans, 

and conchostracans of Ohio. Maintained by 

Stephen Weeks, University of Akron, Ohio, USA. 

North East Atlantic Taxa 

http://www.tmbl.gu.se/libdb/taxon/taxa.html 

Contains PDF files of species checklists, including 

crustaceans from this region, compiled by the Tjärnö 

Marine Biological Laboratory, Sweden. 

Orsten and Crustacean Phylogeny 

http://biosys-serv.biologie.uni-ulm.de/sektion/ 

dieter/dieter.html 

Dieter Walossek’s page introducing the ‘‘orsten’’ 

fossils (Upper Cambrian of Sweden), Eucrustacea, 

nonarthropod crustaceans, and more. Includes photographs 

and drawings of the ‘‘orsten’’ arthropods. 

Ostracod Research Group 

http://users.aber.ac.uk/alm/web/ostrweb2.html 

A site maintained by Robin Whatley and Henry 

Lamb; this is a subgroup of the Micropaleontology 

Research Group in the Institute of Geography and 

Earth Sciences at the University of Wales, Aberystwyth. 

Pesce’s Home Page/Groundwater Fauna of Italy 

http://www.univaq.it/�sc�amb/pesce.html 

Contains information about, and links to, 

groundwater and speleofaunal crustaceans of Italy, 

with links to other sites dealing with amphipods, 

mysids, copepods, and more. 

PHOTOVAULT’s Aquatic Crustacean’s Page 

http://www.photovault.com/Link/Animals/Aquatic� 

Crustacia/AARVolume01.html 

A commercial site that contains many photographs 

of various crustaceans. 

SCAMIT Arthropods of Southern California 

http://www.scamit.org/SpeciesList/arthropd.htm 

An unannotated list of the species of soft bottom 

habitats off southern California, maintained by the 

Southern California Association of Marine Invertebrate 

Taxonomists (SCAMIT). 

(A) Stereo-Atlas of Ostracod Shells 

http://www.nhm.ac.uk/hosted�sites/bms/saos.htm 

A site with information on this and other publications 

of the British Micropaleontological Association. 

(The) Subterranean Amphipod Database 

http://www.odu.edu/�jrh100f/amphipod/ 

Maintained by John Holsinger at Old Dominion 

University, Norfolk, Virginia. 

Systematics of Amphipod Crustaceans (order 

Amphipoda) in the families Crangonyctidae and 

Hadziidae 

http://www.odu.edu/�jrh100f/ 

A U.S. National Science Foundation PEET project 

maintained by John Holsinger (and his students) 

at Old Dominion University, Virginia, USA. 

Includes the Subterranean Amphipod Database. 

Tanaidacea Homepage 

http://tidepool.st.usm.edu/tanaids/index.html 

A comprehensive and searchable listing of all 


tanaid taxa and the literature in which they were 

initially described. Maintained by Richard W. 

Heard and Gary Anderson at the University of 

Southern Mississippi, USA. 

Urzeitkrebse—Lebende Fossilien! 

http://mailbox.univie.ac.at/Erich.Eder/UZK/index2. 

html 

Contains information on large branchiopods 

(Anostraca, Conchostraca, and Notostraca) of Austria, 

maintained by Eric Eder. 

(The) University of South Carolina Meiofaunal 

Laboratory of Bruce Coull 

http://inlet.geol.sc.edu/�nick/ 

A meiofauna page, including harpacticoid copepods, 

maintained by Bruce Coull at the University 

of South Carolina. 

World List of Marine, Freshwater and Terrestrial 

Isopod Crustaceans 

http://www.nmnh.si.edu/iz/isopod 

Contains more than 9,900 isopod records, all described 

species of isopods, and a complete bibliography 

in a searchable Access database. Maintained 

by Brian Kensley (kensley.brian@nmnh.si.edu) and 

Marailyn Schotte (schotte.marilyn@nmnh.si.edu) of 

the USNM, Smithsonian Institution. 

(The) World of Copepoda 

http://www.nmnh.si.edu/iz/copepod/ 

Contains bibliographic databases for all the literature 

contained in the Wilson Library on copepods 

and branchiurans. In total, the website contains 

four databases: (1) a bibliography of all 

known copepod and branchiuran literature, (2) a 

taxonomic list of reported Copepoda and Branchiura 

genera and species, (3) copepod and branchiuran 

researchers of the world, and (4) copepod and 

branchiuran type holdings of the U.S. National 

Museum of Natural History. Maintained by Chad 

Walter. 

Zoological Record Taxonomic Hierarchy 

http://www.biosis.org.uk/zrdocs/zoolinfo/grp� 

crus.htm 

The extensive Internet Resource Guide for Zoology 

provided by Biosis and the Zoological Society 

of London. 

Zooplankton Sensory Motor Systems 

http://www.pbrc.hawaii.edu/�lucifer/ 

Contains information on, and links to, research 

and researchers investigating sensory biology and 

motor processes and systems in zooplankton of pe- 

lagic crustacean and crustacean larvae. Maintained 

by Dan Hartline and Petra Lenz. 

INDIVIDUAL WORKERS WITH HOME PAGES 

CONTAINING CRUSTACEAN 

INFORMATION 

Gary Anderson 

http://tidepool.st.usm.edu/gandrsn/gandrsn.html 

A well-designed site with a large number of links 

to other sites of interest to crustacean workers. 

Raymond Bauer 

http://www.ucs.usl.edu/�rt6933/shrimp/ 

Highlights his research interests in marine habitats 

and the biology of caridean and penaeoid 

shrimp, mating behavior and strategies, hermaphroditism 

and sex change, antifouling (grooming) 

behavior, sperm transfer, latitudinal variation in 

breeding patterns, seagrass fauna, coloration and 

camouflage, and student research. 

Geoffrey Boxshall 

http://www.nhm.ac.uk/science/zoology/project1/ 

index.html 

A single page with general information on copepods, 

linked to The Natural History Museum, 

London site. 

Raul Castro R. 

http://members.xoom.com/renrique/copepoda2. 

html 

List of parasitic copepods on Chilean fishes with 

a list of his publications. 

Paul Hebert 

http://www.uogluelph.ca/�phebert/ 

Summarizes past and current research and multimedia 

projects of the lab. 

Wolfgang Janetzky 

http://www.ifas.ufl.edu/�frank/crbrom.htm 

Highlights his interests in Crustacea inhabiting 

bromeliad phytotelmata. 

Gertraud Krapp-Schickel 

http://hydr.umn.edu/g-k/index.html 

Highlights her interests in amphipods, plus photos 

of amphipodologists. 

Colin McLay 

http://www.zool.canterbury.ac.nz/cm.htm 

Highlights his research interests in population 

and marine ecology, reproductive biology, mating 


strategies, and phylogeny, especially of anomurans 

and brachyurans. 

Jeffrey Shields 

http://www.vims.edu/�jeff/ 

The parasitic isopods of Crustacea (Bopyridae, 

Entoniscidae, and Dajidae). 

Wim Vader 

http//:www.imv.uit.no/ommuseet/enheter/zoo/wim/ 

index.html 

A single page highlighting his interests in Crustacea 

and Amphipoda. 

George (Buz) Wilson 

http://www-personal.usyd.edu.au/�buz/home.html 

Research interests emphasizing asellotan and 

phreatoicidean diversity, with links to many other 

isopod and crustacean sites. 

III-C. CRUSTACEAN LIST SERVERS 

ALCA-L 

majordomo@fenix.ciens.ucv.ve 

List server of the Asociation Latinoamericana de 

Carcinologia, currently maintained by Guido Pereira 

(gpereira@strix.ciens.ucv.ve), Instituto de 

Zoologia Tropical, Universidad Central de Venezuela, 

Caracas, Venezuela. 

BRINE-L 

http://ag.ansc.purdue.edu/aquanic/infosrcs/brine-l. 

htm 

A brine shrimp (Anostraca) discussion list, maintained 

by Lamar Jackson and Harold Pritchett at 

Mercer University, Georgia. Part of AquaNIC, the 

Aquaculture Network Information Center. 

COPEPODA 

copepoda@sciencenet.com 

A list server for discussions of wide ranging copepod 

research. 

CRUST-L 

http://www.vims.edu/�jeff/crust-l.html 

An informal forum for those interested in Crustacea, 

including their biology, ecology, systematics, 

taxonomy, physiology, cell biology, culture, etc. 

Managed by Jeff Shields. 

OSTRACON 

The Ostracoda Discussion List, OSTRACON@ 

LISTSERV.UH.EDU 

A list server for discussions of all things ostracode-like. 

The Vernal Pool ListServ 

vernal@sun.simmons.edu 

The Vernal Pool Association maintains a list on 

the EnvironNet server for those interested in vernal 

pool studies, protection, and education. 

III-D. SOME MUSEUMS WITH CRUSTACEAN 

HOLDINGS ON-LINE 

California Academy of Sciences 

http://web.calacademy.org/research/izg/ 

This will take you directly to the CAS Invertebrate 

Zoology and Geology Department. 

Department of Invertebrate Zoology at the United 

States National Museum 

http://www.nmnh.si.edu/departments/invert.html 

A well-written overview of the history and activities 

of the staff of the world’s largest collection of 

Crustacea. 

Illinois Natural History Survey Crustacean 

Biology Information Page 

http://www.inhs.uiuc.edu/cbd/collections/crustacea. 

html 

One of the largest state collections of crustaceans 

in North America, with a searchable database and 

a well-designed page. 

Muséum National d’Histoire Naturelle (Paris) 

http://www.mnhn.fr/ 

Extensive crustacean holdings, but no information 

available on line yet. 


http://www.nhm.org/ 

The largest natural history museum in the western 

United States, this impressive institution is also 

home to the second largest collection of Crustacea 

in this country. There are an estimated 110,000 to 

120,000 lots, containing 3 to 4 million specimens. 

University of California Berkeley Museum of 

Paleontology 

http://www.ucmp.berkeley.edu 

An interesting page that includes mostly paleontological 

information on arthropods. 

Zoological Museum, University of Copenhagen 

http://www.aki.ku.dk/zmuc/zmuc.htm 

A beautiful home page for one of Europe’s oldest 

and most respected natural history museums. The 

Crustacea collection is extensive and well-curated. 


Addendum 

As might be expected in any attempt to be current in a rapidly changing field, several publications 

or presentations that bear on high-level relationships of the Crustacea have come to light during the 

final months while we prepared this volume for the printer. In particular, the following presentations 

dealing with higher crustacean systematics were selected from among the published abstracts of the 

Fifth International Crustacean Congress in Melbourne, Australia (July 9–13, 2001) (Fifth International 

Crustacean Congress—Program and Abstracts, and List of Participants, 2001): Developmental 

data in crustacean systematics (Koenemann and Schram); Peracarida (Wilson, Watling, Richter, Jarman, 

Spears et al., Wilson and Ahyong, Keable and Wilson, Poore and Brandt, Myers and Lowry); 

malacostracan affinities with insects (K. Wilson); Decapoda (Ahyong and Schram, Porter et al., Brösing 

and Scholtz, Crandall et al., Richter, Pérez-Losada et al., Boyce et al., Wetzer et al., Ngoc-Ho); 

Remipedia (Spears and Yager); Leptostraca (Walker-Smith and Poore); Phosphatocopina (Maas and 

Walossek); Rhizocephala (Glenner and Spears).

An Updated Classification of the Recent Crustacea

Create successful ePaper yourself

Delete template?

Save as template?