An Updated Classification of the Recent Crustacea

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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-
Page 1 and 2: An Updated Classification of the Re
Page 3 and 4: Cover Illustration: Lepidurus packa
Page 5 and 6: SCIENTIFIC PUBLICATIONS COMMITTEE N
Page 7: We sincerely thank the many carcino
Page 11 and 12: did not want to attempt it. We are
Page 13 and 14: ored to credit the person or person
Page 15 and 16: and Boxshall (1996, lichomolgoid co
Page 17 and 18: animal phyla.’’ For the Crustac
Page 19 and 20: same persons to comment on the resu
Page 21 and 22: solved (e.g., Regier and Schultz, 1
Page 23 and 24: ancestral crustacean is of course n
Page 25 and 26: fication (see also Negrea et al., 1
Page 27 and 28: Bowman and Abele in not recognizing
Page 29 and 30: (Müller and Walossek, 1985), the O
Page 31 and 32: cerning barnacle evolution (Schram
Page 33 and 34: parasites of early fish-like verteb
Page 35 and 36: Boxshall and Jaume (2000, see also
Page 37 and 38: that reduction in body segmentation
Page 39 and 40: for a rebuttal of Dahl’s criticis
Page 41 and 42: the Amphipoda are deserving of stat
Page 43 and 44: er than Mysida Boas or Mysida Dana)
Page 45 and 46: tained in our classification, altho
Page 47 and 48: and Dalens (1999) in that we includ
Page 49 and 50: subfamilies of the Cryptoniscidae,
Page 51 and 52: Brusca and Brusca, 1990; Mayrat and
Page 53 and 54: employed recently by Pérez Farfant
Page 55 and 56: the former thaumastochelids being t
Page 57 and 58: Cunningham (1999) data, which sugge
Page 59 and 60:
eventually led us to keep dromiids
Page 61 and 62:
that the trichodactylids may repres
Page 63 and 64:
(1996a-c), Rodríguez (1982, 1986,
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We have thoroughly enjoyed the disc
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Order Pygophora Berndt, 1907 Family
Page 69 and 70:
Pseudocyclopiidae Sars, 1902 Pseudo
Page 71 and 72:
Nucellicolidae Lamb, Boxshall, Mill
Page 73 and 74:
Terrestricytheridae Schornikov, 196
Page 75 and 76:
Eusiridae Stebbing, 1888 Exoedicero
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Parascelidae Bate, 1862 Platyscelid
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Superfamily Oniscoidea Latreille, 1
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Family Alpheidae Rafinesque, 1815 B
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Trichodactylidae Milne Edwards, 185
Page 85 and 86:
H.-E. Gruner and L. B. Holthuis, 1-
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zil, ed. P. S. Young, 635-644. Rio
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mated from nuclear and mitochondria
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tacés Péracarides. Memoires de l
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netic reconstruction from 12S rDNA
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pepods and an analysis of the phylo
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among gammaridean families and amph
Page 99 and 100:
es and consequences. In Proceedings
Page 101 and 102:
tostraca) form Unguja Island (Zanzi
Page 103 and 104:
Saint Laurent, M. de. 1979. Vers un
Page 105 and 106:
. 1983a. Evolution of Tanaidacea. I
Page 107 and 108:
Lysianassoidea. Journal of Natural
Page 109 and 110:
Wills, M. A. 1997. A phylogeny of r
Page 111 and 112:
what your sentence says. Second, ju
Page 113 and 114:
logeny and taxonomy. This focus on
Page 115 and 116:
Submitted by Roger L. Kaesler, Pale
Page 117 and 118:
Bousfield, E. L. 1996. A contributi
Page 119 and 120:
knowledge on the phylogeny and the
Page 121 and 122:
inoidea is difficult, because they
Page 123 and 124:
APPENDIX III. OTHER CRUSTACEAN RESO
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(The) Stomatopod Newsletter Editors
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links to crustacean societies, cons
Page 129 and 130:
(The) Lurker’s Guide to Stomatopo
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strategies, and phylogeny, especial
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An Updated Classification of the Recent Crustacea

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