An Updated Classification of the Recent Crustacea

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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 26 Contributions in Science, Number 39 Rationale
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 Contributions in Science, Number 39 Rationale 27
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 and 8: ACKNOWLEDGMENTS We sincerely thank
Page 9 and 10: An Updated Classification of the Re
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: parasites of early fish-like verteb
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,
Page 65 and 66: CONCLUDING REMARKS We have thorough
Page 67 and 68: 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
Page 77 and 78: Parascelidae Bate, 1862 Platyscelid
Page 79 and 80: Superfamily Oniscoidea Latreille, 1
Page 81 and 82: Family Alpheidae Rafinesque, 1815 B
Page 83 and 84: Trichodactylidae Milne Edwards, 185
Page 85 and 86:
H.-E. Gruner and L. B. Holthuis, 1-
Page 87 and 88:
zil, ed. P. S. Young, 635-644. Rio
Page 89 and 90:
mated from nuclear and mitochondria
Page 91 and 92:
tacés Péracarides. Memoires de l
Page 93 and 94:
netic reconstruction from 12S rDNA
Page 95 and 96:
pepods and an analysis of the phylo
Page 97 and 98:
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
Page 125 and 126:
(The) Stomatopod Newsletter Editors
Page 127 and 128:
links to crustacean societies, cons
Page 129 and 130:
(The) Lurker’s Guide to Stomatopo
Page 131 and 132:
strategies, and phylogeny, especial
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An Updated Classification of the Recent Crustacea

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