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An Updated Classification of the Recent Crustacea

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eventually led us to keep dromiids with <strong>the</strong> o<strong>the</strong>r<br />

‘‘primitive’’ brachyurans in our section Dromiacea,<br />

knowing that by so doing we are continuing to displease<br />

students <strong>of</strong> crab phylogeny who rely mostly<br />

on larval characters and that <strong>the</strong> current arrangement<br />

<strong>of</strong> primitive crabs is not completely in keeping<br />

with <strong>the</strong> molecular evidence in <strong>the</strong> Spears et al.<br />

(1992) study. A detailed discussion <strong>of</strong> <strong>the</strong> situation<br />

within <strong>the</strong> Dromiacea can be found in <strong>the</strong> review<br />

<strong>of</strong> <strong>the</strong> Dynomenidae by McLay (1999).<br />

Superfamily Dromioidea<br />

The families Dromiidae and Dynomenidae are still<br />

listed as valid families, although based on molecular<br />

data (Spears et al., 1992) and sperm morphology<br />

(Jamieson, 1994; Jamieson et al., 1995; Guinot<br />

et al., 1998), <strong>the</strong>ir monophyletic status has been<br />

questioned (but see McLay, 1991, 1999; Števčić,<br />

1995). Earlier classifications, some <strong>of</strong> which have<br />

included <strong>the</strong> Homolidae among <strong>the</strong> dromiacean<br />

families, are reviewed by Števčić (1995), Guinot<br />

and Richer de Forges (1995), and McLay (1999).<br />

Guinot et al. (1998) argue that <strong>the</strong> Dromioidea (referred<br />

to as Dromiacea in that paper, a lapsus calami,<br />

Guinot, pers. comm.), containing <strong>the</strong> three<br />

families Dromiidae, Dynomenidae, and Homolodromiidae,<br />

is a valid monophyletic superfamily, although<br />

<strong>the</strong>y note <strong>the</strong> differences separating <strong>the</strong><br />

homolodromiids. We have maintained <strong>the</strong> separate<br />

status <strong>of</strong> <strong>the</strong> homolodromiids (i.e., placing <strong>the</strong>m in<br />

<strong>the</strong>ir own superfamily Homolodromioidea; see below)<br />

in light <strong>of</strong> <strong>the</strong> many morphological features <strong>of</strong><br />

adults that seem to separate <strong>the</strong>m from <strong>the</strong> dromiids<br />

and dynomenids. In doing so, we follow Guinot<br />

(1995), even though Guinot and Bouchard (1998)<br />

have reverted to treating all three <strong>of</strong> <strong>the</strong>se families<br />

in one superfamily (<strong>the</strong>ir Dromiacea). The families<br />

were reviewed recently by McLay (1991, Dromiidae;<br />

1999, Dynomenidae) with special regard to<br />

<strong>the</strong>ir Indo-Pacific members.<br />

Superfamily Homolodromioidea<br />

Separate superfamily status for <strong>the</strong> Homolodromiidae<br />

appears warranted on <strong>the</strong> basis <strong>of</strong> larval and<br />

adult morphology (see Martin, 1991; Guinot,<br />

1995). Števčić (1998) considers <strong>the</strong> homolodromiids<br />

<strong>the</strong> most primitive extant family <strong>of</strong> brachyuran<br />

crabs. The date <strong>of</strong> Alcock’s establishment <strong>of</strong> <strong>the</strong><br />

Homolodromiidae has been changed from 1899 to<br />

1900 following <strong>the</strong> revision by Guinot (1995).<br />

Superfamily Homoloidea<br />

The alliance <strong>of</strong> homolids with dromiids has been<br />

supported by ultrastructural characters <strong>of</strong> <strong>the</strong><br />

sperm (Guinot et al., 1994; see also <strong>the</strong> extensive<br />

review by Guinot and Richer de Forges, 1995). The<br />

family Poupiniidae was added by Guinot (1991).<br />

SECTION EUBRACHYURA, SUBSECTION<br />

RANINOIDA<br />

Superfamily Raninoidea<br />

Within <strong>the</strong> Raninoidea, <strong>the</strong> subfamily Symethinae<br />

(monogeneric; Symethis Goeke) was elevated to<br />

family level by Tucker (1998), as had been suggested<br />

earlier by Guinot (1993). However, Tucker<br />

did not agree with <strong>the</strong> removal <strong>of</strong> <strong>the</strong> subfamily<br />

Cyrtorhininae from <strong>the</strong> Raninidae, which had been<br />

suggested as a possibility by Guinot (1993).<br />

Superfamily Cyclodorippoidea<br />

The superfamily Tymoloidea has been removed and<br />

in its place is <strong>the</strong> superfamily Cyclodorippoidea, as<br />

<strong>the</strong> family name Cyclodorippidae Ortmann has seniority<br />

over Tymolidae Alcock, according to Guinot<br />

(pers. comm.) and Tavares (1991, 1993). Tavares<br />

(1998) also established a new family, <strong>the</strong> Phyllotymolinidae,<br />

within <strong>the</strong> Cyclodorippoidea. Guinot<br />

and Bouchard (1998) continue to recognize <strong>the</strong> superfamily<br />

Cyclodorippoidea (as did Tavares, 1991,<br />

1993, 1998), stating that this was done ‘‘for convenience’’<br />

while at <strong>the</strong> same time cautioning against<br />

possible paraphyly in <strong>the</strong> assemblage.<br />

Placement <strong>of</strong> this superfamily with <strong>the</strong> raninoids<br />

in <strong>the</strong> Raninoida is possibly a mistake; molecular<br />

data seem to indicate a placement somewhere between<br />

<strong>the</strong> raninids and <strong>the</strong> higher eubrachyurans<br />

(T. Spears, pers. comm.).<br />

SECTION EUBRACHYURA, SUBSECTION<br />

HETEROTREMATA<br />

Superfamily Dorippoidea<br />

The family Orithyiidae Dana has been transferred<br />

to this superfamily based on <strong>the</strong> suggestion <strong>of</strong> Bellwood<br />

(1996, 1998; see below).<br />

Superfamilies Calappoidea and Leucosioidea<br />

The monophyly <strong>of</strong> <strong>the</strong> family Calappidae and its<br />

constituent subfamilies has been questioned recently.<br />

Bellwood (1996, 1998) has recommended that<br />

only <strong>the</strong> families Calappidae and Hepatidae be retained<br />

in <strong>the</strong> superfamily Calappoidea, with <strong>the</strong><br />

Matutidae joining <strong>the</strong> leucosiids in <strong>the</strong> Leucosioidea<br />

and with <strong>the</strong> Orithyiidae transferred to <strong>the</strong> dorippoids.<br />

To some extent, <strong>the</strong>se changes reflect earlier<br />

suggestions based on larval (Rice, 1980) and<br />

adult (Guinot, 1978; Seridji, 1993) morphology,<br />

and <strong>the</strong>re is at least some fossil support for this<br />

arrangement as well (Feldmann and Hopkins,<br />

1999; Schweitzer and Feldmann, 2000). Števčić<br />

(1983) had earlier suggested recognition <strong>of</strong> <strong>the</strong> Matutidae<br />

and Orithyidae and <strong>the</strong>ir separation from<br />

o<strong>the</strong>r Calappidae as well. We have followed Bellwood’s<br />

(1996) recommendations while at <strong>the</strong> same<br />

time not agreeing with her that <strong>the</strong> Oxystomata be<br />

retained. Bellwood’s rearrangement <strong>of</strong> <strong>the</strong> calappids<br />

is not supported by recent molecular data (S.<br />

Boyce, unpublished).<br />

Contributions in Science, Number 39 Rationale 51

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