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An Updated Classification of the Recent Crustacea

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Superfamily Majoidea<br />

Hendrickx (1995, and pers. comm.) brought our<br />

attention to <strong>the</strong> elevation <strong>of</strong> several majid subfamilies<br />

to familial rank, such as <strong>the</strong> elevation <strong>of</strong> some<br />

inachoid groups by Drach and Guinot (1983), who<br />

recognized as families <strong>the</strong> Inachidae and Inachoididae.<br />

We have followed Hendrickx’s recognition <strong>of</strong><br />

former majid subfamilies as families. To some degree,<br />

our treatment (and Hendrickx’s) <strong>of</strong> <strong>the</strong> majoid<br />

families follows <strong>the</strong> seven subfamilies proposed by<br />

Griffin and Tranter (1986) in <strong>the</strong>ir major revision<br />

<strong>of</strong> <strong>the</strong> Majidae <strong>of</strong> <strong>the</strong> Indo-West Pacific. Additional<br />

subfamilies have been proposed by o<strong>the</strong>r workers,<br />

including Števčić (1994), who disagreed with some<br />

<strong>of</strong> <strong>the</strong> subdivisions proposed by Griffin and Tranter<br />

(1986). Diversity <strong>of</strong> <strong>the</strong> former family Majidae is<br />

incredibly high, and recognition or treatment <strong>of</strong> <strong>the</strong><br />

majoids as a superfamily has been noted or suggested<br />

by many earlier workers (e.g., Guinot, 1978;<br />

Drach and Guinot, 1983; Števčić, 1994; Clark and<br />

Webber, 1991, among o<strong>the</strong>rs). M. Wicksten (pers.<br />

comm.) suggests that, if we elevate some <strong>of</strong> <strong>the</strong> former<br />

majid subfamilies to <strong>the</strong> family level, <strong>the</strong>n we<br />

should recognize also <strong>the</strong> family Oregoniidae<br />

Garth, 1958, and possibly also <strong>the</strong> Macrocheiridae<br />

Balss, 1929, ‘‘for consistency.’’ Indeed, Clark and<br />

Webber (1991) proposed recognition <strong>of</strong> both <strong>of</strong><br />

<strong>the</strong>se families based on a reevaluation <strong>of</strong> <strong>the</strong> larval<br />

features <strong>of</strong> Macrocheira and suggested that extant<br />

majoids be partitioned among four families: Oregoniidae,<br />

Macrocheiridae, Majidae, and Inachidae.<br />

Larval morphology indicates <strong>the</strong> distinct nature,<br />

and presumed monophyly, <strong>of</strong> <strong>the</strong>se groups as well<br />

(Pohle and Marques, 2000). We have not taken that<br />

step here, feeling that knowledge <strong>of</strong> larval majoids<br />

is still ra<strong>the</strong>r incomplete, and we recognize here<br />

only <strong>the</strong> families Epialtidae, Inachidae, Inachoididae,<br />

Majidae, Mithracidae, Pisidae, and Tychidae.<br />

Concerning phylogeny among <strong>the</strong> higher (heterotrematous)<br />

crabs, Rice (1983:326) depicts <strong>the</strong> Majidae<br />

(our Majoidea) as basal to <strong>the</strong> primitive xanthid<br />

stock, which in turn gives rise to all o<strong>the</strong>r crab<br />

families and superfamilies. A recent study based on<br />

larval characters (Pohle and Marques, 2000) suggests<br />

that, within <strong>the</strong> Majoidea, <strong>the</strong> Oregoniinae<br />

clade is most basal among those majoid families (or<br />

subfamilies) for which larval morphology is<br />

known.<br />

Superfamily Hymenosomatoidea<br />

According to Guinot and Richer de Forges (1997),<br />

members <strong>of</strong> <strong>the</strong> family Hymenosomatidae (sole<br />

member <strong>of</strong> this superfamily) are thought to be<br />

‘‘highly advanced Heterotremata and not Thoracotremata’’<br />

(Guinot and Richer de Forges, 1997:<br />

454, English abstract). In addition, Guinot and<br />

Richer de Forges (1997) revive <strong>the</strong> idea that <strong>the</strong><br />

closest relatives <strong>of</strong> <strong>the</strong> hymenosomatids may lie<br />

among <strong>the</strong> majoid family Inachoididae. The unusual<br />

sperm morphology <strong>of</strong> one species <strong>of</strong> <strong>the</strong> family,<br />

as reported by Richer de Forges et al. (1997),<br />

would seem to exclude <strong>the</strong> Hymenosomatidae from<br />

<strong>the</strong> Thoracotremata, and even casts doubts as to<br />

<strong>the</strong> family’s inclusion in <strong>the</strong> Heterotremata.<br />

Superfamily Par<strong>the</strong>nopoidea<br />

The superfamily Mimilambroidea and its sole family<br />

Mimilambridae, both originally erected by Williams<br />

(1979) to contain Mimilambrus, have been<br />

removed following <strong>the</strong> suggestion <strong>of</strong> Ng and Rodriguez<br />

(1986) that Mimilambrus can be accommodated<br />

within <strong>the</strong> Par<strong>the</strong>nopidae. Hendrickx (1995)<br />

again alerted us to <strong>the</strong> fact that several former subfamilies<br />

<strong>of</strong> crabs (in this case, former par<strong>the</strong>nopid<br />

subfamilies) had been suggested to be deserving <strong>of</strong>,<br />

or had actually been elevated to, family rank as<br />

long ago as 1978 (Guinot, 1978). Although several<br />

authors (e.g., Hendrickx, 1999) have attributed <strong>the</strong><br />

family name Daldorfiidae to M. J. Rathbun, we<br />

have found no indication that <strong>the</strong> taxon was recognized<br />

by her. Ng and Rodriguez (1986) recognized<br />

<strong>the</strong> suggested par<strong>the</strong>nopoid groupings <strong>of</strong><br />

Guinot as valid families and first used <strong>the</strong> names<br />

Daldorfiidae [as Daldorfidae] and Dairidae, and we<br />

have attributed <strong>the</strong>se families to <strong>the</strong>m. We have followed<br />

Guinot (1978) and Ng and Rodriguez (1986)<br />

and recognize <strong>the</strong> families Aethridae, Dairidae,<br />

Daldorfiidae, and Par<strong>the</strong>nopidae within a superfamily<br />

Par<strong>the</strong>nopoidea, although Hendrickx (1995)<br />

stopped short <strong>of</strong> treating all <strong>of</strong> <strong>the</strong>se as valid families.<br />

Superfamily Retroplumoidea<br />

The family Retroplumidae was given its own superfamily<br />

by Saint Laurent (1989), and its placement<br />

among <strong>the</strong> Heterotremata is based on Saint<br />

Laurent (1989) and Guinot (pers. comm.)<br />

Superfamily Cancroidea<br />

The family Cheiragonidae Ortmann, 1893, containing<br />

<strong>the</strong> genera Telmessus and Erimacrus (formerly<br />

treated by most workers as atelecyclids), was<br />

resurrected and redescribed by Števčić (1988), and<br />

this has been followed by Peter Ng (1998, and pers.<br />

comm., 1997; see also Schweitzer and Salva, 2000),<br />

and so we have included it here as well.<br />

Superfamily Portunoidea<br />

The freshwater family Trichodactylidae has now<br />

been placed in this superfamily, where it joins <strong>the</strong><br />

portunids and geryonids, based primarily on a recent<br />

morphological analysis (Sternberg et al., 1999;<br />

see also below under Potamoidea). Fundamental<br />

differences between trichodactylids and o<strong>the</strong>r freshwater<br />

crabs were recognized by several earlier<br />

workers. Rodriguez (1982, 1986, 1992), Magalhães<br />

and Türkay (1996a–c), Sternberg (1997),<br />

Sternberg et al. (1999), Christoph Schubart (pers.<br />

comm.), and Spears et al. (2000) all acknowledge<br />

<strong>the</strong> unique position <strong>of</strong> <strong>the</strong> Trichodactylidae and all<br />

consider <strong>the</strong> family monophyletic. The hypo<strong>the</strong>sis<br />

52 Contributions in Science, Number 39 Rationale

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