A classification of living and fossil genera of - Raffles Museum of ...
A classification of living and fossil genera of - Raffles Museum of ...
A classification of living and fossil genera of - Raffles Museum of ...
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HOW MANY SPECIES OF DECAPODA<br />
ARE THERE?<br />
Chace (1951) was the last authority to tabulate all <strong>genera</strong><br />
<strong>and</strong> species <strong>of</strong> then-known Recent decapod crustaceans,<br />
based on a comprehensive file card system that he <strong>and</strong><br />
colleagues maintained at the United States National<br />
<strong>Museum</strong> <strong>of</strong> Natural History. Chace estimated the known<br />
number at 8,321 species, distributed among 1,000 <strong>genera</strong>,<br />
with Brachyura accounting for 4,428 species (in 635<br />
<strong>genera</strong>). According to Ng et al. (2008), this latter figure<br />
compares quite favorably with their estimate <strong>of</strong> 4,120<br />
known at that time (raised to 6,793 by 2008), <strong>and</strong> thus we<br />
can place some confidence in Chace’s global decapod<br />
species estimate <strong>of</strong> 8,321 known in 1951. More recent,<br />
somewhat vague, published estimates put the number <strong>of</strong><br />
species somewhere upwards <strong>of</strong> 10,000 species, with only<br />
Martin & Davis (2006) providing an accurate estimate <strong>of</strong><br />
7,905 species (but based on a database estimated to contain<br />
only about half <strong>of</strong> all described crustacean species). Based<br />
on our current effort, we put the total number <strong>of</strong> extant<br />
species <strong>of</strong> Decapoda at 14,756 (in 2,725 <strong>genera</strong>), with<br />
Brachyura accounting for 6,835 species, i.e. 46% (Table 1).<br />
This implies that in the last 50 years, the number <strong>of</strong><br />
described species has nearly doubled! However, we are <strong>of</strong><br />
course still a long way from knowing the true global<br />
diversity <strong>of</strong> Decapoda. The number <strong>of</strong> known <strong>fossil</strong> species<br />
currently st<strong>and</strong>s at 3,300. Here too, discoveries will<br />
continue for some time to come, as new localities are<br />
explored, more revisions are completed, <strong>and</strong> museum<br />
collections are more thoroughly studied.<br />
THE HIGHER CLASSIFICATION OF DECAPODA<br />
The higher <strong>classification</strong> <strong>of</strong> the Decapoda is deeply rooted<br />
in the early works <strong>of</strong> such luminaries as H. Milne Edwards,<br />
Dana, Latreille <strong>and</strong> contemporaries, to which can be added<br />
the <strong>of</strong>ten divergent opinions <strong>of</strong> Borradaile, Bate <strong>and</strong> others.<br />
In the last few decades, a more stable <strong>classification</strong> has<br />
emerged due to the more rigorous application <strong>of</strong> cladistic<br />
techniques to both morphological <strong>and</strong> molecular data, as<br />
well as to the advent <strong>of</strong> more sophisticated morphological<br />
<strong>and</strong> molecular studies. Although the last word on the<br />
subject has not been given (especially within the<br />
Brachyura), we now appear to have broad consensus on the<br />
validity <strong>of</strong> two suborders: Dendrobranchiata <strong>and</strong><br />
Pleocyemata, amply supported by genetic <strong>and</strong><br />
morphological studies. The Pleocyemata are herein further<br />
divided into 10 infraorders, most <strong>of</strong> which have been<br />
recognized for some time <strong>and</strong> appear uncontroversial<br />
(Caridea, Stenopodidea, Astacidea, Achelata, Anomura,<br />
Brachyura). In addition to those, we follow Dixon et al.<br />
(2003), Ahyong & O’Meally (2004), <strong>and</strong> Bracken et al.<br />
(2009) in affording infraordinal status to the glypheid<br />
RAFFLES BULLETIN OF ZOOLOGY 2009<br />
5<br />
lobsters (as infraorder Glypheidea). Following Scholtz &<br />
Richter (1995) <strong>and</strong> Ahyong (2009), we also recognize the<br />
infraorder Polychelida. We follow Robles et al. (2009) in<br />
the recognition <strong>of</strong> two infraorders (Axiidea, Gebiidea) for<br />
the thalassinidean families. Within the Brachyura, <strong>and</strong><br />
departing from Ng et al. (2008), we do not recognize the<br />
section Podotremata, <strong>and</strong> instead follow Ahyong et al.<br />
(2007) with three sections: Dromiacea, Raninoida,<br />
Cyclodrippoida (see also Ahyong, Naruse et al., 2009).<br />
Further notes on the <strong>classification</strong> employed within the<br />
various groups are presented below.<br />
We stress that our arrangement <strong>of</strong> taxa, both at higher<br />
levels (families within superfamilies, sections, etc.) <strong>and</strong> at<br />
lower levels (<strong>genera</strong> within families), is not agreed upon<br />
by all workers.<br />
Dendrobranchiata, Stenopodidea, Caridea. For the<br />
Dendrobranchiata, apart from some additional <strong>genera</strong>, little<br />
has changed since the comprehensive treatment in Pérez<br />
Farfante & Kensley (1997), which in turn formed the basis<br />
<strong>of</strong> the Martin & Davis (2001) <strong>classification</strong>. Recent <strong>and</strong><br />
ongoing molecular studies (e.g. Chan et al., 2008; Ma et<br />
al., 2009) may require changes in the near future. The<br />
stenopodideans, a relatively small group, also have<br />
changed little since the Martin & Davis (2001) list,<br />
although one family <strong>and</strong> several new <strong>genera</strong> have been<br />
added. For the Caridea, an enormous <strong>and</strong> diverse<br />
assemblage, our work is based on ongoing updates to the<br />
Holthuis (1993) catalogue by S. De Grave <strong>and</strong> C. Fransen.<br />
One difference pertains to the subfamilies in Atyidae,<br />
which we do not recognize herein. Another difference<br />
concerns the family status <strong>of</strong> several palaemonoid groups,<br />
where we follow Bruce (1993) in recognizing eight<br />
families. The order <strong>of</strong> superfamilies followed here is not a<br />
reflection <strong>of</strong> phylogenetic order, <strong>and</strong> merely follows Chace<br />
(1992) <strong>and</strong> Holthuis (1993). We acknowledge the recent<br />
study by Bracken et al. (2009), which questioned the<br />
validity <strong>of</strong> the superfamily arrangement within Caridea as<br />
well as the monophyly <strong>of</strong> several families. Christ<strong>of</strong>fersen<br />
(1986, 1987, 1988a, 1988b, 1989, 1990) proposed an<br />
alternative arrangement <strong>of</strong> caridean systematics (see<br />
Holthuis, 1993, for a discussion). Although some <strong>of</strong> his<br />
suggestions have been incorporated over the years (e.g. see<br />
Martin & Davis, 2001), many others were rejected.<br />
However, recent, as yet unpublished, work has forced<br />
caridean workers to look more closely at some <strong>of</strong><br />
Christ<strong>of</strong>fersen’s suggestions. Evidence that an extensive<br />
rearrangement <strong>of</strong> caridean superfamilies <strong>and</strong> families is<br />
needed is forthcoming in ongoing morphological <strong>and</strong><br />
molecular studies, but for now we adhere to the more<br />
traditional <strong>classification</strong>.<br />
Astacidea. The evolutionary relationships <strong>and</strong> taxonomy<br />
<strong>of</strong> groups within the infraorder Astacidea are far from<br />
settled <strong>and</strong> stable. Cladistic methods applied to<br />
morphology (since the mid-1990s) <strong>and</strong> to molecular data<br />
(since 2004) hold promise, but results <strong>of</strong>ten disagree even<br />
at high taxonomic levels. For example, the extinct family