24.10.2014 Views

An Updated Classification of the Recent Crustacea

An Updated Classification of the Recent Crustacea

An Updated Classification of the Recent Crustacea

SHOW MORE
SHOW LESS

Create successful ePaper yourself

Turn your PDF publications into a flip-book with our unique Google optimized e-Paper software.

Brusca and Brusca, 1990; Mayrat and Saint Laurent,<br />

1996; Camp, 1998 (in Camp et al., 1998);<br />

Young, 1998). But as with nearly all o<strong>the</strong>r crustacean<br />

assemblages, this grouping has its opponents<br />

as well. Most <strong>of</strong> <strong>the</strong> disagreement concerns whe<strong>the</strong>r<br />

<strong>the</strong> mysidaceans (i.e., ei<strong>the</strong>r mysids, lophogastrids,<br />

or both) should be placed here (see earlier discussions<br />

on mysids and lophogastrids) and what <strong>the</strong><br />

relationships are among <strong>the</strong> three currently recognized<br />

orders Euphausiacea, Amphionidacea, and<br />

Decapoda (see Jarman et al., 2000; Richter and<br />

Scholtz, in press). Eucarid relationships have been<br />

analyzed by Schram (1984) and by Christ<strong>of</strong>fersen<br />

(1988). Our classification is consistent with both <strong>of</strong><br />

<strong>the</strong>se analyses at higher levels but differs in <strong>the</strong> constituent<br />

suborders.<br />

ORDER EUPHAUSIACEA<br />

The Euphausiacea still contains only <strong>the</strong> two families<br />

Ben<strong>the</strong>uphausiidae (monotypic) and Euphausiidae<br />

(all o<strong>the</strong>r species). The treatment by Baker et<br />

al. (1990) follows this arrangement as well. A recent<br />

analysis <strong>of</strong> 28S rDNA sequence data by Jarman<br />

et al. (2000) suggests that euphausiaceans may<br />

be more closely related to <strong>the</strong> Mysida than to <strong>the</strong><br />

Decapoda.<br />

ORDER AMPHIONIDACEA<br />

This order remains mon<strong>of</strong>amilial and monogeneric<br />

(Amphionides).<br />

ORDER DECAPODA<br />

The decapods have been <strong>the</strong> subject <strong>of</strong> more published<br />

papers than have all o<strong>the</strong>r crustacean groups<br />

combined. This popularity stems in part from <strong>the</strong><br />

economic importance <strong>of</strong> many groups (especially<br />

penaeoid shrimps, palinurid and nephropid lobsters,<br />

and portunid and xanthoid crabs) but also in<br />

part because <strong>of</strong> <strong>the</strong>ir marvelous diversity. The convenient<br />

size <strong>of</strong> most decapods predisposed <strong>the</strong>m to<br />

become subjects <strong>of</strong> some <strong>of</strong> <strong>the</strong> earliest papers using<br />

biochemical and molecular data to resolve crustacean<br />

relationships. Yet we are as far from reaching<br />

a consensus on <strong>the</strong> relationships among <strong>the</strong> decapods<br />

as we are for <strong>the</strong> more obscure groups, and<br />

opinions and datasets remain sharply divided. In<br />

<strong>the</strong> treatment that follows, we have tried to address<br />

<strong>the</strong> many changes and arrangements that have been<br />

suggested since 1982 under <strong>the</strong> taxonomic heading<br />

for each major group <strong>of</strong> decapods. However, we are<br />

certain to have missed several important papers,<br />

and we hasten to remind <strong>the</strong> reader that <strong>the</strong> literature<br />

on this topic is vast. In general, we have settled<br />

on a fairly conservative classification <strong>of</strong> <strong>the</strong><br />

decapods, knowing that, as with all o<strong>the</strong>r crustacean<br />

taxa, this group is destined for revision. Some<br />

<strong>of</strong> <strong>the</strong> many reviews <strong>of</strong> decapod classification that<br />

have appeared since <strong>the</strong> Bowman and Abele (1982)<br />

classification are Felgenhauer and Abele (1983),<br />

Abele and Felgenhauer (1986), Kim and Abele<br />

(1990), Abele (1991), Holthuis (1993a), and<br />

Scholtz and Richter (1995).<br />

The creation <strong>of</strong> two major branches <strong>of</strong> decapods,<br />

Dendrobranchiata and Pleocyemata, by Martin<br />

Burkenroad (1963, 1981) was a ra<strong>the</strong>r bold departure<br />

from previous schemes <strong>of</strong> decapod classification.<br />

According to Fenner Chace (pers. comm.), T.<br />

Bowman more or less accepted Burkenroad’s arguments<br />

without much questioning, and thus <strong>the</strong><br />

use <strong>of</strong> <strong>the</strong> Dendrobranchiata and Pleocyemata in<br />

<strong>the</strong> Bowman and Abele (1982) classification. Chace<br />

(pers. comm.) feels that <strong>the</strong>re is ample evidence for<br />

elevating many <strong>of</strong> <strong>the</strong> major groups <strong>of</strong> <strong>the</strong> Decapoda<br />

as Burkenroad did with <strong>the</strong> penaeoids and<br />

that singling out <strong>the</strong> penaeoid shrimp was to assign<br />

that group an artificial distinction. He is not alone.<br />

Holthuis (1993a; see especially pages 11–13 for a<br />

concise historical overview <strong>of</strong> <strong>the</strong> many attempts to<br />

classify <strong>the</strong> decapods) felt that treating <strong>the</strong> penaeoids<br />

as a separate group (<strong>the</strong> Dendrobranchiata)<br />

equal in rank to <strong>the</strong> combined Natantia Macrura<br />

Reptantia <strong>An</strong>omura Brachyura (<strong>the</strong> Pleocyemata<br />

<strong>of</strong> Burkenroad) was unsatisfactory. Holthuis<br />

(1993a) proposed to revert to <strong>the</strong> older classifications<br />

and treated <strong>the</strong> Natantia and <strong>the</strong> Macrura<br />

Reptantia as ‘‘full suborders <strong>of</strong> equal rank with <strong>the</strong><br />

<strong>An</strong>omura and Brachyura.’’ In his own words (Holthuis,<br />

1993a:6):<br />

I know that this classification will generally be considered<br />

old-fashioned: in several modern handbooks <strong>the</strong><br />

suborder Natantia has been abandoned altoge<strong>the</strong>r; a<br />

small part <strong>of</strong> it, namely <strong>the</strong> Penaeoidea, is elevated to<br />

<strong>the</strong> rank <strong>of</strong> a separate suborder Dendrobranchiata<br />

while <strong>the</strong> rest <strong>of</strong> <strong>the</strong> Natantia plus <strong>the</strong> Macrura Reptantia,<br />

plus <strong>the</strong> <strong>An</strong>omura, plus <strong>the</strong> Brachyura are<br />

placed in a single suborder Pleocyemata. This to me<br />

seems a very artificial and unsatisfactory arrangement,<br />

and I <strong>the</strong>refore still keep to <strong>the</strong> old classification.<br />

This ‘‘old’’ classification to which he refers, probably<br />

because <strong>of</strong> its simplicity and relative lack <strong>of</strong><br />

controversy, is <strong>of</strong>ten encountered in popular and<br />

lay versions <strong>of</strong> crustacean classification. As an example,<br />

<strong>the</strong> publishers <strong>of</strong> <strong>the</strong> BIOSIS and Zoological<br />

Record databases (see URL http://www.york.<br />

biosis.org/zrdocs for <strong>the</strong> BIOSIS/Zoological Record<br />

Taxonomic Hierarchy, Section 10, <strong>Crustacea</strong>) have<br />

‘‘thrown up <strong>the</strong>ir hands in despair’’ (Chace, pers.<br />

comm.) and have reverted to this older and simpler<br />

classification. There, Natantia is treated as a taxon<br />

containing all <strong>of</strong> <strong>the</strong> known shrimp groups (Penaeidea,<br />

Caridea, and Stenopodidea) and <strong>the</strong> Reptantia<br />

is treated as containing <strong>the</strong> anomurans, astacurans,<br />

brachyurans, and palinurans.<br />

Yet <strong>the</strong> distinct nature <strong>of</strong> <strong>the</strong> penaeoids (<strong>the</strong> Dendrobranchiata)<br />

has been supported by additional<br />

morphological (e.g., Schram, 1984, 1986), embryological,<br />

spermatological (e.g., see Jamieson,<br />

1991a), and molecular data. Kim and Abele (1990)<br />

reviewed previous schemes <strong>of</strong> decapod classification<br />

and concluded, based on somewhat limited<br />

data from 18S rRNA, that <strong>the</strong> penaeids were distinct<br />

from o<strong>the</strong>r decapods. This view was support-<br />

Contributions in Science, Number 39 Rationale 43

Hooray! Your file is uploaded and ready to be published.

Saved successfully!

Ooh no, something went wrong!