An Updated Classification of the Recent Crustacea
An Updated Classification of the Recent Crustacea
An Updated Classification of the Recent Crustacea
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