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An Updated Classification of the Recent Crustacea

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subfamilies <strong>of</strong> <strong>the</strong> Cryptoniscidae, for which Bowman<br />

and Abele (1982), followed by Schram (1986),<br />

used <strong>the</strong> name Liriopsidae (see Grygier and Bowman,<br />

1990). Crediting authorship <strong>of</strong> <strong>the</strong> family<br />

Cryptoniscidae (and thus Cryptoniscoidea) to Kossman<br />

ra<strong>the</strong>r than to Gerstaecker is based on <strong>the</strong> correction<br />

published by Grygier and Bowman (1991).<br />

Following Trilles (1999), we also do not recognize<br />

<strong>the</strong> family Microniscidae Müller for <strong>the</strong> genus Microniscus,<br />

although this family is still listed in some<br />

compendia (e.g., by Brasil-Lima, 1998:641, in<br />

Young, 1998). The spelling Cabiropsidae used by<br />

Trilles (1999) and some earlier workers is corrected<br />

to Cabiropidae based on <strong>the</strong> explanation given by<br />

Sassaman (1992).<br />

SUBORDER ONISCIDEA<br />

The relationships <strong>of</strong> <strong>the</strong> terrestrial isopod groups to<br />

one ano<strong>the</strong>r and to marine relatives are still poorly<br />

understood. Although Schmalfuss (1989, in Ferrara,<br />

1989) proposed some relationships among oniscideans<br />

and compared <strong>the</strong> classification <strong>of</strong> oniscideans<br />

presented by Holdich et al. (1984) with a<br />

new one based on his analysis, Schmalfuss’ work<br />

was based on relatively few characters and was criticized<br />

by Brusca (1990). Wägele (pers. comm.) informs<br />

us that <strong>the</strong>re are ‘‘enormous advances that<br />

will be published next year’’ concerning <strong>the</strong> phylogeny<br />

<strong>of</strong> <strong>the</strong> Oniscidea and that several groups<br />

presented here are not monophyletic; fur<strong>the</strong>r, he informs<br />

us that <strong>the</strong> ‘‘section’’ Diplochaeta is currently<br />

being revised. Until <strong>the</strong>se advances become known<br />

to us, we are unsure as to what relationships our<br />

classification should suggest. Holdich et al. (1984)<br />

used two infraorders (<strong>the</strong> Tylidae were placed in a<br />

separate infraorder, Tylomorpha), and within <strong>the</strong><br />

infraorder Ligiamorpha <strong>the</strong>y recognized three sections.<br />

Schmalfuss (1989) did not employ <strong>the</strong> infraorder<br />

level and instead divided all oniscideans<br />

among four major sections. More recent arrangements<br />

<strong>of</strong> <strong>the</strong> oniscidean families have been proposed<br />

by Erhard (1995) and Tabacaru and Danielopol<br />

(1996a, b; see also Roman and Dalens, 1999,<br />

who followed mostly Schmalfuss, 1989, and also<br />

Mattern and Schlegel, 2001). Many workers (e.g.,<br />

Souza-Kury, 1998, in Young, 1998) list <strong>the</strong> oniscidean<br />

families alphabetically.<br />

We have maintained <strong>the</strong> two-infraorder system<br />

and have not recognized <strong>the</strong> new section Microchaeta<br />

proposed by Schmalfuss. The four families<br />

Helelidae, Irmaosidae, Pseudarmadillidae, and<br />

Scleropactidae have been removed from any infraorder<br />

or superfamily, as <strong>the</strong>ir status is indeterminate<br />

(R. Brusca, pers. comm.). For <strong>the</strong> currently<br />

accepted family names (as well as authors and<br />

dates, which were not included by Schmalfuss), we<br />

have had to rely primarily on <strong>the</strong> alphabetical list<br />

<strong>of</strong> oniscidean families maintained on <strong>the</strong> Smithsonian’s<br />

server (Kensley et al., 1998; URL http://<br />

www/nmnh.si.edu/iz/isopod), which is based on<br />

Schmalfuss’ families (<strong>the</strong> terrestrial isopod list is<br />

also accessible via <strong>the</strong> Kensley et al. list <strong>of</strong> marine<br />

isopods, URL gopher://nmnhgoph.si.edu:70/11/.<br />

invertebrate/.crustaceans). Users <strong>of</strong> <strong>the</strong> terrestrial<br />

isopod list are strongly cautioned by <strong>the</strong> authors<br />

(Kensley et al., 1998):<br />

This list is thus intended as a rough guide to <strong>the</strong> astounding<br />

array <strong>of</strong> names and taxa in <strong>the</strong> Oniscidea.<br />

Synonymy will be rampant in <strong>the</strong> list. We have tried to<br />

use <strong>the</strong> most current interpretations <strong>of</strong> some genera and<br />

families. Never<strong>the</strong>less, we realise that in no way do we<br />

even begin to resolve <strong>the</strong> taxonomic confusion that<br />

reigns in this group. There is uncertainty regarding <strong>the</strong><br />

familial placement <strong>of</strong> some genera, and <strong>the</strong>re will certainly<br />

be repetition <strong>of</strong> <strong>the</strong> same specific name under<br />

different genera. There are omissions from <strong>the</strong> list, ei<strong>the</strong>r<br />

<strong>of</strong> names <strong>of</strong> taxa that we’ve completely missed, or<br />

<strong>of</strong> authors and dates <strong>of</strong> publication and/or <strong>of</strong> localities<br />

that we have been unable to find.<br />

We are aware <strong>of</strong> only two newly described oniscidean<br />

families since 1982: Ferrara and Taiti<br />

(1983) described <strong>the</strong> family Irmaosidae, and<br />

Schultz (1995) described <strong>the</strong> Dubioniscidae (see<br />

Souza-Kury, in Young, 1998:656). Establishment <strong>of</strong><br />

<strong>the</strong> family Platyarthridae is credited to Verhoeff<br />

(ra<strong>the</strong>r than to Vandel) by Ferrara and Taiti (1989),<br />

who also note that <strong>the</strong> families Bathytropidae and<br />

<strong>the</strong> Platyarthridae might coincide. G. Poore (pers.<br />

comm.) notes that <strong>the</strong> Styloniscidae Vandel, 1952,<br />

is a replacement name for <strong>the</strong> Patagoniscidae Verhoeff,<br />

1939, and is conserved under ICZN article<br />

40; he <strong>the</strong>refore recommends that <strong>the</strong> earlier date<br />

appear in paren<strong>the</strong>ses, as Styloniscidae Vandel,<br />

1952 (1939). Characters that define <strong>the</strong> various<br />

groupings <strong>of</strong> <strong>the</strong> oniscideans are given by Roman<br />

and Dalens (1999), although workers should note<br />

that <strong>the</strong> characters and groupings based on <strong>the</strong>m<br />

are, in some cases, not universally accepted. A recent<br />

molecular analysis (Mattern and Schlegel,<br />

2001) based on ssu rDNA suggests that Crinochaeta<br />

and Synochaeta are monophyletic, and that <strong>the</strong>se<br />

groups toge<strong>the</strong>r are <strong>the</strong> sister taxon to <strong>the</strong> Diplochaeta.<br />

ORDER TANAIDACEA<br />

Many <strong>of</strong> <strong>the</strong> major taxonomic changes suggested<br />

by <strong>the</strong> late J. Sieg were made prior to 1982 and<br />

were <strong>the</strong>refore incorporated into <strong>the</strong> Bowman and<br />

Abele classification. Subsequent to 1982, <strong>the</strong>re were<br />

also some large-scale rearrangements suggested by<br />

Sieg (1983a, b, 1984, 1986a, b), but <strong>the</strong>re has been<br />

almost no work done at higher levels <strong>of</strong> tanaid systematics<br />

since that time. Unfortunately, it now appears<br />

that many <strong>of</strong> <strong>the</strong> characters established or<br />

used by Sieg do not hold up well under scrutiny<br />

(see Larsen and Wilson, 1998), and it is not clear<br />

how many <strong>of</strong> Sieg’s characters or numerous classificatory<br />

assignments will survive. Kim Larsen (pers.<br />

comm.) is actively studying <strong>the</strong> group and has kindly<br />

updated us, as far as is possible pending a thorough<br />

revision <strong>of</strong> <strong>the</strong> group. Additionally, he has<br />

provided us with many suggested changes. <strong>An</strong> excellent<br />

and comprehensive web site maintained by<br />

Contributions in Science, Number 39 Rationale 41

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