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An Updated Classification of the Recent Crustacea

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crabs and is now restricted to <strong>the</strong> families Grapsidae,<br />

Gecarcinidae, Plagusiidae, Searmidae, and Varunidae<br />

should not influence <strong>the</strong> nomenclature.<br />

Additional References<br />

Schubart, C. D., J. A. Cuesta, R. Diesel, and D. L. Felder.<br />

2000b. Molecular phylogeny, taxonomy, and evolution<br />

<strong>of</strong> non-marine lineages within <strong>the</strong> American<br />

Grapsoidea (<strong>Crustacea</strong>: Brachyura). Molecular Phylogenetics<br />

and Evolution (in press).<br />

Schubart, C. D., J. E. Neigel, and D. L. Felder. 2000a. The<br />

use <strong>of</strong> <strong>the</strong> mitochondrial 16S rRNA gene for phylogenetic<br />

and population studies <strong>of</strong> <strong>Crustacea</strong>. <strong>Crustacea</strong>n<br />

Issues 12 (in press).<br />

Submitted by Christoph Schubart,<br />

Universität Regensburg, Germany<br />

DECAPODA: BRACHYURA<br />

Although recently I published my arrangement <strong>of</strong><br />

<strong>the</strong> brachyuran families, I have some new discoveries<br />

in <strong>the</strong> brachyuran classification, but it is not<br />

finished and it will be published next year. I was<br />

able to classify all dromiacean families into superfamilies,<br />

but not <strong>the</strong> eubrachyuran ones, because<br />

<strong>the</strong>re are many families with obscure systematic position:<br />

Orithyiidae, Calappidae, Matutidae, Astenognathidae,<br />

Hexapodidae, Palicidae, Dairodidae<br />

and many up to now undescribed families (Acidopidae,<br />

Melybiidae, Speocarcinidae, etc.). Here are<br />

some <strong>of</strong> my remarks.<br />

(1) Dynomenidae are <strong>the</strong> most primitive Dromioidea,<br />

because only <strong>the</strong> last pair <strong>of</strong> legs is aberrant.<br />

(2) Among Homoloidea, <strong>the</strong> Poupinidae are <strong>the</strong><br />

most primitive because <strong>the</strong> last pair <strong>of</strong> legs are <strong>of</strong><br />

‘‘normal’’ structure but are partly subdorsal in position.<br />

(3) Raninidae are ‘‘Podotremata’’ (i.e. Dromiacea)<br />

because <strong>the</strong>ir sexual openings in both sexes<br />

are on <strong>the</strong> coxae <strong>of</strong> <strong>the</strong> legs (hence <strong>the</strong> name Podotremata).<br />

(4) The most primitive eubrachyuran<br />

family is <strong>the</strong> Atelecyclidae, because <strong>the</strong>y have <strong>the</strong><br />

antennules and antennae longitudinally directed, a<br />

narrow thoracic sternum, thoracic sternites 4/5–7/8<br />

continuous (entire), and sternites nearly regularly<br />

metamerized. (5) The Dorippidae are highly derived<br />

and aberrant: <strong>the</strong> dorsal position <strong>of</strong> <strong>the</strong> posterior<br />

pair <strong>of</strong> legs, <strong>the</strong> sternite 8 facing dorsally, and<br />

<strong>the</strong> narrowed buccal cavern all are secondarily attained.<br />

The similarity with <strong>the</strong> Dromiacea is thus<br />

superficial. (6) The same could be said for <strong>the</strong> Leucosiidae:<br />

highly derived crabs and consequently<br />

should be placed at <strong>the</strong> end <strong>of</strong> <strong>the</strong> classificatory<br />

scheme <strong>of</strong> <strong>the</strong> Heterotremata. (7) The Majidae are<br />

only one family with many subfamilies. The arrangement<br />

is enclosed [Števčić, Z. 1994. Contribution<br />

to <strong>the</strong> re-classification <strong>of</strong> <strong>the</strong> family Majidae.<br />

Periodicum Biologorum 96:419–420]. (8) The<br />

Par<strong>the</strong>nopidae are more primitive than Majidae,<br />

and <strong>the</strong>refore should be ahead <strong>of</strong> <strong>the</strong> Majidae. (9)<br />

The Retroplumidae are a very derived brachyuran<br />

family. (10) Geryonidae have a similar organization<br />

to <strong>the</strong> Goneplacidae s.s. (11) Your Xanthoidea is a<br />

highly polyphyletic group. (a) The most primitive<br />

‘‘xanthoids’’ are <strong>the</strong> Eriphiidae, not Menippidae!<br />

The most primitive Eriphiidae have sternites 4/5–<br />

7/8 entire, abdominal segments freely articulated in<br />

both sexes, and <strong>the</strong> second gonopod longer than <strong>the</strong><br />

first. They are probably related to Trapeziidae. In<br />

<strong>the</strong> same assemblage with <strong>the</strong> Eriphiidae are <strong>the</strong><br />

Pilumnoididae Guinot and Macpherson, 1987. (b)<br />

Xanthidae s.s. have [some] primitive representatives<br />

(Krausinae, with sternal sutures 4/5–7/8 entire),<br />

but abdominal segments 3–5 in <strong>the</strong> male are<br />

fused, and <strong>the</strong> second gonopod is short. They are<br />

related to <strong>the</strong> Panopeidae/Panopeinae and <strong>the</strong> Pseudorhombilidae.<br />

(c) Pilumnidae have a primitive abdomen<br />

(all segments freely articulated in both sexes)<br />

but specific first and second gonopods, <strong>the</strong> latter<br />

short. They are related to <strong>the</strong> Eumedonidae (in fact<br />

<strong>the</strong> Eumedoninae). (d) Goneplacidae s.s. are in fact<br />

a very small taxon, without any close relationships<br />

with <strong>the</strong> Xanthidae. They are probably close to <strong>the</strong><br />

Geryonidae and Euryplacidae/Euryplacinae. (12)<br />

The Potamidae are in fact a very difficult problem,<br />

however <strong>the</strong> gaps among subfamilies are not quite<br />

distinct. The gaps are not always [clear] and <strong>the</strong>refore<br />

<strong>the</strong> separation <strong>of</strong> <strong>the</strong> freshwater crabs into<br />

families remains uncertain. (13) I think that between<br />

Ocypodidae and Mictyridae and between<br />

Grapsidae and Gecarcinidae <strong>the</strong> gaps are not decisive<br />

and only Ocypodidae and Grapsidae are true<br />

families (this will be published later). (14) Finally,<br />

I think that <strong>the</strong> Cancroidea are not a taxon, <strong>the</strong>y<br />

are only a grade, not a clade (taxon i.e., monophyletic<br />

group). (15) Hepatinae are a subfamily <strong>of</strong> <strong>the</strong><br />

family Aethridae. (16) Palicidae belong to <strong>the</strong> Heterotremata,<br />

with no close affinity with <strong>the</strong> Ocypodidae.<br />

Submitted by Zdravko Števčić,<br />

Rudjer Boskovic Institute, Croatia<br />

DECAPODA: BRACHYURA<br />

Concerning my special knowledge, <strong>the</strong> Brachyura,<br />

I do not agree with all decisions (see my responses),<br />

but I respect <strong>the</strong>m. May I add my feeling, however.<br />

Concerning <strong>the</strong> Podotremata, <strong>the</strong> molecular data<br />

seem to outweigh all o<strong>the</strong>r considerations, despite<br />

<strong>the</strong> fact that <strong>the</strong> first results (Spears and Abele,<br />

1988; Spears, Abele, and Kim, 1992) were fragmentary,<br />

based only on very few taxa (only two<br />

Dromiidae were studied; and <strong>the</strong> conclusion was<br />

made without any Dynomenidae, Homolodromiidae,<br />

Homolidae, Latreilliidae, Cyclodorippidae,<br />

Cymonomidae, nor Phyllotymolinidae) and that <strong>the</strong><br />

new results are not yet published. I am happy to<br />

see that Spears now returns to <strong>the</strong> opinion that <strong>the</strong><br />

Dromiidae are true Brachyura, but we wait her paper<br />

where <strong>the</strong> new demonstration is given.<br />

Concerning your Section Raninoida, you write<br />

(p. 66, 69) that <strong>the</strong>re is ‘‘possibly a mistake.’’ I recognize<br />

that <strong>the</strong> problem <strong>of</strong> <strong>the</strong> placement <strong>of</strong> on <strong>the</strong><br />

one hand Cyclodorippidae, Cymonomidae, and<br />

Phyllotymolinidae, and on <strong>the</strong> o<strong>the</strong>r hand <strong>the</strong> Ran-<br />

112 Contributions in Science, Number 39 Appendix I: Comments and Opinions

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