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20 S. Stöhr and M. Segonzac<br />

The number of disk scales and mouth, tentacle, and genital papillae <strong>in</strong>creases<br />

dur<strong>in</strong>g fur<strong>the</strong>r growth. At 3.1 mm dd, <strong>the</strong> second tentacle pore has moved to its<br />

f<strong>in</strong>al position at <strong>the</strong> mouth slit, framed by five to six papillae (Fig. 4D). Each jaw<br />

edge bears three to four papillae.<br />

At 5mmdd, <strong>the</strong> animals show adult characters, although <strong>the</strong> shape of some<br />

skeletal elements still differs <strong>from</strong> those of <strong>the</strong> largest adults. In addition to <strong>the</strong><br />

outer arm comb on <strong>the</strong> distal genital plate, a short <strong>in</strong>ner comb has formed of a few<br />

papillae on <strong>the</strong> first and second dorsal arm plates (Fig. 4F). These papillae fill <strong>the</strong><br />

gap between <strong>the</strong> genital plates and give <strong>the</strong> appearance of a cont<strong>in</strong>uous comb<br />

across <strong>the</strong> arm base. Of <strong>the</strong> four arm sp<strong>in</strong>es on <strong>the</strong> proximal arm, <strong>the</strong> dorsalmost is<br />

twice as large as <strong>the</strong> o<strong>the</strong>rs. The dorsal arm plates are longer than wi<strong>de</strong> on all but<br />

<strong>the</strong> most proximal segments. The mouth papillae have become elongated, sp<strong>in</strong>elike,<br />

and more numerous (Fig. 4E). The maximum size observed was 10 mm dd.<br />

Comparisons. The dorsal disk and arms of S. jolliveti are similar to those of<br />

Ophiura species, but <strong>the</strong> long, sp<strong>in</strong>e-like shape of <strong>the</strong> mouth papillae and tentacle<br />

scales is unknown <strong>in</strong> that genus. Also, <strong>the</strong> oral shield is ra<strong>the</strong>r small compared to<br />

that of Ophiura. The number of arm sp<strong>in</strong>es <strong>in</strong>creases dur<strong>in</strong>g ontogeny, and s<strong>in</strong>ce<br />

<strong>the</strong> ontogenetic stage of an <strong>in</strong>dividual may be unknown unless complete growth series<br />

are available, arm sp<strong>in</strong>e number is a character that must be used with caution.<br />

The growth series of S. jolliveti suggests that <strong>the</strong> f<strong>in</strong>al number of arm sp<strong>in</strong>es<br />

is five. Many species of Ophiura possess three to five arm sp<strong>in</strong>es, but <strong>the</strong>se are usually<br />

much shorter than an arm jo<strong>in</strong>t and appressed (Lyman 1878; Clark 1911, 1915,<br />

1917). The presence of tentacle scales on <strong>the</strong> first tentacle pore <strong>in</strong>si<strong>de</strong> <strong>the</strong> mouth slit<br />

has not been <strong>de</strong>scribed for any o<strong>the</strong>r species of ophiuroid, but <strong>in</strong> <strong>the</strong> ophiacanthid<br />

Ophiomyces Lyman, 1869, a scale-like papilla is present at <strong>the</strong> first tentacle pore, although<br />

this is an overlooked and unreported feature (F. Hotchkiss, pers. comm.).<br />

The ontogeny of S. jolliveti is similar to that of Ophiura, but a buccal scale is<br />

absent <strong>in</strong> <strong>the</strong> small postlarvae of this species and <strong>the</strong> first tooth forms quite late<br />

dur<strong>in</strong>g <strong>de</strong>velopment. In all analyzed species of Ophiura, both <strong>the</strong> first tooth and <strong>the</strong><br />

buccal scale are present <strong>in</strong> <strong>the</strong> earliest stage (Sumida et al. 1998; Stöhr 2005). It is<br />

unclear which condition is plesiomorphic. The madreporite <strong>in</strong> <strong>the</strong> present species<br />

cannot be dist<strong>in</strong>guished <strong>from</strong> <strong>the</strong> oral shields until almost <strong>the</strong> adult stage, and all<br />

oral shields <strong>de</strong>velop later than <strong>in</strong> many o<strong>the</strong>r species (Sumida et al. 1998; Stöhr<br />

2005). Dur<strong>in</strong>g <strong>de</strong>velopment, some parts of <strong>the</strong>se pentamerous animals <strong>de</strong>velop<br />

faster than o<strong>the</strong>rs, which produces an asymmetrical pattern at some stages, e.g., k-<br />

plates on some but not all radii. In particular, <strong>the</strong> <strong>de</strong>velopment of <strong>the</strong> mouth papillae<br />

differs between jaws and <strong>the</strong> f<strong>in</strong>al number of papillae <strong>in</strong> <strong>the</strong> adult is different on<br />

different jaws.<br />

Sp<strong>in</strong>ophiura jolliveti shares some characters with Ophiura and may be a modified<br />

representative of that genus, but <strong>the</strong> strik<strong>in</strong>g differences also merit a taxonomic<br />

separation, at <strong>the</strong> risk of leav<strong>in</strong>g Ophiura paraphyletic. We thus propose a<br />

new genus, which may be re-evaluated when <strong>the</strong> relationships of <strong>the</strong> species and<br />

its orig<strong>in</strong> are better un<strong>de</strong>rstood.<br />

Habitat and distribution. Sp<strong>in</strong>ophiura jolliveti was found only at vents. The<br />

large number of specimens collected (270) suggests that this species is adapted to<br />

hydro<strong>the</strong>rmal environments. It is present at several vent sites on <strong>the</strong> EPR at 9°N<br />

(Barbecue, Mussel Bed, <strong>East</strong> Wall, and Tra<strong>in</strong> <strong>Station</strong>), 13°N (Genesis, Parigo, Julie,<br />

and Grandbonum), 17°25S (Oasis-Rehu), 17°35S (Wormwood), and 18°36S (Animal

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