Phylogeny of Holothuroidea (Echinodermata) inferred from ...

PHYLOGENY OF SEA CUCUMBERS 79
support via number of unambiguous synapomorphies Mertensio in orbis terrarum circumnavigatione ob-
and bootstrap percentages for branches leading to servatorum. Petropoli.
these clades is not likely to be an artefact of character Bronn HG. 1860. Die Klassen und Ordnungen der Strah-
choice, but a reflection of history. This in turn suggests lenthiere (Actinozoa), wissenschlaftliche dargestellt in
that these internal branches will prove equally difficult Wort und Bild. In: Bronn HG, ed. Bronn’s Klassen und
to resolve using molecular methods.
Ordnungen des Thier-Reiches, Volume II. Leipzig, 403–404.
The Triassic radiation comprises a clade of ho- Cherbonnier G. 1988. Echinodermes: Holothurides. Institut
lothuroids diagnosable in part by a gain of table ossen
Coopération: Faune de Madagascar 70: 1–292.
Français de Recherche Scientifique pour le Développement
icles. This group apparently diverged from the
Clark HL. 1898. Synapta vivipara: a contribution to the
ancestors of Elasipodida by the Middle Devonian (Fig.
morphology of echinoderms. Memoirs of the Boston Society
5). Fossil table ossicles (parafamily Priscopedatidae)
of Natural History 5: 53–88, pl. 11–15.
occurring between this time and the Triassic and as-
Clark HL. 1907. The apodous holothurians. Smithsonian
signed to Dendrochirotida and Dactylochirotida (e.g.
Contributions to Knowledge 35: 1–231.
Clavallus spicaudina; Gutschick & Grill, 1967) prob-
Cuénot L. 1891. Études morphologiques sur les échably
represent stem members of this clade. If Dendro- inodermes. Archives de Biologie 11: 313–680, pl. 24–31.
chirotida and Dactylochirotida do have a Triassic David B, Mooi R. 1998. Major events in the evolution of
origin, as indicated by this study, then very early echinoderms viewed by the light of embryology. In: Mooi
records of these groups based on simple plate ossicles R, Telford M, eds. Echinoderms: San Francisco. Rotterdam:
are also incorrect. Possibly, the plates indicate a Balkema, 21–28.
Palaeozoic divergence by stem members of the Triassic Deflandre-Rigaud M. 1946. Sur les divers types de
radiation. Alternatively, and if they are of holothuroid sclérites d’holothurides Oxfordiens des marnes de Villersur-Mer.
origin at all (e.g. Eocaudina; Reich, 1999), the ossicles
Comptes rendus de l’Académie des Sciences 223:
are from more distantly related forms, suggesting that 513–515.
imbricate plates and a testaceous morphology have Ekman S. 1926. Systematisch-phylogenetische Studien über
evolved and were lost several times in Holothuroidea. Elasipoden und Aspidochiroten. Zoologische Jahrbücher
(Anatomie und Ontogenie der Tiere) 47: 429–540.
Fell HB. 1965. The early evolution of the Echinozoa. Breviora
ACKNOWLEDGEMENTS 219: 1–17.
Fell HB, Moore RC. 1966. Echinozoans. In: Moore RC, ed.
We thank many people for their help. L. Buss, C. Treatise on Invertebrate Paleontology, Part U, Ech-
Massin and D. Pawson reviewed an earlier draft. Dis- inodermata 3. Volume 1. Lawrence, Kansas: Geological
cussions with J. Gauthier, R. Mooi, A. Smith and F. Society of America and University of Kansas Press, U108–
Rowe clarified some important ideas. C. Ahearn, F. U118.
Collier, L. Jarecki, P. Lambert, C. Massin, J. Moore, Ferrigno KF. 1970. Holothurian ossicles from the Dundee
G. Paulay, T. White and R. Woollacott provided lab Limestone in southwestern Ontario. Compass 47: 81–93.
space, specimens or hard-to-find literature. J. Kreychossicles.
Frizzell D, Exline H. 1955. Monograph of fossil holothurian
man and V. Alexandroba translated Russian. Funding
Bulletin of the Missouri University School of Met-
came from an Ernst Mayr Grant (Museum of Com- allurgy and Mines, Technical Series 89: 1–204.
parative Zoology), Lerner Gray Fund (American Mu- Frizzell D, Exline H. 1966. Holothuroidea-fossil record. In:
seum of Natural History), Falconwood Corporation, Moore RC, ed. Treatise on Invertebrate Paleontology, Part
Sigma Xi, John F. Enders Fund (Yale University), U, Echinodermata, Volume 2. Lawrence, Kansas: Geological
US Environmental Protection Agency and Women’s Society of America and University of Kansas Press, U646–
Seamen’s Friend Society of Connecticut.
U672.
Gage JD, Billett DSM. 1986. The family Myriotrochidae
Théel (Echinodermata: Holothuroidea) in the deep northeast
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