Valvatacea

Ventral disc of Petraster speciosus, from Blake & Hotchkiss (2004). Scale bar = 5 mm.

Belongs within: Neoasteroidea.
Contains: Benthopectinidae, Paxillosida, Odontasteridae, Oreasteridae, Ophidiasteridae, Goniasteroidea.

The Valvatacea are a lineage of starfish for which potential synapomorphies are simple-bladed pedicellariae lacking a basal element, and the presence of true paxillae (radially symmetrically abactinal ossicles with a crowning cluster of spinelets; lost in some subgroups) (Blake 1987). In members of the Paxillosina, the mouth-angle plates are prominent, but in the Valvatida they are inconspicuous (Spencer & Wright 1966).

Characters (from Blake 1987): Body shape varied; flattened or arched, arms ranging from undifferentiated to short and rapidly tapering to elongate and columnar, discs small to large, interbrachial angles acute to rounded, area around mouth not depressed. Marginals, abactinals, actinals more or less clearly differentiated, closely spaced in most taxa; encrusting ossicles varied, pedicellariae present in many, few to common where present. Madreporite small to large, medial or near marginals. Abactinals generally closely spaced if skeleton not reduced, abactinal morphologies varied, true paxillae present in some taxa; at metamorphosis, calycinal ring discernible in most, differentiated or not in adults. Terminals small to large. Marginals in two series, mostly phanerozonate; intermarginals rare. If multiple actinal rows occur, longest row almost always adjacent to adambulacrals. Oral frame, ambulacral column protected by skeletal armour, and thickened dermal tissues in some taxa; oral frame dominated by oral ossicles. Ambulacral column rarely moderately compressed, adoral carina lacking; superambulacrals present or lacking. Ossicles present or absent from interbrachial septum. Papulae present, rarely extending beyond abactinals; tube feet almost always in two rows, tips with suckers, spicules absent or present. In development, bipinnaria can occur alone, anterior part of archenteron expands more or less symmetrically and cuts off a coelomic sac on each side; larval mouth, gut present; terminals not arising by fusion.

<==Valvatacea
    |--Paxillosina F67
    |    |--+--Benthopectinidae BH03
    |    |  `--Paxillosida BH03
    |    `--Hemizonina [Gnathasterina, Hemizonida] SW66
    |         |--Lepidasteridae SW66
    |         |    |--Lepidaster Forbes 1850 SW66
    |         |    |    `--*L. grayi Forbes 1850 SW66
    |         |    `--Devonistella Spencer 1927 SW66
    |         |         `--*D. filiciformis (Woodward 1874) [=Helianthaster filiciformis] SW66
    |         |--Palasterinidae [Lindstroemasterininae, Palaesterinidae, Palaeosolasteridae, Protactininae] SW66
    |         |    |--Palaeosolaster Stürtz 1899 (see below for synonymy) SW66
    |         |    |    `--*P. gregoryi Stürtz 1899 SW66
    |         |    `--Palasterina McCoy 1854 BH04 (see below for synonymy)
    |         |         |--*P. primaeva (Forbes 1848) [=Uraster primaevus] SW66
    |         |         `--P. antiqua SW66
    |         `--Petrasteridae SW66
    |              |--Uranaster Gregory 1899 BH04
    |              |    `--U. kinahani (Baily 1878) BH04
    |              `--Petraster Billings 1858 BH04 [Petrasteridae SW66]
    |                   |--*P. rigidus (Billings 1857) [=Palasterina rigidus] BH04
    |                   |--P. ramseyensis [=Uranaster ramseyensis] BE03
    |                   |--P. speciosus (Miller 1880) [=Palasterina speciosa] BH04
    |                   `--P. wigleyi Branstrator 1982 BH04
    `--Valvatida [Valvata] M09
         |  i. s.: Mithrodiidae [Mithrodiinae] M09
         |           |--Mithrodia Gray 1840 [incl. Heresaster Michelin 1844] SW66
         |           |    `--M. clavigera (Lamarck 1816) [=Asterias clavigera; incl. *M. spinulosa Gray 1840] SW66
         |           `--Thromidia Pope & Rowe 1977 M09
         |                |--*T. catalai Pope & Rowe 1977 M09
         |                |--T. brycei Marsh 2009 M09
         |                |--T. gigas (Mortensen 1935) [=Mithrodia gigas] M09
         |                `--T. seychellesensis Pope & Rowe 1977 M09
         |--Granulosina [Granulosa] SW66
         |    |  i. s.: Odontasteridae SW66
         |    |         Chaetaster Müller & Troschel 1840 [Chaetasteridae] SW66
         |    |           `--C. longipes (Retzius 1805) (see below for synonymy) SW66
         |    |         Archaster Müller & Troschel 1840 [Archasteridae] SW66
         |    |           `--*A. typicus Müller & Troschel 1840 SW66
         |    |         Oreasteridae SW66
         |    |         Ophidiasteridae SW66
         |    `--Goniasteroidea SW66
         `--Tumulosina SW66
              |--Monaster Etheridge 1892 BE03 [incl. Etheridgaster Gregory 1899 SW66; Monasteridae SW66]
              |    |--*M. clarkei (de Koninck 1878) BE03 (see below for synonymy)
              |    `--M. stutchburii (Etheridge 1892) [=Palaeaster (Monaster) stutchburii] F71
              |--Sphaerasteridae SW66
              |    |--Sphaeraster Quenstedt 1875 FR02 [=Sphaerites Quenstedt 1852 non Duftschmid 1805 SW66]
              |    |    `--*S. punctatus (Quenstedt 1852) [=*Sphaerites punctatus] SW66
              |    `--Valettaster Lambert 1914 FR02 (see below for synonymy)
              |         `--*V. ocellatus [=Oreaster ocellatus, *Tholaster ocellatus] SW66
              `--Stauranderasteridae SW66
                   |--Hadranderaster Spencer 1907 [incl. Stauraster Valette 1928] SW66
                   |    `--H. simplex (Geinitz 1871) (see below for synonymy) SW66
                   |--Stauranderaster Spencer 1907 SW66 [incl. Gasterometra Gislén 1925 WR78]
                   |    |--*S. boysii (Forbes 1848) [=Oreaster boysii] SW66
                   |    |--S. coronatus SW66
                   |    `--‘*Gasterometra’ polycirra Gislén 1925 WR78
                   `--Aspidaster de Loriol 1884 SW66
                        |--*A. delgadoi de Loriol 1884 SW66
                        `--A. bulbiferus SW66

Chaetaster longipes (Retzius 1805) [=Asterias longipes; incl. A. subulata Lamarck 1816, *Chaetaster subulatus] SW66

Hadranderaster simplex (Geinitz 1871) [=Oreaster simplex; incl. Pentaceros abbreviatus Spencer 1905, *Hadranderaster abbreviatus] SW66

*Monaster clarkei (de Koninck 1878) BE03 [=Palaeaster (*Monaster) clarkei F71, Etheridgaster clarkei F71]

Palaeosolaster Stürtz 1899 [incl. Echinasterias Stürtz 1899, Echinodiscaster Delage & Hérouard 1904, Echinodiscites Schuchert 1914, Echinodiscus Stürtz 1899 nec Leske 1778 nec Worthen & Miller 1883, Echinostella Stürtz 1899] SW66

Palasterina McCoy 1854 BH04 [=Palaeasterina Etheridge 1881 SW66; incl. Archasterina Lehmann 1957 SW66, Hisingeraster Stürtz 1900 SW66, Lindstroemaster Gregory 1899 SW66, Protactis Spencer 1927 SW66, Pseudopalasterina Stürtz 1900 SW66]

Valettaster Lambert 1914 FR02 [=Tholaster Spencer 1913 non Seunes 1890 SW66; incl. Asteriaceros Valette 1934 SW66]

*Type species of generic name indicated

REFERENCES

Blake, D. B. 1987. A classification and phylogeny of post-Palaeozoic sea stars (Asteroidea: Echinodermata). Journal of Natural History 21 (2): 481-528.

[BE03] Blake, D. B., & D. R. Elliott. 2003. Ossicular homologies, systematics, and phylogenetic implications of certain North American Carboniferous asteroids (Echinodermata). Journal of Paleontology 77 (3): 476–489.

[BH03] Blake, D. B., & H. Hagdorn. 2003. The Asteroidea (Echniodermata) of the Muschelkalk (Middle Triassic of Germany). Paläontologische Zeitschrift 77 (1): 23–58.

[BH04] Blake, D. B., & F. H. C. Hotchkiss. 2004. Recognition of the asteroid (Echinodermata) crown group: implications of the ventral skeleton. Journal of Paleontology 78: 359–370.

[F67] Fell, H. B. 1967. Echinoderm ontogeny. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt S. Echinodermata 1. General characters. Homalozoa—Crinozoa (except Crinoidea) vol. 1 pp. S60–S85. The Geological Society of America, Inc., and The University of Kansas: Lawrence (Kansas).

[F71] Fletcher, H. O. 1971. Catalogue of type specimens of fossils in the Australian Museum, Sydney. Australian Museum Memoir 13: 1–167.

[FR02] Fujita, T., & F. W. E. Rowe. 2002. Podosphaerasteridae fam. nov. (Echinodermata: Asteroidea: Valvatida), with a new species, Podosphaeraster toyoshiomaruae, from southern Japan. Species Diversity 7: 317–332.

[M09] Marsh, L. M. 2009. A new species of Thromidia (Echinodermata: Asteroidea) from Western Australia. Records of the Western Australian Museum 25 (2): 145–151.

[SW66] Spencer, W. K., & C. W. Wright. 1966. Asterozoans. In: Moore, R. C. (ed.) Treatise on invertebrate Paleontology pt U. Echinodermata 3 vol. 1 pp. U4–U107. The Geological Society of America, Inc., and The University of Kansas Press.

[WR78] Wienberg Rasmussen, H. 1978. Articulata. In: Moore, R. C., & C. Teichert (eds) Treatise on Invertebrate Paleontology pt. T. Echinodermata 2. Crinoidea vol. 3 pp. T813–T927. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).