Valvatacea

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

Belongs within: Ambuloasteroidea.
Contains: Benthopectinidae, Paxillosida, Odontasteridae, Asterinidae, Pseudarchasterinae, Goniasteroidea, Oreasteridae, Ophidiasteridae.

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
    |--PaxillosinaF67
    |    |--+--BenthopectinidaeBH03
    |    |  `--PaxillosidaBH03
    |    `--Hemizonina [Gnathasterina, Hemizonida]SW66
    |         |--LepidasteridaeSW66
    |         |    |--Lepidaster Forbes 1850SW66
    |         |    |    `--*L. grayi Forbes 1850SW66
    |         |    `--Devonistella Spencer 1927SW66
    |         |         `--*D. filiciformis (Woodward 1874) [=Helianthaster filiciformis]SW66
    |         |--Palasterinidae [Lindstroemasterininae, Palaesterinidae, Palaeosolasteridae, Protactininae]SW66
    |         |    |--Palaeosolaster Stürtz 1899 (see below for synonymy)SW66
    |         |    |    `--*P. gregoryi Stürtz 1899SW66
    |         |    `--Palasterina McCoy 1854BH04 (see below for synonymy)
    |         |         |--*P. primaeva (Forbes 1848) [=Uraster primaevus]SW66
    |         |         `--P. antiquaSW66
    |         `--PetrasteridaeSW66
    |              |--Uranaster Gregory 1899BH04
    |              |    `--U. kinahani (Baily 1878)BH04
    |              `--Petraster Billings 1858BH04
    |                   |--*P. rigidus (Billings 1857) [=Palasterina rigidus]BH04
    |                   |--P. ramseyensis [=Uranaster ramseyensis]BE03
    |                   |--P. speciosus (Miller 1880) [=Palasterina speciosa]BH04
    |                   `--P. wigleyi Branstrator 1982BH04
    `--Valvatida [Granulosa, Granulosina, Valvata]M09
         |  i. s.: Arthraster Forbes 1848SW66 [ArthrasteridaeSG93, Arthrasterinae]
         |           |--*A. dixoni Forbes 1848SW66
         |           `--A. cristatus Spencer 1913SG93
         |         Pycinasteridae [Pycinasterinae]SG93
         |           |--Phocidaster Spencer 1913SW66
         |           |    `--*P. grandis Spencer 1913SW66
         |           `--Pycinaster Spencer 1907 [=Pycnaster Sladen 1891 non Pomel 1883]SW66
         |                |--*P. angustatus (Forbes 1848) [=Goniaster (Goniodiscus) angustatus, *Pycnaster angustatus]SW66
         |                |--P. mortenseni Mercier 1935SG93
         |                `--P. peyroti Valette 1925SG93
         |         OdontasteraceaB87
         |           |--OdontasteridaeB87
         |           `--ChaetasteridaeB87
         |                |--Chaetasterina gracilis Hess 1970SG93
         |                `--Chaetaster Müller & Troschel 1840SW66
         |                     `--C. longipes (Retzius 1805) (see below for synonymy)SW66
         |--GaneriaceaB87
         |    |--AsterinidaeBTH00
         |    `--Ganeriidae [Cryasteridae]B87
         |         |--Ganeria Gray 1847SW66
         |         |    `--*G. falklandica Gray 1847SW66
         |         |--Aleutiaster Clark 1939SW66
         |         |    `--*A. schafferi Clark 1939SW66
         |         |--Cycethra Bell 1881 [incl. Lebrunaster Perrier 1891]SW66
         |         |    `--C. verrucosa (Philippi 1857) [=Goniodiscus verrucosus; incl. *C. simplex Bell 1881]SW66
         |         |--Leilaster Clark 1938SW66
         |         |    `--*L. radians (Perrier 1881) [=Korethraster radians]SW66
         |         |--Magdalenaster Koehler 1907SW66
         |         |    `--*M. arcticus Koehler 1907SW66
         |         |--Perknaster Sladen 1889 [incl. Cribellopsis Koehler 1917, Cryaster Koehler 1906]SW66
         |         |    `--*P. fuscus Sladen 1889SW66
         |         |--Scotiaster Koehler 1908SW66
         |         |    `--*S. inornatus Koehler 1908SW66
         |         `--Tarachaster Fisher 1913SW66
         |              `--*T. tenuis Fisher 1913SW66
         `--+--Archaster Müller & Troschel 1840BTH00, SW66 [Archasteracea, Archasteridae]
            |    |--*A. typicus Müller & Troschel 1840SW66
            |    |--A. angulatus Müller & Troschel 1842 [incl. A. laevis Clark 1938]SM15
            |    |--A. patersoni Spencer 1915SG93
            |    `--A. tenuispinusN79
            `--+--PseudarchasterinaeBTH00
               `--+--GoniasteroideaBTH00
                  `--+--OreasteraceaB87
                     |    |--OreasteridaeSW66
                     |    `--Acanthaster Gervais 1841SW66 (see below for synonymy)
                     |         `--A. planci (Linnaeus 1758) [=Asterias planci; incl. *Ac. echinus Gervais 1841]SW66
                     `--OphidiasteraceaB87
                          |--OphidiasteridaeSW66
                          `--Mithrodiidae [Mithrodiinae]B87
                               |--Mithrodia Gray 1840 [incl. Heresaster Michelin 1844]SW66
                               |    `--M. clavigera (Lamarck 1816) [=Asterias clavigera; incl. *M. spinulosa Gray 1840]SW66
                               `--Thromidia Pope & Rowe 1977M09
                                    |--*T. catalai Pope & Rowe 1977M09
                                    |--T. brycei Marsh 2009M09
                                    |--T. gigas (Mortensen 1935) [=Mithrodia gigas]M09
                                    `--T. seychellesensis Pope & Rowe 1977M09

Acanthaster Gervais 1841SW66 [=Echinaster Gray 1840 non Müller & Troschel 1840SW66; AcanthasteridaeB87, Echinasterina]

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

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

*Type species of generic name indicated

References

[B87] 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.

[BTH00] Blake, D. B., A. Tintori & H. Hagdorn. 2000. A new, early crown-group asteroid (Echinodermata) from the Norian (Triassic) of northern Italy. Rivista Italiana di Paleontologia e Stratigrafia 106: 141–156.

[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).

[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.

[N79] Norman, A. M. 1879. The Mollusca of the fiords near Bergen, Norway. Journal of Conchology 2: 8–77.

[SM15] Sampey, A., & L. M. Marsh. 2015. Kimberley marine biota. Historical data: echinoderms. Records of the Western Australian Museum Supplement 84: 207–246.

[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.

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