Neoasteroidea

Poraniomorpha hispida, copyright Claude Nozères.

Belongs within: Ambuloasteroidea.
Contains: Valvatacea, Spinulosacea, Asteriadina.

The Neoasteroidea represents the starfish crown group, including all living starfish and all known Asteroidea from the Mesozoic and Cenozoic eras (Blake & Hagdorn 2003).

Characters (from Blake & Hagdorn 2003): Axillary or axillary series absent. More or less extensive actinal series differentiated ventral to marginal series. Actinals in basal taxa more or less irregular in form and arrangement but with longest series adjacent to marginal series. Adambulacrals overlapping in distal direction. Adambulacrals little wider than ambulacral articular flanges; ambulacrals typically larger than adambulacrals. Longitudinal ambulacral connective surfaces inclined proximally such that proximal flange of each ambulacral more or less clearly overlapping distal side of more proximal ossicle, ossicular form is not bilateral.

Neoasteroidea [Valvatina]
    |--+--+--Valvatacea BH03
    |  |  `--Spinulosacea BH03
    |  `--Poraniidae BH03
    |       |--Noriaster Blake, Tintori & Hagdorn 2000 BH04
    |       |    `--*N. barberoi Blake et al. 2000 BH03
    |       |--Chondraster Verrill 1895 SW66
    |       |    `--*C. grandis (Verrill 1878) [=Porania grandis] SW66
    |       |--Marginaster Perrier 1881 [incl. Cheilaster Bell 1892] SW66
    |       |    `--*M. pectinatus Perrier 1881 SW66
    |       |--Petricia Gray 1847 SW66
    |       |    `--P. vernicina (Lamarck 1816) [=Asterias vernicina; incl. *P. punctata Gray 1847] SW66
    |       |--Poraniella Verrill 1914 SW66
    |       |    `--*P. regularis Verrill 1914 SW66
    |       |--Poranisca Verrill 1914 SW66
    |       |    `--*P. lepida Verrill 1914 SW66
    |       |--Pseudoporania Dons 1936 SW66
    |       |    `--*P. stormi Dons 1936 SW66
    |       |--Sphaeriaster Dons 1939 [=Sphaeraster Dons 1938 non Quenstedt 1875] SW66
    |       |    `--*S. berthae (Dons 1938) [=*Sphaeraster berthae] SW66
    |       |--Spoladaster Fisher 1940 SW66
    |       |    `--*S. brachyactis (Clark 1923) [=Cryaster brachyactis] SW66
    |       |--Tylaster Danielsson & Koren 1881 SW66
    |       |    `--*T. willei Danielsson & Koren 1881 SW66
    |       |--Porania Gray 1840 BH04 [incl. Glabraster Clark 1916 SW66]
    |       |    `--P. pulvillus (Müller 1788) (see below for synonymy) SW66
    |       |         |--P. p. pulvillus BH04
    |       |         `--P. p. insignis Muller 1776 BH04
    |       `--Poraniomorpha Danielsson & Koren 1881 (see below for synonymy) SW66
    |            |--*P. rosea Danielsson & Koren 1881 SW66
    |            |--P. hispida BK77
    |            `--P. tumida BK77
    `--Forcipulatacea [Forcipulatae, Forcipulatida] BH03
         |  i. s.: Argoviaster Hess 1972 B02
         |         Hystrixasterias Blake 1990 B02
         |--Asteriadina SW66
         `--Brisingidae [Brisingina, Euclasteroides] SW66
              |--Astrocles Fisher 1917 SW66
              |    `--*A. actinodetus Fisher 1917 SW66
              |--Astrolirus Fisher 1917 SW66
              |    `--*A. panamensis (Ludwig 1905) [=Brisinga panamensis] SW66
              |--Astrostephane Fisher 1917 SW66
              |    `--*A. moluccana (Fisher 1916) [=Brisinga moluccana] SW66
              |--Belgicella Ludwig 1903 SW66
              |    `--*B. racowitzana Ludwig 1903 SW66
              |--Brisingaster de Loriol 1883 SW66
              |    `--*B. robillardi de Loriol 1883 SW66
              |--Brisingenes Fisher 1917 SW66
              |    `--*B. mimica (Fisher 1916) [=Brisinga mimica] SW66
              |--Colpaster Sladen 1889 SW66
              |    `--*C. scutigerulus Sladen 1889 SW66
              |--Craterobrisinga Fischer 1916 SW66
              |    `--*C. panopla (Fisher 1906) [=Brisinga panopla] SW66
              |--Freyella Perrier 1885 SW66
              |    `--*F. spinosa Perrier 1885 SW66
              |--Freyellaster Fisher 1918 SW66
              |    `--*F. fecundus (Fisher 1905) [=Freyella fecunda] SW66
              |--Hymenodiscus Perrier 1884 SW66
              |    `--*H. agassizi Perrier 1884 SW66
              |--Odinella Fisher 1940 SW66
              |    `--*O. nutrix Fisher 1940 SW66
              |--Parabrisinga Hayashi 1948 SW66
              |    `--*P. pellucida Hayashi 1948 SW66
              |--Stegnobrisinga Fisher 1916 SW66
              |    `--*S. placoderma (Fisher 1916) [=Brisinga (*Stegnobrisinga) placoderma] SW66
              |--Brisinga Asbjornsen 1856 SW66
              |    |--*B. endecacnemos Asbjornsen 1856 SW66
              |    `--B. mediterranea SW66
              |--Brisingella Fisher 1917 SW66
              |    |--*B. fragilis (Fisher 1906) [=Brisinga fragilis] SW66
              |    `--B. coronata BK77
              `--Odinia Perrier 1885 SW66
                   |--*O. semicoronata Perrier 1885 SW66
                   `--O. pandina BK77

Porania pulvillus (Müller 1788) [=Asterias pulvillus; incl. A. gibbosus Leach 1817 non A. gibbosa Pennant 1777, *Porania gibbosa] SW66

Poraniomorpha Danielsson & Koren 1881 [incl. Culcitopsis Verrill 1914, Lasiaster Sladen 1889, Rhegaster Sladen 1883] SW66

*Type species of generic name indicated

REFERENCES

[BK77] Barel, C. D. N., & P. G. N. Kramers. 1977. A survey of the echinoderm associates of the north-east Atlantic area. Zoologische Verhandelingen 156: 1–159.

[B02] Blake, D. B. 2002. Compsaster formosus Worthen & Miller (Asteroidea; Echinodermata): A Carboniferous homeomorph of the post-Paleozoic Asteriidae. Paläontologische Zeitschrift 76 (2): 357–367.

[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.

[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.

Goniasteroidea

Podosphaeraster sp., from Christopher Mah.

Belongs within: Valvatacea.
Contains: Goniasterinae, Anthenoidinae, Hippasteriinae, Pseudarchasterinae.

The Goniasteroidea are a group of starfish with flat, plate-like abactinals (Blake 1987).

Characters (from Blake 1987, as Goniasteracea): Body shape usually more or less flattened with discs large, interbrachial arcs rounded; some species subspherical; multi-armed species lacking; most species moderate to large in size but some small. Abactinals, marginals, actinals almost always differentiated, body moderately stout with closely fitted ossicles, abactinal surface somewhat flexible unless body is subspherical; dermal and skeletal layer thickened in Asterodiscididae. Encrusting ossicles varied, mostly in the form of spinelets, granules, also spines, tubercles; pedicellariae either absent or simple, common; blades varied, sessile or alveolar. Abactinals in form of paxillae, tabulae or flattened plates; marginals enlarged, varied, intermarginals almost always lacking. Ambulacrals, adambulacrals varied, from thin to quite stout; spicules lacking from tube feet; superambulacrals present or absent, interbrachial septa almost always lacking ossicles.

<==Goniasteroidea [Goniasteracea]
    |--Asterodiscides MG-H11 [Asterodiscididae FR02]
    |    |--A. macroplax Rowe 1985 MG-H11
    |    `--A. tessellatus Rowe 1977 MG-H11
    |--Podosphaeraster Clark in Clark & Wright 1962 [Podosphaerasteridae] FR02
    |    |--*P. polyplax Clark in Clark & Wright 1962 FR02
    |    |--P. gustavei Rowe 1985 FR02
    |    |--P. pulvinatus Rowe & Nichols 1980 FR02
    |    |--P. thalassae Cherbonnier 1970 FR02
    |    `--P. toyoshiomaruae Fujita & Rowe 2002 FR02
    `--Goniasteridae FR02
         |  i. s.: Lithosoma Fisher 1911 SW66
         |           |--*L. actinometra Fisher 1911 SW66
         |           `--L. penichra Fisher 1917 MG-H11
         |         Milteliphaster Alcock 1893 SW66
         |           |--*M. woodmasoni Alcock 1893 SW66
         |           `--M. regenerator (Doderlein 1922) MG-H11
         |         Rosaster Perrier 1894 [incl. Nereidaster Verrill 1899] SW66
         |           `--*R. alexandri (Perrier 1881) [=Pentagonaster alexandri] SW66
         |         Codellaster Blake & Kues 2002 BK02
         |           `--*C. keepersae Blake & Kues 2002 BK02
         |         Comptonia Gray 1840 E01
         |           |--*C. elegans Gray 1840 SW66
         |           `--C. comptoni (Forbes 1848) E01
         |         Noviaster Vallette 1929 E01
         |           `--*N. lissajousi Valette 1929 SW66
         |         Mediaster Stimpson 1857 [incl. Isaster Verrill 1894 non Desor 1858; Mediasterinae] SW66
         |           |--*M. aequalis Stimpson 1857 SW66
         |           |--M. australiensis Clark 1916 MG-H11
         |           `--M. hayi Blake 1986 BK02
         |         Nymphaster Sladen 1889 SW66
         |           |--N. arenatus (Perrier 1881) [=Pentagonaster arenatus, *N. protentus Sladen 1889] SW66
         |           |--N. moebii Studer 1884 MG-H11
         |           `--N. moluccanus Fisher 1913 MG-H11
         |         Calliaster Gray 1840 SW66
         |           `--*C. childreni Gray 1840 SW66
         |         Calliderma Gray 1847 [incl. Tomidaster Sladen 1891] SW66
         |           |--*C. emma Gray 1847 SW66
         |           `--C. smithiae SW66
         |         Cenomanaster Wright 1951 [=Jacobella Mercier 1935 nec Jeannet 1908 nec Patte 1926] SW66
         |           `--*C. cenomanensis (Mercier 1935) [=*Jacobella cenomanensis] SW66
         |         Chomataster Spencer 1913 [incl. Huraeaster Valette 1915] SW66
         |           `--*C. acules Spencer 1913 SW66
         |         Crateraster Spencer 1913 [incl. Austinaster Adkins 1928] SW66
         |           `--*C. quinqueloba (Goldfuss 1822) [=Asterias quinqueloba] SW66
         |         Forbesiaster de Loriol 1909 SW66
         |           `--*F. wrighti de Loriol 1909 SW66
         |         ‘Indiaster’ Rao 1957 non Lambert 1920 SW66
         |           `--*I. krishna Rao 1957 SW66
         |         Leptogonium Pomel 1887 SW66
         |           `--*L. mauritanicum Pomel 1887 SW66
         |         Mastaster Mercier 1935 SW66
         |           `--*M. villersensis Mercier 1935 SW66
         |         Miopentagonaster Mercier 1935 SW66
         |           `--*M. calloviensis Mercier 1935 SW66
         |         Ophryaster Spencer 1913 SW66
         |           |--*O. oligoplax (Sladen 1891) [=Nymphaster oligoplax] SW66
         |           `--O. magnus SW66
         |         Pachyaster de Loriol 1909 SW66
         |           `--*P. aegyptiacus de Loriol 1909 SW66
         |         Spenceraster Lambert 1913 [=Trachyaster Spencer 1913 non Pomel 1883] SW66
         |           `--*S. rugosus (Spencer 1907) [=Nymphaster rugosus, *Trachyaster rugosus] SW66
         |         Teichaster Spencer 1913 SW66
         |           `--*T. favosus Spencer 1913 SW66
         |         Tylasteria Valette 1930 [=Tylaster Spencer 1913 non Danielsson & Koren 1881] SW66
         |           `--*T. jurensis (Goldfuss 1822) [=Asterias jurensis, *Tylaster jurensis] SW66
         |         Amphiaster Verrill 1868 SW66
         |           `--*A. insignis Verrill 1868 SW66
         |         Astroceramus Fisher 1906 SW66
         |           `--*A. callimorphus Fisher 1906 SW66
         |         Astrothauma Fisher 1913 SW66
         |           `--*A. euphylacteum Fisher 1913 SW66
         |         Circeaster Koehler 1909 SW66
         |           `--*C. marcelli Koehler 1909 SW66
         |         Eugoniaster Verrill 1899 SW66
         |           `--*E. investigatoris (Alcock 1893) [=Pentagonaster investigatoris] SW66
         |         Gigantaster Döderlein 1924 SW66
         |           `--*G. weberi Döderlein 1924 SW66
         |         Gilbertaster Fisher 1906 SW66
         |           `--*G. anacanthus Fisher 1906 SW66
         |         Johannaster Koehler 1909 SW66
         |           `--*J. superbus Koehler 1909 SW66
         |         Litonotaster Verrill 1899 SW66
         |           `--*L. intermedius (Perrier 1884) [=Pentagonaster intermedius] SW66
         |         Lydiaster Koehler 1909 SW66
         |           `--*L. johannae Koehler 1909 SW66
         |         Mahabissaster Macan 1938 SW66
         |           `--*M. zengi Macan 1938 SW66
         |         Mariaster Clark 1916 SW66
         |           `--*M. giganteus (Goto 1914) [=Johannaster giganteus] SW66
         |         Notioceramus Fisher 1940 SW66
         |           `--*N. anomalus Fisher 1940 SW66
         |         Peltaster Verrill 1899 SW66
         |           `--P. nidarosiensis (Storm 1881) [=Goniaster nidarosiensis; incl. *P. hebes Verrill 1899] SW66
         |         Progoniaster Döderlein 1924 SW66
         |           `--*P. atavus Döderlein 1924 SW66
         |         Pseudogoniodiscaster Livingstone 1930 SW66
         |           `--*P. wardi Livingstone 1930 SW66
         |         Sibogaster Döderlein 1924 SW66
         |           `--*S. digitatus Döderlein 1924 SW66
         |         Styphlaster Clark 1938 SW66
         |           `--*S. notabilis Clark 1938 SW66
         |--Goniasterinae E01
         |--Anthenoidinae SW66
         |--Hippasteriinae SW66
         |--Pseudarchasterinae SW66
         |--Chitonaster Sladen 1889 [Chitonasterinae] SW66
         |    `--*C. cataphractus Sladen 1889 SW66
         |--Nectriinae SW66
         |    |--Nectria Gray 1840 SW66
         |    |    `--*N. ocellifera (Lamarck 1816) [=Asterias ocellifera] SW66
         |    `--Nectriaster Clark 1946 SW66
         |         `--*N. monacanthus (Clark 1916) [=Mediaster monacanthus] SW66
         `--Pycinasterinae SW66
              |--Pycinaster Spencer 1907 [=Pycnaster Sladen 1891 non Pomel 1883] SW66
              |    `--*P. angustatus (Forbes 1848) [=Goniaster (Goniodiscus) angustatus, *Pycnaster angustatus] SW66
              `--Phocidaster Spencer 1913 SW66
                   `--*P. grandis Spencer 1913 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.

[BK02] Blake, D. B, & B. S. Kues. 2002. Homeomorphy in the Asteroidea (Echinodermata); a new late Cretaceous genus and species from Colorado. Journal of Paleontology 76 (6): 1007–1013.

[E01] Eagle, M. K. 2001. A new species of Cottreauaster (Asteroidea: Echinodermata) from the Middle Jurassic of New Zealand. Records of the Auckland Museum 37: 93–100.

[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.

[MG-H11] McEnnulty, F. R., K. L. Gowlett-Holmes, A. Williams, F. Althaus, J. Fromont, G. C. B. Poore, T. D. O’Hara, L. Marsh, P. Kott, S. Slack-Smith, P. Alderslade & M. V. Kitahara. 2011. The deepwater megabenthic invertebrates on the western continental margin of Australia (100–1100 m depths): composition, distribution and novelty. Records of the Western Australian Museum Supplement 80: 1–191.

[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.

Paxillosida

Seven-armed starfish Luidia ciliaris, copyright Fiona Crouch.

Belongs within: Valvatacea.
Contains: Astropectininae.

The Paxillosida are a group of starfish associated with sandy or muddy habitats. They mostly lack a number of features found in other starfish, such as an anus, extra-oral feeding, suckered tube feet, or a brachiolaria larval stage (Matsubara et al. 2005).

<==Paxillosida [Paxillosa]
    |  i. s.: Radiasteridae [Mimasterinae] MK05
    |           |--Radiaster Perrier 1881 [incl. Mimaster Sladen 1882] SW66
    |           |    `--*R. elegans Perrier 1881 SW66
    |           `--Gephyriaster Fisher 1910 SW66
    |                `--*G. swifti (Fisher 1905) [=Mimaster swifti] SW66
    |--Astropectinidae [Diplozonina, Plutonasteridae, Primasterinae] MK05
    |    |  i. s.: Proserpinaster euryactis (Fisher 1913) MG-H11
    |    |--Astropectininae SW66
    |    `--Craspidaster Sladen 1889 [incl. Nauricia Gray 1840 (nom. dub.); Craspidasterinae] SW66
    |         `--*C. hesperus (Müller & Troschel 1840) [=Archaster hesperus] SW66
    |--Luidia Forbes 1839 BH04 (see below for synonymy)
    |    |  i. s.: L. avicularia Fisher 1913 MG-H11
    |    |         L. hardwicki (Gray 1840) MG-H11
    |    |         L. quinaria MK05
    |    |         L. sarsi Düben & Koren 1884 W01 [incl. Bipinnaria asterigera H04]
    |    |--L. (Luidia) ciliaris (Philippi 1837) [=Asterias ciliaris; incl. *L. fragilissima Forbes 1839] SW66
    |    `--L. (Platasterias Gray 1871) BK02, B02 [Platasteriidae]
    |         `--L. (*P.) latiradiata Gray 1871 BK02, SW66
    `--Cribellina SW66
         |--Goniopectinidae SW66
         |    |--Goniopectininae SW66
         |    |    |--Goniopecten Perrier 1881 SW66
         |    |    |    `--*G. demonstrans Perrier 1881 SW66
         |    |    `--Prionaster Verrill 1889 SW66
         |    |         `--*P. elegans Verrill 1889 SW66
         |    `--Ctenodiscinae [Ctenodiscidae] SW66
         |         |--Pectinidiscus Ludwig 1900 SW66
         |         |    `--*P. annae Ludwig 1900 SW66
         |         `--Ctenodiscus Müller & Troschel 1842 [incl. Anodiscus Perrier 1869] SW66
         |              |--C. crispatus (Retzius 1805) (see below for synonymy) SW66
         |              `--C. orientalis Fisher 1913 MG-H11
         `--Porcellanasteridae SW66
              |--Sidonaster Koehler 1909 SW66
              |    `--*S. vaneyi Koehler 1909 SW66
              |--Porcellanaster Wyville-Thomson 1877 (see below for synonymy) SW66
              |    `--*P. ceruleus Wyville-Thomson 1877 SW66
              |--Abyssaster Madsen 1961 SW66
              |    `--*A. tara (Wood-Mason & Alcock 1891) [=Hyphalaster tara] SW66
              |--Benthogenia Fisher 1911 SW66
              |    `--*B. cribellosa Fisher 1911 SW66
              |--Eremicaster Fisher 1905 SW66
              |    `--E. gracilis (Sladen 1883) (see below for synonymy) SW66
              |--Hyphalaster Sladen 1883 SW66
              |    `--*H. inermis Sladen 1883 SW66
              |--Lysaster Bell 1909 SW66
              |    `--*L. lorioli Bell 1909 SW66
              |--Styracaster Sladen 1883 [incl. Chunaster Ludwig 1907, Machairaster Perrier 1884 (n. n.)] SW66
              |    `--*S. horridus Sladen 1883 SW66
              `--Thoracaster Sladen 1883 SW66
                   `--*T. cylindratus Sladen 1883 SW66

Ctenodiscus crispatus (Retzius 1805) [=Asterias crispata; incl. A. polaris Sabine 1821, *Ctenodiscus polaris] SW66

Eremicaster gracilis (Sladen 1883) [=Porcellanaster gracilis; incl. P. tenebrarius Fisher 1905, *Eremicaster tenebrarius] SW66

Luidia Forbes 1839 BH04 [=Luydia Düben & Koren 1847 SW66; incl. Alternaster Döderlein 1920 SW66, Armaster Döderlein 1920 SW66, Astrella Perrier 1882 SW66, Denudaster Döderlein 1920 SW66, Hemicnemis Müller & Troschel 1840 SW66, Integraster Döderlein 1920 SW66, Maculaster Döderlein 1920 SW66, Penangaster Döderlein 1920 SW66, Petalaster Gray 1840 SW66, Quinaster Döderlein 1920 SW66, Senegaster Döderlein 1920 SW66; Luidiidae MK05]

Porcellanaster Wyville-Thomson 1877 [incl. Albatrossaster Ludwig 1907, Albatrossia Ludwig 1905 non Jordan & Evermann 1898, Caulaster Perrier 1882] SW66

*Type species of generic name indicated

REFERENCES

[B02] Blake, D. B. 2002. Compsaster formosus Worthen & Miller (Asteroidea; Echinodermata): a Carboniferous homeomorph of the post-Paleozoic Asteriidae. Paläontologische Zeitschrift 76 (2): 357–367.

[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.

[BK02] Blake, D. B, & B. S. Kues. 2002. Homeomorphy in the Asteroidea (Echinodermata); a new late Cretaceous genus and species from Colorado. Journal of Paleontology 76 (6): 1007–1013.

[H04] Haeckel, E. 1899–1904. Kunstformen der Natur. Bibliographisches Institut: Leipzig und Wien.

[MK05] Matsubara, M., M. Komatsu, T. Araki, S. Asakawa, S. Yokobori, K. Watanabe & H. Wada. 2005. The phylogenetic status of Paxillosida (Asteroidea) based on complete mitochondrial DNA sequences. Molecular Phylogenetics and Evolution 36: 598–605.

[MG-H11] McEnnulty, F. R., K. L. Gowlett-Holmes, A. Williams, F. Althaus, J. Fromont, G. C. B. Poore, T. D. O’Hara, L. Marsh, P. Kott, S. Slack-Smith, P. Alderslade & M. V. Kitahara. 2011. The deepwater megabenthic invertebrates on the western continental margin of Australia (100–1100 m depths): composition, distribution and novelty. Records of the Western Australian Museum Supplement 80: 1–191.

[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.

[W01] Williamson, D. I. 2001. Larval transfer and the origins of larvae. Zoological Journal of the Linnean Society 131: 111–122.

Ophiacanthidae

Ophiacantha bidentata, copyright Claude Nozères.

Belongs within: Ophiuroidea.

The Ophiacanthidae are a group of brittle stars with slender arms bearing numerous long, often glassy spines (Spencer & Wright 1966).

Characters (from Spencer & Wright 1966): Arms slender, commonly constricted at nodes. Dorsal and ventral arm plates very small. Arm spines long, numerous, at angle to arm, commonly glassy and serrate. Disc with granules and spinules. Distal vertebrae may be partly divided longitudinally by series of pores.

<==Ophiacanthidae [Ophiomycetidae]
    |--Sinophiura multispina (Koehler 1922) MG-H11
    |--Ophiomedea Koehler 1906 F61
    |    `--*O. duplicata Koehler 1906 SW66
    |--Ophiocamax Lyman 1878 SW66
    |    `--*O. vitrea Lyman 1878 SW66
    |--Ophiocymbium Lyman 1880 SW66
    |    `--*O. cavernosum Lyman 1880 SW66
    |--Ophiolimna Verrill 1899 SW66
    |    `--*O. bairdi (Lyman 1883) [=Ophiacantha bairdi] SW66
    |--Ophiotoma Lyman 1883 [incl. Ophiopora Verrill 1899] SW66
    |    `--*O. coriacea Lyman 1883 [incl. O. bartletti Lyman 1883] sW66
    |--Ophiotrema Koehler 1896 SW66
    |    `--*O. alberti Koehler 1896 SW66
    |--Glaciacantha Fell 1961 F61
    |    `--*G. jason Fell 1961 F61
    |--Amphipsila Verrill 1899 SW66
    |    `--*A. maculata Verrill 1899 SW66
    |--Microphiura Mortensen 1911 SW66
    |    `--*M. decipiens Mortensen 1911 SW66
    |--Ophiacanthella Verrill 1899 SW66
    |    `--*O. troscheli (Lyman 1878) [=Ophiacantha troscheli] SW66
    |--Ophialcaea Verrill 1899 SW66
    |    `--*O. tuberculosa (Lyman 1878) [=Ophiacantha tuberculosa] SW66
    |--Ophiambix Lyman 1880 SW66
    |    `--*O. aculeatus Lyman 1880 SW66
    |--Ophientrema Verrill 1899 SW66
    |    `--*O. scolopendrica (Lyman 1883) [=Ophiacantha scolopendrica] SW66
    |--Ophioblenna Lütken 1859 SW66
    |    `--*O. antillensis Lütken 1859 SW66
    |--Ophiochondrella Verrill 1899 SW66
    |    `--*O. squamosa (Lyman 1883) [=Ophiochondrus squamosus] SW66
    |--Ophiocopa Lyman 1883 SW66
    |    `--*O. spatula Lyman 1883 SW66
    |--Ophiodaces Koehler 1922 SW66
    |    `--*O. inanis Koehler 1922 SW66
    |--Ophiodelos Koehler 1931 SW66
    |    `--*O. insignis Koehler 1931 SW66
    |--Ophiodictys Koehler 1922 SW66
    |    `--*O. uncinatus Koehler 1922 SW66
    |--Ophiogema Koehler 1922 SW66
    |    `--*O. punctata Koehler 1922 SW66
    |--Ophioglyphoida Chapman 1934 SW66
    |    `--*O. fosteri (Chapman 1934) [=Ophiacantha (*Ophioglyphoida) fosteri] SW66
    |--Ophiolebes Lyman 1878 SW66
    |    `--*O. scorteus Lyman 1878 SW66
    |--Ophiologimus Clark 1911 SW66
    |    `--*O. hexactis Clark 1911 SW66
    |--Ophiomelina Koehler 1922 SW66
    |    `--*O. placida (Koehler 1904) [=Ophiomitrella placida] SW66
    |--Ophiomitra Lyman 1869 SW66
    |    `--*O. valida Lyman 1869 SW66
    |--Ophiomytis Koehler 1904 SW66
    |    `--*O. weberi Koehler 1904 SW66
    |--Ophiophrura Clark 1911 SW66
    |    `--*O. liodisca Clark 1911 SW66
    |--Ophiopinna Hess 1960 SW66
    |    `--*O. elegans (Heller 1858) [=Geocoma elegans] SW66
    |--Ophioplinthaca Verrill 1899 SW66
    |    `--*O. dipsacos (Lyman 1878) [=Ophiomitra dipsacos] SW66
    |--Ophioprium Clark 1915 SW66
    |    `--*O. curvicornis (Lyman 1883) [=Ophiacantha curvicornis] SW66
    |--Ophioripa Koehler 1922 SW66
    |    `--*O. marginata Koehler 1922 SW66
    |--Ophiosparte Koehler 1922 SW66
    |    `--*O. gigas Koehler 1922 SW66
    |--Ophiothauma Clark 1938 SW66
    |    `--*O. heptactis Clark 1938 SW66
    |--Ophiotholia Lyman 1880 SW66
    |    `--*O. supplicans Lyman 1880 SW66
    |--Ophiomitrella Verrill 1899 SW66
    |    |--*O. laevipellis (Lyman 1883) [=Ophiacantha laevipellis] SW66
    |    `--O. clavigera BK77
    |--Ophiomyces Lyman 1869 SW66
    |    |--*O. frutectosus Lyman 1869 SW66
    |    `--O. delata Koehler 1904 MG-H11
    |--Ophiophthalmus Matsumoto 1917 [incl. Ophiosemnotes Matsumoto 1917] SW66
    |    |--*O. cataleimmoidus (Clark 1911) [=Ophiacantha cataleimmoida] SW66
    |    `--O. relictus (Koehler 1904) MG-H11
    |--Ophiothamnus Lyman 1869 [incl. Ophioleda Koehler 1906] SW66
    |    |--*O. vicarius Lyman 1869 SW66
    |    `--O. biocal O’Hara & Stöhr 2006 MG-H11
    |--Ophiurothamnus Matsumoto 1917 SW66
    |    |--*O. dicylus (Clark 1911) [=Ophiomitra dicyla] SW66
    |    `--O. clausa (Lyman 1878) MG-H11
    |--Ophioplinthaca MG-H11
    |    |--O. plicata (Lyman 1878) MG-H11
    |    `--O. rudis (Koehler 1897) MG-H11
    |--Amphilimna Verrill 1899 MG-H11, SW66
    |    |--*A. olivacea (Lyman 1869) [=Ophiocnida olivacea] SW66
    |    |--A. granulosa Liao 1989 MG-H11
    |    `--A. transacta (Koehler 1930) MG-H11
    |--Ophiotreta Verrill 1899 MG-H11, SW66
    |    |--O. eximia (Koehler 1904) MG-H11
    |    |--O. stimulea (Lyman 1878) MG-H11
    |    `--O. valenciennesi (Lyman 1879) MG-H11
    `--Ophiacantha Müller & Troschel 1842 (see below for synonymy) SW66
         |--O. bidentata (Retzius 1805) [=Asterias bidentata; incl. *O. spinulosa Müller & Troschel 1842] SW66
         |--O. abyssicola K-MC02
         |--O. alternata Koehler 1907 MG-H11
         |--O. anomala BK77
         |--O. brachygnatha Clark 1928 MG-H11
         |--O. clavigera Clark 1966 MG-H11
         |--O. dallasi (Duncan 1879) MG-H11
         |--O. disjuncta (Koehler 1911) LR06
         |--O. indica Ljungman 1867 MG-H11
         |--O. pacata Koehler 1922 MG-H11
         |--O. rosea Lyman 1878 MG-H11
         `--O. spectabilis Sars 1871 MG-H11

Ophiacantha Müller & Troschel 1842 [incl. Ophiectodia Verrill 1899, Ophientodia Verrill 1899, Ophiodiplax Koehler 1911, Ophiopristis Verrill 1899, Ophioscalus Verrill 1899] SW66

*Type species of generic name indicated

REFERENCES

[BK77] Barel, C. D. N., & P. G. N. Kramers. 1977. A survey of the echinoderm associates of the north-east Atlantic area. Zoologische Verhandelingen 156: 1–159.

[F61] Fell, H. B. 1961. New genera and species of Ophiuroidea from Antarctica. Transactions of the Royal Society of New Zealand 88 (4): 839–841.

[K-MC02] Klein-MacPhee, G., & B. B. Collette. 2002. Scorpionfishes. Family Scorpaenidae. In: Collette, B. B., & G. Klein-MacPhee (eds) Bigelow and Schroeder’s Fishes of the Gulf of Maine 3^rd ed. pp. 331–338. Smithsonian Institute Press: Washington.

[LR06] Lanterbecq, D., G. W. Rouse, M. C. Milinkovitch & I. Eeckhaut. 2006. Molecular phylogenetic analyses indicate multiple independent emergences of parasitism in Myzostomida (Protostomia). Systematic Biology 55 (2): 208–227.

[MG-H11] McEnnulty, F. R., K. L. Gowlett-Holmes, A. Williams, F. Althaus, J. Fromont, G. C. B. Poore, T. D. O’Hara, L. Marsh, P. Kott, S. Slack-Smith, P. Alderslade & M. V. Kitahara. 2011. The deepwater megabenthic invertebrates on the western continental margin of Australia (100–1100 m depths): composition, distribution and novelty. Records of the Western Australian Museum Supplement 80: 1–191.

[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.

Ophiodermatidae

Ophiopeza spinosa, copyright Moorea Biocode.

Belongs within: Ophiurida.

The Ophiodermatidae are a group of brittle stars with the relatively stout arms inserted laterally and firmly fused with the disc (Spencer & Wright 1966).

Characters (from Spencer & Wright 1966): Arms inserted laterally and firmly fused with disc, moderately long, stout, widest at base. Granules cover disc scales of both surfaces and commonly jaws also. Unpaired infradental papillae at apex of each jaw.

Ophiodermatidae [Ophiarachnidae]
    |--Cryptopelta Clark 1909 [incl. Ophiodyscrita Clark 1938] SW66
    |    `--*C. aster (Lyman 1879) [=Ophiopeza aster] SW66
    |--Ophiarachna Müller & Troschel 1842 SW66
    |    `--*O. incrassata (Lamarck 1816) [=Ophiura incrassata] SW66
    |--Ophiochasma Grube 1868 [incl. Ophiopinax Bell 1884] SW66
    |    `--O. stellata (Ljungman 1867) [=Ophiarachna stellata; incl. *Ophiochasma adspersa Grube 1868] SW66
    |--Ophiopsammus Lütken 1869 MG-H11, SW66
    |    `--O. assimilis (Bell 1888) MG-H11
    |--Bathypectinura Clark 1909 SW66
    |    `--*B. lacertosa (Lyman 1883) [=Pectinura lacertosa] SW66
    |--Diopederma Clark 1913 SW66
    |    `--*D. daniana (Verrill 1867) [=Ophiura daniana] SW66
    |--Distichophis Ely 1942 SW66
    |    `--*D. clarki Ely 1942 SW66
    |--Ophiochaeta Lütken 1869 SW66
    |    `--*O. hirsuta Lütken 1869 SW66
    |--Ophioclastus Murakami 1943 SW66
    |    `--*O. hataii Murakami 1943 SW66
    |--Ophiocormus Clark 1915 [incl. Ophiostegastus Murakami 1944] SW66
    |    `--*O. notabilis Clark 1915 SW66
    |--Ophiocryptus Clark 1915 SW66
    |    `--*O. maculosus Clark 1915 SW66
    |--Ophioncus Ives 1889 SW66
    |    `--*O. granulosus Ives 1889 SW66
    |--Ophiopaepale Ljungman 1871 SW66
    |    `--*O. goesiana Ljungman 1871 SW66
    |--Ophiopetra Hess 1962 SW66
    |    `--*O. lithographica Hess 1962 SW66
    |--Ophiopezella Ljungman 1871 SW66
    |    `--*O. spinosa (Ljungman 1867) [=Ophiarachna spinosa] SW66
    |--Ophiurochaeta Matsumoto 1915 SW66
    |    `--*O. mixta (Lyman 1878) [=Ophiochaeta mixta] SW66
    |--Ophiuroconis Matsumoto 1915 SW66
    |    `--*O. monolepis Matsumoto 1915 SW66
    |--Ophiurodon Matsumoto 1915 SW66
    |    `--*O. grandisquama (Koehler 1904) [=Ophioconis grandisquama] SW66
    |--Schizoderma Nielsen 1932 SW66
    |    `--*S. diplax Nielsen 1932 SW66
    |--Toporkovia Djakonov 1954 SW66
    |    `--*T. fragilis Djakonov 1954 SW66
    |--Ophioderma Müller & Troschel 1840 SW66
    |    |--*O. longicauda (Retzius 1805) [=Asterias longicauda] SW66
    |    `--O. brevispinum F67
    |--Pectinura Forbes 1843 SW66
    |    |--*P. vestita Forbes 1843 SW66
    |    `--P. cylindrica F67
    |--Ophioconis Lütken 1869 SW66
    |    |--*O. forbesi (Heller 1862) [=Pectinura forbesi] SW66
    |    `--O. opacum (Clark 1928) MG-H11
    |--Ophiarachnella Ljungman 1872 SW66
    |    |--*O. gorgonia (Müller & Troschel 1842) [=Ophiarachna gorgonia] SW66
    |    `--O. septemspinosa (Müller & Troschel 1842) BM09
    |--Palaeocoma d’Orbigny 1850 SW66
    |    |--*P. milleri (Phillips 1829) [=Ophiura milleri] SW66
    |    `--P. escheri SW66
    `--Ophiopeza Peters 1851 SW66
         |--*O. fallax Peters 1851 SW66
         |--O. cylindrica (Hutton 1872) MG-H11
         `--O. spinosa (Ljungman 1867) MG-H11

*Type species of generic name indicated

REFERENCES

[BM09] Bryce, C., & L. Marsh. 2009. Echinodermata (Asteroidea, Echinoidea and Holothuroidea) of Mermaid (Rowley Shoals), Scott and Seringapatam Reefs, Western Australia. Records of the Western Australian Museum Supplement 77: 209–220.

[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).

[MG-H11] McEnnulty, F. R., K. L. Gowlett-Holmes, A. Williams, F. Althaus, J. Fromont, G. C. B. Poore, T. D. O’Hara, L. Marsh, P. Kott, S. Slack-Smith, P. Alderslade & M. V. Kitahara. 2011. The deepwater megabenthic invertebrates on the western continental margin of Australia (100–1100 m depths): composition, distribution and novelty. Records of the Western Australian Museum Supplement 80: 1–191.

[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.

Ophiotrichidae

Common brittle stars Ophiothrix fragilis, copyright Bernard Picton.

Belongs within: Gnathophiurina.

The Ophiotrichidae are a family of stout-toothed brittle stars.

Characters (from Spencer & Wright 1966): Teeth stout, quadrangular, spiniform tooth papillae clustered at apex of each jaw. No oral papillae.

Ophiotrichidae
    |--Dougaloplus BM09
    |--Ophiomaza Lyman 1871 [incl. Luetkenia Brock 1888 nec Claus 1864 nec Duncan 1878] SW66
    |    `--*O. cacaotica Lyman 1871 SW66
    |--Ophiopteron Ludwig 1888 SW66
    |    `--*O. elegans Ludwig 1888 SW66
    |--Amphiophiothrix Clark 1946 SW66
    |    `--*A. demessa (Lyman 1861) [=Ophiothrix demessa] SW66
    |--Gymnolophus Brock 1888 SW66
    |    `--*G. holdsworthi (Smith 1878) [=Ophiothela holdsworthi] SW66
    |--Lissophiothrix Clark 1938 SW66
    |    `--*L. delicata Clark 1938 SW66
    |--Ophioaethiops Brock 1888 [incl. Ophiohelix Koehler 1895] SW66
    |    `--*O. unicolor Brock 1888 SW66
    |--Ophiocnemis Müller & Troschel 1842 SW66
    |    `--*O. marmorata (Lamarck 1816) [=Ophiura marmorata] SW66
    |--Ophiolophus Marktanner-Turneretscher 1887 SW66
    |    `--*O. novarae Marktanner-Turneretscher 1887 SW66
    |--Ophiopeltis Düben & Koren 1846 SW66
    |    `--*O. securigera Düben & Koren 1846 SW66
    |--Ophiophthirius Döderlein 1898 SW66
    |    `--*O. actinometrae Döderlein 1898 sW66
    |--Ophiopsammium Lyman 1874 SW66
    |    `--*O. semperi Lyman 1874 SW66
    |--Ophiosphaera Brock 1888 SW66
    |    `--*O. insignis Brock 1888 SW66
    |--Ophiotrichoides Ludwig 1882 SW66
    |    `--*O. lymani Ludwig 1882 SW66
    |--Ophiothela Verrill 1867 [incl. Ophioteresis Bell 1892] SW66
    |    |--*O. mirabilis Verrill 1867 SW66
    |    `--O. danae Verrill 1869 MG-H11
    |--Ophiogymna Ljungman 1866 [incl. Ophiocampsis Duncan 1887] SW66
    |    |--*O. elegans Ljungman 1866 SW66
    |    |--O. pellicula (Duncan 1887) BM09
    |    `--O. pulchella (Koehler 1905) MG-H11
    |--Macrophiothrix Clark 1938 [incl. Placophiothrix Clark 1938] SW66
    |    |--*M. longipeda (Lamarck 1816) [=Ophiura longipeda] SW66
    |    |--M. demessa (Lyman 1861) BM09
    |    |--M. koehleri Clark 1968 MG-H11
    |    `--M. variabilis (Duncan 1887) MG-H11
    `--Ophiothrix Müller & Troschel 1840 B01
         |--O. pentaphylla (Pennant 1777) (see below for synonymy) SW66
         |--O. aristulata Lyman 1879 MG-H11
         |--O. armata Koehler 1905 BM09
         |--O. caespitosa Lyman 1879 MG-H11
         |--O. capillaris H04
         |--O. ciliaris (Lamarck 1816) MG-H11
         |--O. exigua Lyman 1874 BM09
         |--O. foveolata Marktanner-Turneretscher 1887 MG-H11
         |--O. fragilis AB01 [incl. O. alopecurus BK77, O. echinata BK77, O. versicolor BK77]
         |--O. nereidina (Lamarck 1816) BM09
         |--O. purpurea von Martins 1867 BM09
         |--O. quinquemaculata BK77
         |--O. savignyi F67
         |--O. spiculata CH97
         |--O. spongicola Stimpson 1855 MG-H11
         `--O. viridialba von Martens 1867 MG-H11

Ophiothrix pentaphylla (Pennant 1777) [=Asterias pentaphylla; incl. Ophiura rosula Fleming 1828, *Ophiothrix rosula] SW66

*Type species of generic name indicated

REFERENCES

[AB01] Aronson, R. B., & D. B. Blake. 2001. Global climate change and the origin of modern benthic communities in Antarctica. American Zoologist 41: 27–39.

[BK77] Barel, C. D. N., & P. G. N. Kramers. 1977. A survey of the echinoderm associates of the north-east Atlantic area. Zoologische Verhandelingen 156: 1–159.

[B01] Boczarowski, A. 2001. Isolated sclerites of Devonian non-pelmatozoan echinoderms. Palaeontologia Polonica 59: 1–219.

[BM09] Bryce, C., & L. Marsh. 2009. Echinodermata (Asteroidea, Echinoidea and Holothuroidea) of Mermaid (Rowley Shoals), Scott and Seringapatam Reefs, Western Australia. Records of the Western Australian Museum Supplement 77: 209–220.

[CH97] Castro, P., & M. E. Huber. 1997. Marine Biology 2^nd ed. WCB McGraw-Hill: Boston.

[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).

[H04] Haeckel, E. 1899–1904. Kunstformen der Natur. Bibliographisches Institut: Leipzig und Wien.

[MG-H11] McEnnulty, F. R., K. L. Gowlett-Holmes, A. Williams, F. Althaus, J. Fromont, G. C. B. Poore, T. D. O’Hara, L. Marsh, P. Kott, S. Slack-Smith, P. Alderslade & M. V. Kitahara. 2011. The deepwater megabenthic invertebrates on the western continental margin of Australia (100–1100 m depths): composition, distribution and novelty. Records of the Western Australian Museum Supplement 80: 1–191.

[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.

Ophiactidae

Savigny's brittle star Ophiactis savignyi, copyright Moorea Biocode.

Belongs within: Gnathophiurina.

The Ophiactidae are a family of brittle stars in which the disc bears a covering of granules or spinules.

Characters (from Spencer & Wright 1966): Arms slender, commonly constricted at nodes. Disc with granules or spinules. Jaws with lateral oral papillae separated by gap from dissimilar infradental papillae at apex.

Ophiactidae [Ophiactinae]
    |--Histampica duplicata (Lyman 1874) MG-H11
    |--Ophiopus Ljungman 1866 [incl. Ophiaregma Sars 1872] SW66
    |    `--*O. arcticus Ljungman 1866 SW66
    |--Hemipholis Lyman 1865 SW66
    |    |--*H. elongata (Say 1825) [=Ophiura elongata] SW66
    |    `--H. cordifera H04
    |--Ophiopholis Müller & Troschel 1840 SW66
    |    |--O. aculeata (Retzius 1783) (see below for synonymy) SW66
    |    `--O. japonica H04
    `--Ophiactis Lütken 1856 [incl. Amphiactis Matsumoto 1915] SW66
         |--*O. savignyi (Müller & Troschel 1842) [=Ophiolepis savignyi; incl. *Ophiactis krebsii Lütken 1856] SW66
         |--O. abyssicola BK77
         |--O. balli BK77
         |--O. brevis Clark 1938 MG-H11
         |--O. hirta Lyman 1879 MG-H11
         |--O. macrolepidota Marktanner-Turneretscher 1887 MG-H11
         |--O. modesta Brock 1888 MG-H11
         |--O. nidarosiensis BK77
         `--O. profundi Lütken & Mortensen 1899 MG-H11

Ophiopholis aculeata (Retzius 1783) [=Asterias aculeatus; incl. Ophiolepis scolopendrica Müller & Troschel 1840, *Ophiopholis scolopendrica] SW66

*Type species of generic name indicated

REFERENCES

[BK77] Barel, C. D. N., & P. G. N. Kramers. 1977. A survey of the echinoderm associates of the north-east Atlantic area. Zoologische Verhandelingen 156: 1–159.

[H04] Haeckel, E. 1899–1904. Kunstformen der Natur. Bibliographisches Institut: Leipzig und Wien.

[MG-H11] McEnnulty, F. R., K. L. Gowlett-Holmes, A. Williams, F. Althaus, J. Fromont, G. C. B. Poore, T. D. O’Hara, L. Marsh, P. Kott, S. Slack-Smith, P. Alderslade & M. V. Kitahara. 2011. The deepwater megabenthic invertebrates on the western continental margin of Australia (100–1100 m depths): composition, distribution and novelty. Records of the Western Australian Museum Supplement 80: 1–191.

[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.

Amphiura

Amphiura filiformis, copyright Judith Oakley.

Belongs within: Gnathophiurina.

Amphiura is a cosmopolitan genus of brittle stars with the central disc covered by fine scales.

Characters (from Spencer & Wright 1966): Oral papillae not forming continuous row along jaw, having single infradental separated by gap from single outer papilla with internal one invisible above gap. Disc with fine, flat, imbricating scales. Tentacle scales 2.

Amphiura Forbes 1843 [incl. Hemilepis Ljungman 1871] SW66
    |--*A. chiajii Forbes 1843 SW66
    |--A. belgicae BK77
    |--A. borealis BK77
    |--A. dolia Clark 1938 MG-H11
    |--A. filiformis BK77
    |--A. fragilis BK77
    |--A. griegi BK77
    |--A. leptotata Clark 1915 MG-H11
    |--A. mediterranea BK77
    |--A. microplax BK77
    |--A. octacantha (Clark 1915) MG-H11
    |--A. roseus HS01
    `--A. securigera BK77

*Type species of generic name indicated

REFERENCES

[BK77] Barel, C. D. N., & P. G. N. Kramers. 1977. A survey of the echinoderm associates of the north-east Atlantic area. Zoologische Verhandelingen 156: 1–159.

[HS01] Hayward, B. W., A. B. Stephenson, M. S. Morley, W. M. Blom, H. R. Grenfell, F. J. Brook, J. L. Riley, F. Thompson & J. J. Hayward. 2001. Marine biota of Parengarenga Harbour, Northland, New Zealand. Records of the Auckland Museum 37: 45–80.

[MG-H11] McEnnulty, F. R., K. L. Gowlett-Holmes, A. Williams, F. Althaus, J. Fromont, G. C. B. Poore, T. D. O’Hara, L. Marsh, P. Kott, S. Slack-Smith, P. Alderslade & M. V. Kitahara. 2011. The deepwater megabenthic invertebrates on the western continental margin of Australia (100–1100 m depths): composition, distribution and novelty. Records of the Western Australian Museum Supplement 80: 1–191.

[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.

Asteriidae

Common starfish Asterias rubens, photographed by Jason Hall-Spencer.

Belongs within: Asteriadina.
Contains: Leptasterias.

The Asteriidae are a family of starfish with a swollen body and both straight and crossed pedicellariae. Of the three subfamilies listed below, the Pycnopodiinae has the aboral skeleton obsolescent, whereas Neomorphaster has the primary aboral plates conspicuously enlarged. The Asteriinae is composed of more generalised forms that lack the distinctive characters of the other subfamilies (Spencer & Wright 1966).

Characters (from Spencer & Wright 1966): Arms 5 to many; body swollen; ambulacrals and adambulacrals numerous, compressed; adambulacrals transversely elongate, ambulacrals narrow. Tube feet normally in 4 rows; pedicellariae straight and crossed. Aboral skeleton generally reticulate.

Asteriidae
    |  i. s.: ‘Compsaster’ spiniger (Wright 1880) B02
    |         Germanasterias BH03
    |--Neomorphaster Sladen 1889 (see below for synonymy) SW66
    |    `--*N. eustichus Sladen 1889 [incl. Stichaster talismani Perrier 1891] SW66
    |--Pycnopodiinae [Pycnopodiidae] SW66
    |    |--Pycnopodia Stimpson 1862 SW66
    |    |    `--*P. helianthoides (Brandt 1835) [=Asterias helianthoides] SW66
    |    `--Lysastrosoma Fisher 1922 SW66
    |         `--*L. anthosticta Fisher 1922 SW66
    `--Asteriinae [Coscinasteriinae, Notasteriinae, Stichasteridae] SW66
         |--Leptasterias SW66
         |--Adelasterias Koehler 1914 SW66
         |    `--*A. papillosa (Koehler 1906) [=Diplasterias papillosa] SW66
         |--Allostichaster Verrill 1914 SW66
         |    `--*A. polyplax (Müller & Troschel 1844) [=Asteracanthion polyplax] SW66
         |--Aphanasterias Fisher 1923 SW66
         |    `--*A. pycnopodia Fisher 1923 SW66
         |--Aphelasterias Fisher 1923 SW66
         |    `--*A. japonica (Bell 1881) [=Asterias japonica] SW66
         |--Astrometis Fisher 1923 SW66
         |    `--*A. sertulifera (Xantus 1860) [=Asterias sertulifera] SW66
         |--Astrostole Fisher 1923 SW66
         |    `--*A. scaber (Hutton 1872) [=Margaraster scaber] SW66
         |--Australiaster Fisher 1923 SW66
         |    `--*A. dubius (Clark 1909) [=Coscinasterias dubia] SW66
         |--Caimanaster Clark 1962 SW66
         |    `--*C. acutus Clark 1962 SW66
         |--Carlasterias da Costa 1952 [incl. Mortensenia da Costa 1941 non Döderlein 1906] SW66
         |    `--*C. lusitanica (da Costa 1941) [=Mortensenia lusitanica] SW66
         |--Cosmasterias Sladen 1889 [=Comasterias Perrier 1891; incl. Quadraster Perrier 1896] SW66
         |    `--C. lurida (Philippi 1858) (see below for synonymy) SW66
         |--Cryptasterias Verrill 1914 SW66
         |    `--*C. turqueti (Koehler 1906) [=Diplasterias turqueti] SW66
         |--Displasterias Perrier 1888 (see below for synonymy) SW66
         |    `--*D. brandti (Bell 1881) [=Asterias brandti] SW66
         |--Distolasterias Perrier 1896 SW66
         |    `--*D. stichantha (Sladen 1889) [=Asterias (Stolasterias) stichantha] SW66
         |--Evasterias Verrill 1914 SW66
         |    `--*E. troscheli (Stimpson 1862) [=Asterias troscheli] SW66
         |--Gastraster Perrier 1894 SW66
         |    `--*G. margaritaceus (Perrier 1882) [=Pedicellaster margaritaceus] SW66
         |--Granaster Perrier 1894 [incl. Hemiasterias Verrill 1914] SW66
         |    `--*G. nutrix (Studer 1885) [=Stichaster nutrix] SW66
         |--Icasterias Fisher 1923 SW66
         |    `--*I. panopla (Stuxberg 1878) [=Asterias panopla] SW66
         |--Kenrickaster Clark 1962 SW66
         |    `--*K. pedicellaris Clark 1962 SW66
         |--Lethasterias Fisher 1923 SW66
         |    `--*L. nanimensis (Verrill 1914) [=Asterias nanimensis] SW66
         |--Lysasterias Fisher 1908 [incl. Anasterias Ludwig 1903 non Perrier 1885, Paedasterias Verrill 1914] SW66
         |    `--*L. perrieri (Studer 1885) [=Anasterias perrieri] SW66
         |--Marthasterias Jullien 1878 SW66
         |    `--M. glacialis (Linné 1758) SW66 (see below for synonymy)
         |--Meyenaster Verrill 1913 SW66
         |    `--*M. gelatinosus (Meyen 1834) [=Asterias gelatinosus] SW66
         |--Neosmilaster Fisher 1930 SW66
         |    `--*N. georgianus (Studer 1885) [=Asterias georgianus] SW66
         |--Notasterias Koehler 1911 [incl. Autasterias Koehler 1911] SW66
         |    `--*N. armata Koehler 1911 SW66
         |--Orthasterias Verrill 1914 SW66
         |    `--O. koehleri (de Loriol 1897) [=Asterias koehleri; incl. *O. columbiana Verrill 1914] SW66
         |--Perissasterias Clark 1923 SW66
         |    `--*P. polyacantha Clark 1923 SW66
         |--Pisaster Müller & Troschel 1840 [incl. Calliasterias Fewkes 1889] SW66
         |    |--P. ochraceus (Brandt 1835) (see below for synonymy) SW66
         |    `--P. giganteus (Stimpson 1857) CWH77
         |--Psalidaster Fisher 1940 SW66
         |    `--*P. mordax Fisher 1940 SW66
         |--Pseudechinaster Clark 1962 SW66
         |    `--*P. rubens Clark 1962 SW66
         |--Saliasterias Koehler 1920 SW66
         |    `--*S. bracheata Koehler 1920 SW66
         |--Sclerasterias Perrier 1891 [incl. Eustolasterias Fisher 1923] SW66
         |    `--*S. guernei Perrier 1891 SW66
         |--Smilasterias Sladen 1889 [incl. Nanaster Perrier 1894] SW66
         |    `--*S. scalprifera (Sladen 1889) [=Asterias (*Smilasterias) scalprifera] SW66
         |--Stenasterias Verrill 1914 SW66
         |    `--*S. macropora (Verrill 1909) [=Asterias (Leptasterias) macropora] SW66
         |--Stephanasterias Verrill 1871 SW66
         |    `--*S. albula (Stimpson 1853) [=Asteracanthion albulus] SW66
         |--Stichaster Müller & Troschel 1840 [incl. Coelasterias Verrill 1871, Tonia Gray 1840] SW66
         |    `--*S. aurantiacus (Meyen 1834) [=Asterias aurantiaca] SW66
         |--Stichastrella Verrill 1914 SW66
         |    `--*S. rosea (Müller 1776) [=Asterias rosea] SW66
         |--Stylasterias Verrill 1914 SW66
         |    `--*S. forreri (de Loriol 1887) [=Asterias forreri] SW66
         |--Tarsastrocles Fisher 1923 SW66
         |    `--*T. verrilli (Fisher 1903) [=Hydrasterias verrilli] SW66
         |--Triplasterias Engels & Schroevers 1961 SW66
         |    `--*T. mercatoris Engels & Schroevers 1961 SW66
         |--Uniophora Gray 1840 SW66
         |    `--U. granifera (Lamarck 1816) [=Asterias granifera; incl. *U. globifera Gray 1840] SW66
         |--Urasterias Verrill 1909 SW66
         |    `--*U. linckii (Müller & Troschel 1842) [=Asteracanthion linckii] SW66
         |--Calvasterias Perrier 1875 [incl. Stichorella Koehler 1920] SW66
         |    |--*C. asterinoides Perrier 1875 SW66
         |    `--C. suteri F67
         |--Coscinasterias Verrill 1867 SW66
         |    |  i. s.: C. acutispina WOR04
         |    |         C. calamaria B64
         |    |--*C. (Coscinasterias) muricata Verrill 1867 SW66
         |    `--C. (Stolasterias Sladen 1889) [incl. Lytaster Perrier 1894, Polyasterias Perrier 1894] SW66
         |         `--C. (*S.) tenuispina (Lamarck 1816) [=Asterias tenuispina] SW66
         |--Anasterias Perrier 1875 (see below for synonymy) SW66
         |    |--*A. (Anasterias) minuta Perrier 1875 SW66
         |    |--A. (Kalyptasterias Koehler 1923) SW66
         |    |    `--A. (*K.) conferta (Koehler 1923) [=*Kalyptasterias conferta] SW66
         |    `--A. (Sporasterias Perrier 1894) SW66
         |         `--A. (S.) antarctica (Lütken 1856) (see below for synonymy) SW66
         `--Asterias Linnaeus 1758 B02 (see below for synonymy)
              |--*A. rubens Linnaeus 1758 SW66 [=Asteracanthion rubens BK77; incl. Asterias violacea BK77]
              |--A. amurensis MK05
              |--A. forbesi BK77
              |--A. gaveyi SW66
              |--A. lincki BK77
              `--A. vulgaris CH97

Anasterias Perrier 1875 [incl. Asteroderma Perrier 1888, Eremasterias Fisher 1930, Parastichaster Koehler 1920] SW66

Anasterias (Sporasterias) antarctica (Lütken 1856) [=Asteracanthion antarcticum; incl. Asterias rugispina Stimpson 1860, *Sporasterias rugispina] SW66

Asterias Linnaeus 1758 B02 [incl. Allasterias Verrill 1909 SW66, Asteracanthion Müller & Troschel 1840 SW66, Parasterias Verrill 1914 SW66, Stellonia Nardo 1834 SW66, Uraster Agassiz 1835 SW66]

Cosmasterias lurida (Philippi 1858) [=Asteracanthion luridum; incl. A. sulcifer Perrier 1869, *Cosmasterias sulcifer] SW66

Displasterias Perrier 1888 [incl. Bathyasterias Fisher 1930, Koehleraster Fisher 1922 non Lambert & Thiéry 1921, Podasterias Perrier 1894] SW66

Marthasterias glacialis (Linné 1758) SW66 [=Asterias glacialis SW66, Asteracanthion glaciale BK77; incl. *Marthasterias foliacea Jullien 1878 SW66]

Neomorphaster Sladen 1889 [=Glyptaster Sladen 1885 non Hall 1852; incl. Calycaster Perrier 1891; Neomorphasterinae] SW66

Pisaster ochraceus (Brandt 1835) [=Asterias ochraceus; incl. Asteracanthion margaritifer Müller & Troschel 1840, *Pisaster margaritifer] SW66

*Type species of generic name indicated

REFERENCES

[BK77] Barel, C. D. N., & P. G. N. Kramers. 1977. A survey of the echinoderm associates of the north-east Atlantic area. Zoologische Verhandelingen 156: 1–159.

[B64] Bennett, E. W. 1964. The marine fauna of New Zealand: Crustacea Brachyura. New Zealand Department of Scientific and Industrial Research Bulletin 153: 1–120.

[B02] Blake, D. B. 2002. Compsaster formosus Worthen & Miller (Asteroidea; Echinodermata): a Carboniferous homeomorph of the post-Paleozoic Asteriidae. Paläontologische Zeitschrift 76 (2): 357–367.

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

[CH97] Castro, P., & M. E. Huber. 1997. Marine Biology 2^nd ed. WCB McGraw-Hill: Boston.

[CWH77] Cooper, J., M. Wieland & A. Hines. 1977. Subtidal abalone populations in an area inhabited by sea otters. Veliger 20 (2): 163–167.

[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).

[MK05] Matsubara, M., M. Komatsu, T. Araki, S. Asakawa, S. Yokobori, K. Watanabe & H. Wada. 2005. The phylogenetic status of Paxillosida (Asteroidea) based on complete mitochondrial DNA sequences. Molecular Phylogenetics and Evolution 36: 598–605.

[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.

[WOR04] Waters, J. M., P. M. O’Loughlin & M. S. Roy. 2004. Molecular systematics of some Indo-Pacific asterinids (Echinodermata, Asteroidea): does taxonomy reflect phylogeny? Molecular Phylogenetics and Evolution 30: 872–878.