Mellitidae

Encope michelini, copyright Gustav Paulay.

Belongs within: Clypeasteroida.

The Mellitidae are a group of sea biscuits characterised by a series of perforations in the disc, found from the Lower Miocene to the present of tropical and warm temperate regions of the Americas (Durham 1966b).

Characters (from Durham 1966b): Medium-sized to large, flattened; petals well defined, moderately closed, outer member of pore pair greatly elongated; internal supports well developed; posterior interambulacral and paired ambulacral lunules or notches; paired interambulacra not continuous, posterior interambulacrum variable; on oral surface interambulacra widest at ambitus, about as wide as ambulacra; basicoronal plates small; periproct oral, between posterior lunule and peristome; food grooves bifurcating just outside primordial plates.

<==Mellitidae
    |--Leodia Gray 1852 D66b
    |    `--*L. sexiesperforata (Leske 1778) M89 (see below for synonymy)
    |--Mellitella Duncan 1889 D66b
    |    `--*M. stokesii (Agassiz 1841) [=Encope stokesii] M89
    |--Mellita Agassiz 1841 (nom. cons.) M89
    |    |--*M. quinquiesperforata (Leske 1778) M89 (see below for synonymy)
    |    |--M. caroliniana M89
    |    |--M. lata D66a
    |    |--M. mangrovi MGV06
    |    |--M. sexiesperforata W79
    |    `--M. tenuis K-M02
    `--Encope Agassiz 1840 (see below for synonymy) D66b
         |--*E. grandis Agassiz 1840 M89
         |--E. aberrans M89
         |--‘Scutella’ cassidulina Desmoulins 1837 (see below for synonymy) D66b
         |--E. emarginata (Leske 1778) [=Echinodiscus emarginatus, *Echinoglyphus emarginatus, Scutella emarginata] D66b
         |--‘Scutella’ macrophora Ravenel 1843 [=*Macrophora macrophora, *Ravenellia macrophora] D66b
         |--E. michelini K-M02
         |--E. micropora M89
         `--E. wetmorei D66b

Encope Agassiz 1840 [incl. Desmoulinaster Lambert & Thiéry 1914, Echinoglyphus Gray 1852, Echinoglycus (l. c.) non Leske 1778, Macrophora Conrad 1865, Moulinia Agassiz 1841, Moulinsia Agassiz 1847 non Grateloup 1841, Ravenellia Lütken 1864] D66b

*Leodia sexiesperforata (Leske 1778) M89 [=Echinodiscus sexiesperforatus M89; incl. Echinus hexaporus Gmelin 1789 D66b, Leodia richardsoni Gray 1852 D66b, *Scutella sexforis Lamarck 1816 D66b]

*Mellita quinquiesperforata (Leske 1778) M89 [=Echinodiscus quinquiesperforatus M89; incl. M. testudinata Klein 1734 (pre-Linnean) D66b]

‘Scutella’ cassidulina Desmoulins 1837 [=*Desmoulinaster cassidulina, *Moulinia cassidulina, *Moulinsia cassidulina] D66b

*Type species of generic name indicated

REFERENCES

[D66a] Durham, J. W. 1966a. Echinoids—ecology and paleoecology. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt U. Echinodermata 3 vol. 1 pp. U257–U265. The Geological Society of America, Inc., and The University of Kansas Press.

[D666b] Durham, J. D. 1966b. Clypeasteroids. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt U. Echinodermata 3 vol. 2 pp. U450–U491. The Geological Society of America, Inc., and The University of Kansas Press.

[K-M02] Klein-MacPhee, G. 2002. Triggerfishes. Family Balistidae. In: Collette, B. B., & G. Klein-MacPhee (eds) Bigelow and Schroeder’s Fishes of the Gulf of Maine 3^rd ed. pp. 590–592. Smithsonian Institute Press: Washington.

[MGV06] Mai, A. C. G., A. M. Garcia & J. P. Vieira. 2006. Ecologia alimentar do barrigudinho Jenynsia multidentata (Jenyns, 1842) (Pisces: Cyprinodontiformes) no estuário da Laguna dos Patos, Rio Grande do Sul, Brasil. Comunicações do Museo de Ciências e Tecnologia da PUCRS, Serie Zoologia 19 (1): 3–18.

[M89] Mooi, R. 1989. Living and fossil genera of the Clypeasteroida (Echinoidea: Echinodermata): an illustrated key and annotated checklist. Smithsonian Contributions to Zoology 488: 1–51.

[W79] Warner, G. F. 1979. Aggregation in echinoderms. In: Larwood, G., & B. R. Rosen (eds) Biology and Systematics of Colonial Organisms pp. 375–396. Academic Press: London.

Neolaganidae

(Left to right) lateral, aboral, oral and diagrammatic oral views of Neolaganum archerensis, from Durham (1966).

Belongs within: Clypeasteroida.

The Neolaganidae are a group of sand dollars known from the Eocene and Oligocene of the Gulf of Mexico and the Caribbean (Durham 1966).

Characters (from Durham 1966): Usually with pseudocompound plates in petals, outer member of pore pairs greatly elongated, basicoronal plates in regular pentagon with ambulacral plates at apices, first pair of coronal plates markedly larger than succeeding plates, terminal adapical interambulacral plate rectangular.

<==Neolaganidae D66
    |--Neorumphia Durham 1954 D66
    |    `--*N. elegans (Sánchez Roig 1949) [=Rumphia elegans] M89
    |--Sanchezella Durham 1954 D66
    |    `--*S. sanchezi (Lambert 1926) [=Jacksonaster sanchezi] M89
    |--Weisbordella Durham 1954 D66
    |    `--*W. caribbeana (Weisbord 1934) [=Peronella caribbeana] M89
    |--Wythella Durham 1954 D66
    |    `--*W. eldridgei (Twitchell 1915) [=Laganum eldridgei] M89
    |--Cubanaster Sánchez Roig 1952 D66
    |    |--*C. torrei (Lambert in Sánchez Roig 1962) [=Jacksonaster torrei] M89
    |    `--C. acunai M89
    `--Neolaganum Durham 1954 M89
         |--*N. archerensis (Twitchell 1915) [=Laganum archerensis] M89
         |--N. dalli M89
         `--N. durhami M89

*Type species of generic name indicated

REFERENCES

[D66] Durham, J. D. 1966. Clypeasteroids. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt U. Echinodermata 3 vol. 2 pp. U450–U491. The Geological Society of America, Inc., and The University of Kansas Press.

[M89] Mooi, R. 1989. Living and fossil genera of the Clypeasteroida (Echinoidea: Echinodermata): an illustrated key and annotated checklist. Smithsonian Contributions to Zoology 488: 1–51.

Echinarachniidae

Aboral and diagrammatic oral view of Vaquerosella andersoni, from Durham (1966).

Belongs within: Clypeasteroida.

The Echinarachniidae are a group of sand dollars known from the Oligocene to the present of the North Pacific, as well as the present day north-western North Atlantic (Durham 1966).

Characters (from Durham 1966): Medium-sized to large; petals well developed, anterior petal more open than paired petals; interambulacrum 5 may be discontinuous; interambulacra 0.7 or less width of ambulacra at ambitus; four genital pores; periproct marginal to inframarginal; food grooves with central trunk.

<==Echinarachniidae D66
    |--Echinarachnius Gray 1825 (nom. cons.) M89 [=Phelsumaster Lambert & Thiéry 1914 D66, Phelsumia Pomel 1883 D66]
    |    `--*E. parma (Lamarck 1816) M89 [=Scutella parma M89, *Phelsumaster parma D66, *Phelsumia parma D66]
    |--Kewia Nisiyama 1935 D66
    |    `--*K. blancoensis (Kew 1920) [=Scutella blancoensis] M89
    |--Nipponaster Durham 1952 [=Pseudoastrodapsis Durham 1953] D66
    |    `--*N. nipponicus (Nisiyama 1934) M89 [=Astrodapsis nipponicus M89, *Pseudoastrodapsis nipponicus D66]
    |--Tenuiarachnius Durham 1955 D66
    |    `--*T. kleinpelli (Grant & Hertlein 1938) M89 (see below for synonymy)
    |--Remondella Durham 1955 D66
    |    |--*R. gabbi (Rémond 1863) [=Clypeaster gabbi] M89
    |    `--R. waldroni M89
    |--Astrodapsis Conrad 1856 [=Asterodapsis Agassiz 1872, Astrodaspis (l. c.)] D66
    |    |--*A. antiselli Conrad 1856 M89 [=*Asterodapsis antiselli D66]
    |    |--A. brewerianus D66
    |    `--A. spatiosus M89
    `--Vaquerosella Durham 1955 D66
         |--*V. andersoni (Twitchell 1915) [=Scutella andersoni] M89
         |--V. norrisi M89
         `--V. vaquerosensis D66

*Tenuiarachnius kleinpelli (Grant & Hertlein 1938) M89 [=Echinarachnius gabbii kleinpelli M89; incl. Scutella gabbi tenuis Kew 1915 non Echinarachnius tenuis Yoshikawa 1898 D66]

*Type species of generic name indicated

REFERENCES

[D66] Durham, J. D. 1966. Clypeasteroids. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt U. Echinodermata 3 vol. 2 pp. U450–U491. The Geological Society of America, Inc., and The University of Kansas Press.

[M89] Mooi, R. 1989. Living and fossil genera of the Clypeasteroida (Echinoidea: Echinodermata): an illustrated key and annotated checklist. Smithsonian Contributions to Zoology 488: 1–51.

Scutellidae

Scutella subrotunda, copyright Parent Géry.

Belongs within: Clypeasteroida.

The Scutellidae are a group of sand dollars known from the Oligocene and Miocene (Durham 1966).

Characters (from Durham 1966): Medium-sized to large, flattened, internal supports well developed; petals closed, outer member of pore pair subdivided; interambulacra continuous, usually as wide as ambulacra at ambitus; primordial ambulacral and interambulacral plates about equal; four genital pores; periproct on oral surface; food grooves bifurcating close to peristome.

<==Scutellidae D66
    |--Parascutella Durham 1953 D66
    |    |--*P. leognanensis (Lambert 1903) [=Scutella leognanensis] M89
    |    `--P. bonali M89
    |--Parmulechinus Lambert 1910 [incl. Stenaster Lambert 1905 non Billings 1858] D66
    |    |--*P. agassizi (Oppenheim 1902) M89 (see below for synonymy)
    |    `--P. subtetragona M89
    `--Scutella Lamarck 1816 M89 [incl. Lambertiella Checchia-Rispoli 1917 D66]
         |--*S. subrotunda (Leske 1778) [=Echinodiscus subrotundus] M89
         `--S. vindobonensis D66
              |--S. v. vindobonensis D66
              `--S. v. secunda D66

*Parmulechinus agassizi (Oppenheim 1902) M89 [=Scutella agassizi M89; incl. *Stenaster labrei Lambert 1905 D66, Sc. striatula Agassiz 1841 non de Serres 1829 D66]

*Type species of generic name indicated

REFERENCES

[D66] Durham, J. D. 1966. Clypeasteroids. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt U. Echinodermata 3 vol. 2 pp. U450–U491. The Geological Society of America, Inc., and The University of Kansas Press.

[M89] Mooi, R. 1989. Living and fossil genera of the Clypeasteroida (Echinoidea: Echinodermata): an illustrated key and annotated checklist. Smithsonian Contributions to Zoology 488: 1–51.

Holasteroida

Lateral and oral views of Echinocorys scutatus, from Wagner & Durham (1966).

Belongs within: Atelostomata.

The Holasteroida are a diverse group of mostly deep-water irregular echinoids known from the Lower Jurassic to the present. They exhibit a wide variety of forms; the primarily bathyal Pourtalesiidae are among the most unusual of all echinoids, with a more or less elongate, bottle-shaped test (Wagner & Durham 1966).

Characters (from Wagner & Durham 1966): Apical system typically elongate or disjunct, no genital 5; plastron feebly differentiated or meridosternous; petals not always differentiated, paired petals typically not impressed; no floscelle; apical system and peristome may be opposite to one another; fascioles variable.

<==Holasteroida
    |--Calymne Thomson 1877 [Calymnidae] WD66
    |    `--*C. relicta Thomson 1877 WD66
    |--Stenonaster Lambert 1922 [=Stenocorys Lambert & Thiéry 1917, Stenonia Desor 1858 non Gray 1853] WD66
    |    `--*S. tuberculata (Defrance 1816) (see below for synonymy) WD66
    |--Somaliasteridae WD66
    |    |--Somaliaster Hawkins 1935 WD66
    |    |    `--*S. magniventer Hawkins 1935 WD66
    |    |--Brightonia Kier 1957 WD66
    |    |    `--*B. macfadyeni Kier 1957 WD66
    |    |--Iraniaster Cotteau & Gauthier in Morgan 1895 WD66
    |    |    `--*I. morgani Cotteau & Gauthier in Morgan 1895 WD66
    |    `--Leviechinus Kier 1957 WD66
    |         `--*L. gregoryi (Currie 1927) [=Pericosmus gregoryi] WD66
    |--Urechinidae WD66
    |    |--Chelonechinus Bather 1934 WD66
    |    |    `--*C. suvae Bather 1934 WD66
    |    |--Pilematechinus Agassiz 1904 WD66
    |    |    `--*P. rathbuni (Agassiz 1898) [=Cystechinus rathbuni] WD66
    |    |--Plexechinus Agassiz 1896 WD66
    |    |    `--*P. cinctus Agassiz 1896 WD66
    |    |--Sanchezaster Lambert in Sánchez Roig 1924 WD66
    |    |    `--*S. habanensis Lambert in Sánchez Roig 1924 WD66
    |    |--Sternopatagus de Maijere 1902 WD66
    |    |    `--*S. sibogae de Meijere 1902 WD66
    |    `--Urechinus Agassiz 1879 [incl. Cystechinus Agassiz 1879] WD66
    |         |--*U. naresianus Agassiz 1879 WD66
    |         `--‘*Cystechinus’ wyvillei Agassiz 1879 WD66
    |--Pourtalesiidae WD66
    |    |--Ceratophysa Pomel 1883 [=Rodocystis Lambert & Thiéry 1924] WD66
    |    |    `--*C. ceratopyga (Agassiz 1879) [=*Rodocystis ceratopyga] WD66
    |    |--Cystocrepis Mortensen 1907 WD66
    |    |    `--*C. setigera (Agassiz 1898) [=Echinocrepis setigera] WD66
    |    |--Echinocrepis Agassiz 1879 WD66
    |    |    `--*E. cuneata Agassiz 1879 WD66
    |    |--Echinosigra Mortensen 1907 WD66
    |    |    `--*E. phiale (Thomson 1873) [=Pourtalesia phiale] WD66
    |    |--Helgocystis Mortensen 1907 WD66
    |    |    `--*H. carinata (Agassiz 1879) [=Pourtalesia carinata] WD66
    |    |--Spatagocystis Agassiz 1879
    |    |    `--*S. challengeri Agassiz 1879 WD66
    |    `--Pourtalesia Agassiz 1869 [incl. Phyale Pomel 1883 non Koch 1847, Phyalopsis Pomel 1883] WD66
    |         |--*P. miranda Agassiz 1869 WD66
    |         |--P. jeffreysi Thomson 1873 [=*Phyale jeffreysi] WD66
    |         `--P. laguncula Agassiz 1879 [=*Phyalopsis laguncula] WD66
    |--Collyritidae WD66
    |    |--Cardiopelta Pomel 1883 WD66
    |    |    `--*C. trigonalis (Desor in Desor & de Loriol 1872) [=Collyrites trigonalis] WD66
    |    |--Cyclolampas Pomel 1883 WD66
    |    |    `--*C. voltzii (Agassiz 1839) [=Disaster voltzii] WD66
    |    |--Grasia Michelin 1854 WD66
    |    |    `--*G. elongata (Gras 1852) [=Hyboclypus elongata] WD66
    |    |--Orbignyana Ebray 1860 [=Spatoclypus Pomel 1883] WD66
    |    |    `--*O. ebrayi (Cotteau 1873) [=Collyrites ebrayi, *Spatoclypus ebrayi] WD66
    |    |--Proholaster Gauthier 1896 WD66
    |    |    `--*P. auberti Gauthier 1896 WD66
    |    |--Pygomalus Pomel 1883 WD66
    |    |    `--*P. ovalis (Leske 1778) [=Spatangites ovalis] WD66
    |    |--Pygorhytis Pomel 1883 WD66
    |    |    `--*P. ringens (Agassiz 1836) [=Disaster ringens] WD66
    |    `--Collyrites Desmoulins 1835 WD66
    |         |--*C. elliptica (Lamarck 1816) [=Ananchytes elliptica] WD66
    |         |--C. bicordata [=Cardiopelta bicordata; incl. Co. bicordata var. baltica Beurlen 1934] B01
    |         `--C. dorsalis F66
    |--Disasteridae WD66
    |    |--Disaster Agassiz 1836 [=Dysaster Agassiz 1839] WD66
    |    |    `--*D. granulosus (Goldfuss 1826) [=Nucleolites granulosus, *Dysaster granulosus] WD66
    |    |--Acrolusia Lambert 1920 WD66
    |    |    `--*A. gauthieri Lambert 1920 WD66
    |    |--Cardiolampas Pomel 1883 WD66
    |    |    `--*C. friburgensis (Ooster 1865) [=Collyrites friburgensis] WD66
    |    |--Corthya Pomel 1883 WD66
    |    |    `--*C. hemisphaerica (Gras 1848) [=Disaster hemisphaericus] WD66
    |    |--Dialyaster Pomel 1883 WD66
    |    |    `--*D. gueymardi (Gras 1848) [=Metaporinus gueymardi] WD66
    |    |--Oustechinus Lambert 1931 WD66
    |    |    `--*O. bassae Lambert 1931 WD66
    |    |--Tithonia Pomel 1883 WD66
    |    |    `--*T. convexus (Catullo 1827) [=Nucleolites convexus] WD66
    |    |--Collyropsis Gauthier 1896 [incl. Procollyropsis Beurlen 1934] WD66
    |    |    |--*C. carinatus (Leske 1778) [=Spatangites carinatus] WD66
    |    |    `--‘Disaster’ platypygus Quenstedt 1874 [=*Procollyropsis platypygus] WD66
    |    `--Metaporinus Agassiz 1844 (see below for synonymy) WD66
    |         |--*M. michelini Agassiz 1844 WD66
    |         `--‘Collyrites’ censoriensis Cotteau 1849-1856 [=*Perioxus censoriensis] WD66
    `--Holasteridae WD66
         |--Aurelianaster Lambert & Thiéry 1925 (see below for synonymy) WD66
         |    `--*A. valettei (Lambert in Vallette 1913) [=*Leiocorys valettei, *Vallettaster valettei] WD66
         |--Basseaster Lambert 1936 WD66
         |    `--*B. rostratus Lambert 1936 WD66
         |--Cardiaster Forbes 1850 WD66
         |    `--C. granulosus (Goldfuss 1826) (see below for synonymy) WD66
         |--Cardiotaxis Lambert 1917 WD66
         |    `--*C. peroni (Lambert 1887) [=Cardiaster peroni] WD66
         |--Cibaster Pomel 1883 WD66
         |    `--*C. bourgeoisanus d’Orbigny 1853) [=Cardiaster bourgeoisanus] WD66
         |--Duncaniaster Lambert 1896 WD66
         |    `--*D. australiae (Duncan 1887) [=Holaster australiae] WD66
         |--Entomaster Gauthier 1888 WD66
         |    `--*E. rousseli Gautheir 1888 WD66
         |--Galeaster Seunes 1889 WD66
         |    `--*G. bertrandi Seunes 1889 WD66
         |--Ganbirretia Gauthier 1903 WD66
         |    `--*G. douvillei Gauthier 1903 WD66
         |--Garumnaster Lambert 1907 WD66
         |    `--*G. michaleti Lambert 1907 WD66
         |--Guettaria Gauthier 1888 WD66
         |    `--*G. angladei Gauthier 1888 WD66
         |--Hagenowia Duncan 1889 [=Hagenovia (l. c.), Martinosigra Nielson 1942] WD66
         |    `--*H. rostrata (Forbes 1889) [=Cardiaster rostratus, *Martinosigra rostrata] WD66
         |--Hemipneustes Agassiz 1836 WD66
         |    `--H. striatoradiatus (Leske 1778) (see below for synonymy) WD66
         |--Infulaster Desor 1858 WD66
         |    `--I. excentricus (Woodward 1833) (see below for synonymy) WD66
         |--Ismidaster Boehm 1927 WD66
         |    `--*I. toulai Boehm 1927 WD66
         |--Jeronia Seunes 1888 WD66
         |    `--*J. pyrenaica Seunes 1888 WD66
         |--Labrotaxis Casey 1960 WD66
         |    `--*L. cantianus (Casey 1960) [=Holaster (*Labrotaxis) cantianus] WD66
         |--Lampadaster Cotteau 1889 WD66
         |    `--*L. gradidieri Cotteau 1889 WD66
         |--Lampadocorys Pomel 1883 WD66
         |    `--*L. sulcatus (Cotteau 1873) [=Holaster sulcatus] WD66
         |--Messaoudia Lambert 1917 WD66
         |    `--*M. pyriformis (Peron & Gauthier 1878) [=Holaster pyriformis] WD66
         |--Offaster Desor 1858 WD66
         |    `--*O. pilula (Lamarck 1816) [=Ananchytes pilula] WD66
         |--Opisopneustes Gauthier 1889 WD66
         |    `--*O. cossoni Gauthier 1889 WD66
         |--Paronaster Airaghi 1906 WD66
         |    `--*P. cupuliformis Airaghi 1906 WD66
         |--Pseudholaster Pomel 1883 WD66
         |    `--*P. bicarinatus (Agassiz 1847) [=Holaster bicarinatus] WD66
         |--Pseudoffaster Lambert 1924 WD66
         |    `--*P. caucasicus (Dru 1884) [=Holaster caucasicus] WD66
         |--Rispolia Lambert 1917 WD66
         |    `--*R. subtrigonata (Catullo 1827) [=Nucleolites subtrigonatus] WD66
         |--Scagliaster Munier-Chalmas 1891 WD66
         |    `--*S. concavus (Catullo 1827) [=Ananchytes concavus] WD66
         |--Stereopneustes de Meijere 1902 WD66
         |    `--*S. relictus de Meijere 1902 WD66
         |--Taphraster Pomel 1883 WD66
         |    `--*T. campicheanus [=Holaster campicheanus] WD66
         |--Tholaster Seunes 1890 [=Gibbaster Seunes 1889 non Gauthier 1887] WD66
         |    `--*T. munieri (Seunes 1889) [=*Gibbaster munieri] WD66
         |--Titanaster Szörényi 1929 WD66
         |    `--*T. labiostoma Szörényi 1929 WD66
         |--Zumoffenia Fourtau 1912 WD66
         |    `--*Z. ludovici Fourtau 1912 WD66
         |--Pseudananchys Pomel 1883 [incl. Craginaster Lambert 1903] WD66
         |    |--*P. algiris (Coquand 1862) [=Ananchytes algiris] WD66
         |    `--‘Holaster’ completa Cragin 1893 [=*Craginaster completa] WD66
         |--Sternotaxis Lambert 1893 [incl. Plesiocorys Pomel 1883] WD66
         |    |--*S. planus (Mantell 1822) [=Spatangus planus] WD66
         |    `--‘Holaster’ placenta Agassiz in Agassiz & Desor 1847 [=*Plesiocorys placenta] WD66
         |--Toxopatagus Pomel 1883 [incl. Heteropneustes Pomel 1883] WD66
         |    |--*T. italicus (Manzoni 1878) [=Hemipneustes italicus] WD66
         |    `--‘*Heteropneustes’ deletrei Pomel 1883 WD66
         |--Stegaster Pomel 1883 [incl. Seunaster Lambert in Blayac 1912, Synochitis Lambert 1917] WD66
         |    |--*S. gillieroni (de Loriol 1873) [=Cardiaster gillieroni] WD66
         |    |--‘Holaster’ bouillei Cotteau in de Bouille 1873 [=*Seunaster bouillei] WD66
         |    `--S. cotteaui WD66
         |--Holaster Agassiz 1836 [incl. Ananchothuria Fossa-Mancini 1919, Holasteropsis Elbert 1902] WD66
         |    |--*H. nodulosus (Goldfuss 1829) [=Spatangus nodulosus] WD66
         |    |--‘*Holasteropsis’ credneriana Elbert 1902 WD66
         |    |--H. planus K79
         |    |--H. subglobosus K79
         |    `--‘*Ananchothuria’ tesselata Fossa-Mancini 1919 WD66
         `--Echinocorys Leske 1778 (see below for synonymy) WD66
              |--*E. scutatus Leske 1778 (see below for synonymy) WD66
              |--‘Anachytes’ corculum Goldfuss 1826 [=*Corculum corculum] WD66
              |--‘*Oolaster’ mattseensis Laube 1869 WD66
              |--‘*Ananchytes’ ovatus Lamarck 1816 WD66
              `--‘*Galeola’ papillosa Quenstedt 1874 WD66

Holasteroida incertae sedis:
  Coraster Cotteau 1886 WD66
    `--*C. vilanovae Cotteau 1886 WD66
  Corechinus Kongiel 1936 WD66
    `--*C. pulaviensis Kongiel 1936 WD66
  Habanaster Lambert in Sánchez Roig 1924 WD66
    `--*H. sanchezi Lambert in Sánchez Roig 1924 WD66
  Menuthiaster Lambert 1896 WD66
    `--*M. cotteaui Lambert 1896 WD66
  Nordenskjoeldaster Lambert 1910 WD66
    `--*N. antarcticus Lambert 1910 WD66
  Physaster Pomel 1883 [incl. Inflataster Anthula 1899] WD66
    `--*P. inflatus (d’Orbigny 1854) [=Echinospatagus inflatus] WD66
  Turanglaster Solovyev & Melikov 1963 WD66
    `--*T. mazkii Solovyev & Melikov 1963 WD66

Aurelianaster Lambert & Thiéry 1925 [=Leiocorys Lambert in Vallette 1913, Vallettaster Lambert in Vallette 1924] WD66

Cardiaster granulosus (Goldfuss 1826) [=Spatangus granulosus; incl. S. cordiformis Woodward 1853, *Cardiaster cordiformis] WD66

Echinocorys Leske 1778 [=Echinocorytes Leske 1778, Galea Smith 1817 non Meuschen 1787, Spatagoides Bayle 1878; incl. Ananchites Lamarck 1801, Ananchytes Lamarck 1816, Corculum Goldfuss 1826, Galeola Quenstedt 1874, Oolaster Laube 1869] WD66

*Echinocorys scutatus Leske 1778 [=*Echinocorytes scutatus, *Galea scutata, *Spatagoides scutatus; incl. E. vulgaris Breynius 1732 (pre-Linnean)] WD66

Hemipneustes striatoradiatus (Leske 1778) [=Spatangus striatoradiatus; incl. S. radiatus Lamarck 1840, *Hemipneustes radiatus] WD66

Infulaster excentricus (Woodward 1833) [=Spatangus excentricus; incl. Cardiaster hagenowi d’Orbigny 1853, *Infulaster hagenowi] WD66

Metaporinus Agassiz 1844 [=Metaporhinus (l. c.); incl. Perioxus Pomel 1883, Thesaporhinus Ebray 1859 (n. n.), Thecaporinus (l. c.)] WD66

*Stenonaster tuberculata (Defrance 1816) [=Anachytes tuberculata, *Stenocorys tuberculata, *Stenonia tuberculata] WD66

*Type species of generic name indicated

REFERENCES

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

[F66] Fischer, A. G. 1966. Spatangoids. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt U. Echinodermata 3 vol. 2 pp. U543–U628. The Geological Society of America, Inc., and The University of Kansas Press.

[K79] Kauffman, E. G. 1979. Cretaceous. In: Robison, R. A., & C. Teichert (eds) Treatise on Invertebrate Paleontology pt A. Introduction. Fossilisation (Taphonomy), Biogeography and Biostratigraphy pp. A418–A487. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

[WD66] Wagner, C. D., & J. W. Durham. 1966. Holasteroids. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt U. Echinodermata 3 vol. 2 pp. U523–U543. The Geological Society of America, Inc., and The University of Kansas Press.

Dendrasteridae

Diagrammatic oral view of Scutellaster interlineatus, from Durham (1955).

Belongs within: Clypeasteroida.

The Dendrasteridae are a group of sand dollars known from the Pliocene to the present (Durham 1966b).

Characters (from Durham 1966b): Medium-sized to large; petals well developed; anterior petal more widely open than paired petals; interambulacrum 5 discontinuous; interambulacra nearly as wide as ambulacra at ambitus; four genital pores; food grooves bifurcating or trifurcating; periproct inframarginal to supramarginal.

<==Dendrasteridae D66b
    |--Dendraster Agassiz 1847 M89
    |    |--*D. excentricus (Eschscholtz 1831) M89 [=Scutella excentricus M89, Echinarachnius excentricus D66b]
    |    `--D. laevis D66a
    |--Merriamaster Lambert 1911 [incl. Orchophorus Lambert & Thiéry 1914, Twitchellia Lambert 1916] D66b
    |    `--*M. perrini (Weaver 1908) M89 [=Scutella perrini M89; incl. Orchophorus koehleri Lambert & Thiéry 1914 D66b]
    |--Scaphechinus Agassiz 1863 [=Chaetodiscus Lütken 1864] D66b
    |    `--*S. mirabilis Agassiz 1863 M89 [incl. Chaetodiscus scutella Lütken 1864 D66b]
    `--Scutellaster Cragin 1895 [=Anorthoscutum Lambert & Thiéry 1914, Calaster Kew 1920] D66b
         |--*S. interlineatus (Stimpson 1856) M89 (see below for synonymy)
         `--S. oregonensis M89
              |--S. o. oregonensis M89
              `--S. o. major M89

*Scutellaster interlineatus (Stimpson 1856) M89 [=Scutella interlineata M89, *Anorthoscutum interlineatum D66b, *Calaster interlineatus D66b; incl. Scutellaster cretaceus Cragin 1895 D66b]

*Type species of generic name indicated

REFERENCES

[D66a] Durham, J. W. 1966a. Echinoids—ecology and paleoecology. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt U. Echinodermata 3 vol. 1 pp. U257–U265. The Geological Society of America, Inc., and The University of Kansas Press.

[D66b] Durham, J. D. 1966b. Clypeasteroids. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt U. Echinodermata 3 vol. 2 pp. U450–U491. The Geological Society of America, Inc., and The University of Kansas Press.

[M89] Mooi, R. 1989. Living and fossil genera of the Clypeasteroida (Echinoidea: Echinodermata): an illustrated key and annotated checklist. Smithsonian Contributions to Zoology 488: 1–51.

Toxasteridae

Toxaster retusus, from the Natural History Museum.

Belongs within: Atelostomata.

The Toxasteridae are a group of heart urchins known from the Lower Cretaceous to the present, characterised by possession of a relatively primitive apical system, together with a typical lack of fascioles and/or primary spines. They are represented in the modern fauna by the bathyal Indonesian genus Isopatagus, which is the only living spatangoid urchin with circumoral tube feet possessing double pores (Fischer 1966).

Characters (from Fischer 1966): Apical system normally ethmophract (posterior genital plates mutually contiguous), gonopores 3 or 4; paired ambulacra petaloid, generally open, frontal ambulacrum petaloid or not; ambital and circumoral ambulacral pores double; fascioles usually absent; primary spines absent; plastron protamphisternous to mesamphisternous.

<==Toxasteridae [Toxasterina] F66
    |--Adytaster Lambert 1895 [=Adetaster Delage & Hérouard 1904] F66
    |    `--*A. lucentinus (Cotteau 1890) [=Cyclaster lucentinus] F66
    |--Aphelaster Lambert 1920 F66
    |    `--*A. integer Lambert 1920 F66
    |--Douvillaster Lambert 1917 [=Hypsaster Pomel 1883 non Pomel 1869] F66
    |    `--*D. vatonnei (Coquand 1878) [=Epiaster vatonnei, *Hypsaster vatonnei] F66
    |--Enallopneustes Pomel 1883 F66
    |    `--*E. jullieni (Peron & Gauthier 1881) [=Holaster jullieni] F66
    |--Isaster Desor 1858 F66
    |    `--*I. aquitanicus (Grateloup 1836) [=Spatangus aquitanicus] F66
    |--Isomicraster Lambert 1901 F66
    |    `--*I. stolleyi Lambert 1901 F66
    |--Isopatagus Mortensen 1948 F66
    |    `--*I. obovatus Mortensen 1948 F66
    |--Macraster Roemer 1888 F66
    |    `--*M. texanus Roemer 1888 F66
    |--Mokotibaster Lambert 1933 F66
    |    `--*M. hourequi Lambert 1933 F66
    |--Polydesmaster Lambert 1920 F66
    |    `--*P. fourtaui Lambert 1920 F66
    |--Palmeraster Sánchez Roig 1949 F66
    |    |--*P. palmeri Sánchez Roig 1949 F66
    |    `--P. herrerai F66
    |--Heteraster d’Orbigny 1853 (see below for synonymy) F66
    |    |--*H. oblongus (Brongniart 1821) [=Spatangus oblongus] F66
    |    |--H. grenovii (Forbes 1852) [=Hemipneustes grenovii, *Enallaster grenovii] F66
    |    `--H. trigonalis (Desor 1847) [=Micraster trigonalis, *Epiaster trigonalis] F66
    `--Toxaster Agassiz 1840 (see below for synonymy) F66
         |--*T. retusus (Lamarck 1816) [=Spatangus retusus] F66
         |--T. africanus F66
         |--‘Spatangus’ argilaceus Phillips 1829 [=*Hypsaster argilaceus] F66
         |--‘Echinospatagus’ breyniusi d’Orbigny 1859 [=*Miotoxaster breyniusi] F66
         |--‘Echinospatagus’ hofmanni M79
         |--T. lafittei F66
         `--‘*Pliotaxaster’ lyonsi Fourtau 1907 F66

Pre-Linnean: Echinospatagus cordiformis Breynius 1732 F66

Heteraster d’Orbigny 1853 [incl. Enallaster d’Orbigny 1853, Epiaster d’Orbigny 1854, Pseudoepiaster Seunes 1888, Taeniaster Lambert 1895 non Billings 1858] F66

Toxaster Agassiz 1840 [incl. Echinospatagus d’Orbigny 1853, Hypsaster Pomel 1883, Miotoxaster Pomel 1883, Pliotaxaster Fourtau 1907] F66

*Type species of generic name indicated

REFERENCES

[F66] Fischer, A. G. 1966. Spatangoids. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt U. Echinodermata 3 vol. 2 pp. U543–U628. The Geological Society of America, Inc., and The University of Kansas Press.

[M79] Müller, A. H. 1979. Fossilization (taphonomy). In: Robison, R. A., & C. Teichert (eds) Treatise on Invertebrate Paleontology pt A. Introduction. Fossilisation (Taphonomy), Biogeography and Biostratigraphy pp. A2–A78. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

Asterostomatidae

Platybrissus roemeri, copyright Koghisberg's Collections.

Belongs within: Atelostomata.

The Asterostomatidae is a possibly polyphyletic group of heart urchins characterised by the loss of petals and/or fascioles (Fischer 1966).

Characters (from Fischer 1966): Peristome labiate, phyllodes well developed, test generally fragile; radioles usually present; petals weakly developed or absent, fascioles of various types or absent; primary spines present or absent; apical system ethmolytic; plastron mesamphisternous, holamphisternous or ultramphisternous.

<==Asterostomatidae [Antillasterinae, Asterostomatina, Palaeopneustidae, Paleopneustidae] F66
    |--Asterostoma Agassiz 1847 F66
    |    `--*A. excentricum Agassiz 1847 F66
    |--Antillaster Lambert 1909 [=Pseudasterostoma Sánchez Roig 1952 non Duncan 1889] F66
    |    `--*A. cubensis (Cotteau 1871) [=Asterostoma cubensis, *Pseudasterostoma cubensis] F66
    |--Brissolampas Pomel 1883 F66
    |    `--*B. conicus (Dames 1877) [=Paleopneustes conicus] F66
    |--Brissomorpha Laube 1871 F66
    |    `--*B. fuchsi Laube 1871 F66
    |--Cleistechinus De Loriol 1882 F66
    |    `--*C. canaverii De Loriol 1882 F66
    |--Delopatagus Koehler 1907 F66
    |    `--*D. brucei Koehler 1907 F66
    |--Elipneustes Koehler 1914 [incl. Eurypneustes Koehler 1914 non Duncan & Sladen 1882] F66
    |    `--*E. denudatus (Koehler 1914) [=Eurypneustes denudatus] F66
    |--Genicopatagus Agassiz 1879 F66
    |    `--*G. affinis Agassiz 1879 F66
    |--Linopneustes Agassiz 1881 F66
    |    `--*L. murrayi (Agassiz 1873) [=Palaeopneustes murrayi] F66
    |--Megapetalus Clark 1929 F66
    |    `--*M. lovenioides Clark 1929 F66
    |--Moronaster Sánchez Roig 1952 F66
    |    `--*M. moronensis Sánchez Roig 1952 F66
    |--Palaeobrissus Agassiz 1883 [=Palaeobryssus Meissner 1903] F66
    |    `--*P. hilgardi Agassiz 1883 [=*Palaeobryssus hilgardi] F66
    |--Palaeotropus Lovén 1872 F66
    |    `--*P. josephinae Lovén 1872 F66
    |--Paleopneustes Agassiz 1873 [=Palaeopneustes Duncan 1889] F66
    |    `--*P. cristatus Agassiz 1873 [=*Palaeopneustes cristatus] F66
    |--Paleotrema Koehler 1914 [=Palaeotrema (l. c.)] F66
    |    `--*P. loveni (Agassiz 1881) [=Palaeotropus loveni] F66
    |--Peripatagus Koehler 1895 F66
    |    `--*P. cinctus Koehler 1895 F66
    |--Plesiozonus de Meijere 1902 F66
    |    `--*P. hirsutus de Meijere 1902 F66
    |--Prosostoma Pomel 1883 [=Pseudasterostoma Duncan 1889] F66
    |    `--*P. jimenoi (Cotteau 1870) [=Asterostoma jimenoi, *Pseudasterostoma jimenoi] F66
    |--Pycnolampas Agassiz & Clark 1907 F66
    |    `--*P. oviformis Agassiz & Clark 1907 F66
    |--Pygospatangus Cotteau 1888 F66
    |    `--*P. salvae Cotteau 1888 F66
    |--Argopatagus Agassiz 1879 [incl. Meijeria Döderlein 1906, Phrissocystis Agassiz 1898] F66
    |    |--*A. vitreus Agassiz 1879 F66
    |    |--‘*Phrissocystis’ aculeata Agassiz 1904 F66
    |    `--‘Phrissocystis’ humilis de Meijere 1902 [=*Meijeria humilis] F66
    |--Heterobrissus Manzoni & Mazzetti 1877 [incl. Archaeopneustes Gregory 1892] F66
    |    |--*H. montesi Manzoni & Mazzetti 1877 F66
    |    |--‘Palaeopneustes’ hystrix Agassiz 1880 [=*Archaeopneustes hystrix] F66
    |    `--H. niasicus F66
    `--Platybrissus Grube 1865 [=Platybryssus Meissner 1903] F66
         |--*P. (Platybrissus) roemeri Grube 1865 [=*Platybryssus roemeri] F66
         `--P. (Eurypatagus Mortensen 1948) F66
              `--P. (*E.) ovalis (Mortensen 1948) [=*Eurypatagus ovalis] F66

*Type species of generic name indicated

REFERENCES

[F66] Fischer, A. G. 1966. Spatangoids. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt U. Echinodermata 3 vol. 2 pp. U543–U628. The Geological Society of America, Inc., and The University of Kansas Press.

Hemiaster

Clockwise from lower left, (a) aboral, (b) oral, (c) lateral and (d) posterior views of Hemiaster bufo, from Fischer (1966).

Belongs within: Hemiasterina.

Hemiaster is a genus of broad and high heart urchins known from the Lower Cretaceous to the present (Fischer 1966).

See also: Hemiaster: an echinoid with heart.

Characters (from Fischer 1966): Test broad, high relative to length, abruptly truncated in rear, showing slight frontal sinus; apical system ethmophract, with four gonopores; frontal ambulacrum nonpetaloid or semipetaloid, with small round pores; paired ambulacra having relatively short petals, frontal pair longest.

<==Hemiaster Agassiz 1847 [incl. Leucaster Gauthier in Peron 1887, Peroniaster Gauthier 1887, Perionaster (l. c.)] F66
    |  i. s.: ‘*Peroniaster’ cotteaui Gauthier 1887 F66
    |         ‘*Leucaster’ remensis Gauthier 1887 F66
    |         H. texanus F66
    |--H. (Hemiaster) [incl. Integraster Lambert & Thiéry 1924 non Döderlein 1920] F66
    |    |--*H. (H.) bufo (Brongniart 1822) [=Spatangus bufo] F66
    |    `--H. (H.) ligeriensis d’Orbigny 1853 [=H. (*Integraster) ligeriensis] F66
    |--H. (Bolbaster Pomel 1869) F66
    |    `--H. (*B.) prunella (Lamarck 1816) [=Spatangus prunella] F66
    |--H. (Gregoryaster Lambert 1907) F66
    |    `--H. (*G.) coranguinum (Gregory 1892) [=Pericosmus coranguinum] F66
    |--H. (Holanthus Lambert & Thiéry 1924) F66
    |    `--H. (*H.) hickmanni Koehler 1914 F66
    |--H. (Leymeriaster Lambert & Thiéry 1924) F66
    |    `--H. (*L.) leymeriei Agassiz 1847 F66
    |--H. (Mecaster Pomel 1883) F66
    |    `--H. (*M.) fourneli Agassiz in Agassiz & Desor 1847 F66
    `--H. (Trachyaster Pomel 1869) F66
         `--H. (*T.) globosus (Pomel 1869) [=*Trachyaster globosus] F66

*Type species of generic name indicated

REFERENCES

[F66] Fischer, A. G. 1966. Spatangoids. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt U. Echinodermata 3 vol. 2 pp. U543–U628. The Geological Society of America, Inc., and The University of Kansas Press.

Hemiasterina

Apical view of Pericosmus latus, from the Natural History Museum.

Belongs within: Atelostomata.
Contains: Hemiaster, Schizaster.

The Hemiasterina are a lineage of heart urchins known from the Lower Cretaceous to the present, characterised by the presence of a peripetalous fasciole (Fischer 1966).

Characters (from Fischer 1966): Petaloid spatangoids having peripetalous fasciole, sometimes combined with lateroanal or marginal fasciole, but none with subanal fasciole. Apical system ethmophract to ethmolytic. Primary spines rarely developed.

<==Hemiasterina F66
    |--Aeropsidae F66
    |    |--Aeropsis Mortensen 1907 [=Aerope Jeffreys 1876 nec Leach 1813 nec Albers 1860] F66
    |    |    `--*A. rostrata (Wyville Thomson 1877) [=*Aerope rostrata] F66
    |    `--Aceste Wyville Thomson 1877 [=Acestina Lambert & Thiéry 1924] F66
    |         `--*A. bellidifera Wyville Thomson 1877 [=*Acestina bellidifera] F66
    |--Pericosmus Agassiz 1847 [incl. Megalaster Duncan 1877; Pericosmidae] F66
    |    |  i. s.: P. compressus (Duncan 1877) H86, F66 [=*Megalaster compressus F66; incl. Meoma crawfordi H86]
    |    |         P. melanostomus MD66
    |    |--*P. (Pericosmus) latus (Agassiz 1847) [=Micraster (*Pericosmus) latus] F66
    |    |--P. (Lambertona Sánchez Roig 1952) F66
    |    |    `--P. (*L.) lamberti (Sánchez Roig 1924) [=Victoriaster lamberti] F66
    |    `--P. (Victoriaster Lambert 1920) F66
    |         `--P. (*V.) gigas M’Coy 1882 F66
    |--Palaeostomatidae F66
    |    |--Homoeaster Pomel 1883 F66
    |    |    `--*H. tunetanus Pomel 1883 F66
    |    |--Lambertiaster Gauthier 1892 F66
    |    |    `--*L. douvillei Gauthier 1892 F66
    |    |--Leiostomaster Lambert 1920 F66
    |    |    `--*L. gentili Lambert 1920 F66
    |    |--Ornithaster Cotteau 1886 F66
    |    |    `--*O. evaristei Cotteau 1886 F66
    |    `--Palaeostoma Lovén in Agassiz 1872 (see below for synonymy) F66
    |         |--*P. mirabile (Gray 1851) [=*Leskia mirabilis] F66
    |         `--‘*Skouraster’ rochi Lambert 1937 F66
    |--Hemiasteridae F66
    |    |--Hemiaster F66
    |    |--Cheopsia Fourtau 1908 F66
    |    |    `--*C. mortenseni Fourtau 1908 F66
    |    |--Crucibrissus Lambert 1920 F66
    |    |    `--*C. integer (de Loriol 1891) [=Macropneustes integer] F66
    |    |--Distefanaster Checchia-Rispoli 1902 F66
    |    |    `--*D. garganicus Checchia-Rispoli 1902 F66
    |    |--Ditremaster Munier-Chalmas 1885 F66
    |    |    `--*D. nux (Desor 1853) [=Hemiaster nux] F66
    |    |--Hernandezaster Sánchez Roig 1949 F66
    |    |    `--*H. hernandezi Sánchez Roig 1949 F66
    |    |--Heterolampas Cotteau 1862 F66
    |    |    `--*H. maresi Cotteau 1862 F66
    |    |--Holcopneustes Cotteau 1889 F66
    |    |    `--*H. gourdoni (Cotteau 1887) [=Trachyaster gourdoni] F66
    |    |--Isopetalum Lambert 1911 [incl. Homoianthoides Lambert 1920] F66
    |    |    `--*I. pseudoverticale Oppenheim 1900 [=Linthia pseudoverticale] F66
    |    |--Opissaster Pomel 1883 F66
    |    |    `--*O. polygonalis Pomel 1883 F66
    |    |--Palhemiaster Lambert 1916 F66
    |    |    `--*P. peroni Lambert 1916 F66
    |    |--Sarsiaster Mortensen 1950 F66
    |    |    `--*S. griegii Mortensen 1950 F66
    |    |--Vomeraster Lambert 1920 F66
    |    |    `--*V. verrucosus (Coquand 1862) [=Hemiaster verrucosus] F66
    |    |--Washitaster Lambert 1927 F66
    |    |    `--*W. riovitsae (Adkins 1920) [=Hemiaster riovitsae] F66
    |    `--Hypsopatagus Pomel 1869 [=Hypsospatangus Cotteau 1890; incl. Trachypneustes Munier-Chalmas 1891] F66
    |         |--*H. (Hypsopatagus) meneghini (Desor 1858) [=Macropneustes meneghini, *Hypsospatangus meneghini] F66
    |         `--H. (Leiopneustes Cotteau 1885) [incl. Stenopatagus Lambert 1911] F66
    |              `--H. (*L.) antiquus (Agassiz 1847) [=Brissus antiquus] F66
    `--Schizasteridae [Schizasterinae] F66
         |--Schizaster F66
         |--Bathyspatus Pomel 1883 (n. d.) A66
         |--Amphipneustes Koehler 1900 [=Antipneustes Koehler 1926] F66
         |    `--*A. lorioli Koehler 1900 [=*Antipneustes lorioli] F66
         |--Hemifaorina Jeannet & Martin 1937 F66
         |    `--*H. tuber (Herklots 1854) [=Hemiaster tuber] F66
         |--Hemigymnia Arnaud 1898 F66
         |    `--*H. aturica Arnaud 1898 F66
         |--Parabrissus Bittner 1880 F66
         |    `--*P. pseudoprenaster Bittner 1880 F66
         |--Peribrissus Pomel 1869 F66
         |    `--*P. saheliensis Pomel 1869 F66
         |--Pseudobrissus Lambert 1905 F66
         |    `--*P. corsicus (Cotteau 1877) [=Brissus corsicus] F66
         |--Schizopneustes Thiéry 1907 [=Dipneustes Arnaud 1891 non Hoernes 1866] F66
         |    `--*S. aturicus (Arnaud in Cotteau 1891) [=*Dipneustes aturicus] F66
         |--Diploporaster Mortensen 1950 F66
         |    |--*D. barbatus Mortensen 1950 F66
         |    `--D. savignyi F66
         |--Faorina Gray 1851 [=Favorina (l. c.); incl. Atrapus Troschel 1851, Sinaechinus Hayasaka 1948] F66
         |    |--*F. chinensis Gray 1851 F66
         |    `--F. kawaguchii (Hayasaka 1948) [=*Sinaechinus kawaguchii] F66
         |--Periaster d’Orbigny 1853 F66
         |    |--*P. elatus (Desmoulins 1837) [=Spatangus elatus] F66
         |    `--P. obesus O75
         |--Tripylus Philippi 1845 [=Hamaxitus Troschel 1851; incl. Parapneustes Koehler 1912] F66
         |    |--*T. excavatus Philippi 1845 [=*Hamaxitus excavatus] F66
         |    `--‘*Parapneustes’ cordatus Koehler 1912 F66
         |--Proraster Lambert 1895 [incl. Sanfilippaster Checchia-Rispoli 1932] F66
         |    |--*P. atavus (Arnaud in Cotteau 1883) [=Schizaster atavus] F66
         |    |--P. geayi Cotteau 1908 [=*Sanfilippaster geayi] F66
         |    `--P. jukesii (Gray 1851) MG-H11
         |--Abatus Troschel 1851 [incl. Pseudabatus Koehler 1911, Parabatus (l. c.), Spatagodesma Agassiz 1898] F66
         |    |--*A. cavernosus (Philippi 1845) [=Spatangus (Tripylus) cavernosus] F66
         |    |--‘*Spatagodesma’ diomedae Agassiz 1898 F66
         |    `--‘*Pseudabatus’ nimrodi Koehler 1911 F66
         |--Brisaster Gray 1855 [incl. Indiaster Lambert 1920, Lymanaster Lambert 1920] F66
         |    |--*B. fragilis (Düben & Koren 1844) F66 [=Brissus fragilis F66, Tripylus fragilis BK77]
         |    |--B. indicus Koehler 1914 [=*Indiaster indicus] F66
         |    `--B. townsendi (Agassiz 1898) BK77, F66 [=Schizaster townsendi F66, *Lymanaster townsendi F66]
         |--Moira Agassiz 1872 [=Moera Michelin 1855 nec Leach 1814 nec Adams 1851] F66
         |    |--*M. (Moira) atropos (Lamarck 1816) (nom. cons.) [=Spatangus atropos, *Moera atropos] F66
         |    `--M. (Moiropsis Agassiz 1881) F66
         |         `--M. (*M.) claudicans (Agassiz 1879) [=Schizaster claudicans] F66
         |--Agassizia Agassiz & Desor 1847 [=Agassisia] F66
         |    |--*A. (Agassizia) scrobiculata Agassiz & Desor 1847 F66
         |    `--A. (Anisaster Pomel 1886) [incl. Eoagassizia Grant & Hertlein 1938] F66
         |         |--A. (*A.) gibberula Cotteau 1876 F66
         |         `--‘*Eoagassizia’ alta Grant & Hertlein 1938 F66
         |--Linthia Desor 1853 [incl. Escheria Desor 1853 non Heer 1847] F66
         |    |--L. (Linthia) F66
         |    |    |--*L. (L.) insignis Desor 1853 F66
         |    |    `--L. (L.) sudanensis F66
         |    `--L. (Lutetiaster Lambert 1920) F66
         |         `--L. (*L.) subglobosa (Lamarck 1816) [=Spatangus subglobosus] F66
         `--Prenaster Desor 1853 F66
              |--*P. (Prenaster) alpinus Desor 1853 F66
              |--P. (Protenaster Pomel 1883) [=Desoria Gray 1851 non Nicollet 1842] F66
              |    `--P. (*P.) australis (Gray 1851) [=*Desoria australis] F66
              `--P. (Salviniaster Lambert 1911) F66
                   `--P. (*S.) migueli (Lambert 1911) [=*Salviniaster migueli] F66

Palaeostoma Lovén in Agassiz 1872 [=Leskia Gray 1851 non Robineau Desvoidy 1830; incl. Skouraster Lambert 1937] F66

*Type species of generic name indicated

REFERENCES

[A66] Anon. 1966. Doubtful nominal genera of echinoids. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt U. Echinodermata 3 vol. 2 pp. U633. The Geological Society of America, Inc., and The University of Kansas Press.

[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.

[F66] Fischer, A. G. 1966. Spatangoids. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt U. Echinodermata 3 vol. 2 pp. U543–U628. The Geological Society of America, Inc., and The University of Kansas Press.

[H86] Hutton, F. W. 1886. Notes on some Australian Tertiary fossils. Proceedings of the Linnean Society of New South Wales, series 2, 1 (2): 481–482.

[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.

[MD66] Melville, R. V., & J. W. Durham. 1966. Echinoids—skeletal morphology. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt U. Echinodermata 3 vol. 1 pp. U220-U251. The Geological Society of America, Inc., and The University of Kansas Press.

[O75] Owen, R. 1875. On fossil evidences of a sirenian mammal (Eotherium aegyptiacum, Owen) from the nummulitic Eocene of the Mokattam cliffs, near Cairo. Quarterly Journal of the Geological Society of London 31: 100–105.

Micrasterina

Red heart urchin Meoma ventricosa, copyright John Miller.

Belongs within: Atelostomata.
Contains: Spatangidae, Loveniidae, Eupatagus.

The Micrasterina is a lineage of heart urchins known from the Cretaceous to the present (Fischer 1966). They first appear in the fossil record in the form of the family Micrasteridae, known from the Upper Cretaceous to the Eocene, which possessed subanal fascioles, a mesamphisternous plastron and ethmophract to transitional apical systems with three or four gonopores (Fischer 1966).

Characters (from Fischer 1966): Petaloid spatangoids with subanal fasciole (absent in some members of Loveniidae), which may be combined with peripetalous or internal fasciole. Primary spines developed except in Micrasteridae.

<==Micrasterina [Prymnodesmia] F66
    |--+--Spatangidae F66
    |  `--Loveniidae F66
    |--Micrasteridae [Micrasterinae] F66
    |    |--Brissopneustes Cotteau 1887 F66
    |    |    `--*B. villanovae Cotteau 1887 F66
    |    |--Isopneustes Pomel 1883 F66
    |    |    |--*I. bourgeoisi (Cotteau 1869) [=Cyclaster bourgeoisi] F66
    |    |    `--I. subquadratus F66
    |    |--Ovulaster Cotteau 1884 F66
    |    |    |--*O. gauthieri Cotteau 1884 F66
    |    |    `--O. auberti F66
    |    `--Micraster Agassiz 1836 [=Pycnaster Pomel 1883] F66
    |         |  i. s.: M. cortestudinarium K79
    |         |--*M. (Micraster) coranguinum (Leske 1778) [=Spatangus coranguinum, *Pycnaster coranguinum] F66
    |         `--M. (Gibbaster Gauthier 1887) F66
    |              `--M. (*G.) fastigatus Gauthier 1887 F66
    `--Brissidae [Brissina, Cyclasteridae, Unifasciidae] F66
         |--Eupatagus F66
         |--Aguayoaster Sánchez Roig 1952 F66
         |    `--*A. aguayoi Sánchez Roig 1952 F66
         |--Anabrissus Mortensen 1950 F66
         |    `--*A. damesi (Agassiz 1881) [=Brissus damesi] F66
         |--Arcaechinus Kier 1957 F66
         |    `--*A. auraduensis Kier 1957 F66
         |--Brissopatagus Cotteau 1863 [=Brissospatagus Agassiz 1874, Brissospatangus Cotteau 1866] F66
         |    `--*B. caumonti Cotteau 1863 [=*Brissospatangus caumonti] F66
         |--Cionobrissus Agassiz 1879 [=Cionobryssus Meissner 1903] F66
         |    `--*C. revinctus Agassiz 1879 [=*Cionobryssus revinctus] F66
         |--Fernandezaster Sánchez Roig 1952 F66
         |    `--*F. mortenseni Sánchez Roig 1952 F66
         |--Fourtaunia Lambert 1902 F66
         |    `--*F. santamariae (Gauthier in Fourtau 1900) [=Hypsospatangus santamariae] F66
         |--Gillechinus Fell 1964 F66
         |    `--*G. cudmorei Fell 1964 F66
         |--Herreraster Sánchez Roig 1951 F66
         |    `--*H. herrerae Sánchez Roig 1951 F66
         |--Hikelaster Lambert & Thiéry 1920 [=Troschelia Duncan & Sladen 1883] F66
         |    `--*H. tuberculatus (Duncan & Sladen 1883) [=*Troschelia tuberculata] F66
         |--Idiobryssus Clark 1939 F66
         |    `--*I. coelus Clark 1939 F66
         |--Lajanaster Sánchez Roig 1926 F66
         |    `--*L. jacksoni Sánchez Roig 1926 F66
         |--Lissospatangus Mortensen 1950 F66
         |    `--*L. hirsutus Mortensen 1950 F66
         |--Mariania Airaghi 1901 [=Airaghia Lambert 1910] F66
         |    `--*M. marmorae (Agassiz 1847) [=Macropneustes marmorae, *Airaghia marmorae] F66
         |--Mauritianaster Lambert 1920 F66
         |    `--*M. gentili Lambert 1920 F66
         |--Megapneustes Gauthier 1898 F66
         |    `--*M. grandis Gauthier 1898 F66
         |--Migliorinia Checchia-Rispoli 1942 F66
         |    `--*M. migiurtina Checchia-Rispoli 1942 F66
         |--Neopneustes Duncan 1889 F66
         |    `--*N. micrasteroides (Agassiz 1878) [=Rhynobrissus micrasteroides] F66
         |--Plesiaster Pomel 1883 F66
         |    `--*P. peini (Coquand 1862) [=Micraster peini] F66
         |--Plesiopatagus Pomel 1883 [=Plesiospatangus Cotteau 1886] F66
         |    `--*P. cotteaui (de Loriol 1880) [=Eupatagus cotteaui, *Plesiospatangus cotteaui] F66
         |--Radiobrissus Fourtau 1913 F66
         |    `--*R. gneffensis Fourtau 1913 F66
         |--Rhynobrissus Agassiz 1872 [=Rhinobrissus Quenstedt 1874, Rhinobryssus Meissner 1903] F66
         |    `--*R. pyramidalis Agassiz 1872 [=*Rhinobrissus pyramidalis, *Rhinobryssus pyramidalis] F66
         |--Rojasia Sánchez Roig 1951 F66
         |    `--*R. rojasi Sánchez Roig 1951 F66
         |--Spatagobrissus Clark 1923 F66
         |    `--*S. mirabilis Clark 1923 F66
         |--Spatangomorpha Boehm 1882 F66
         |    `--*S. eximia Boehm 1882 F66
         |--Stomaporus Cotteau 1888 F66
         |    `--*S. hispanicus Cotteau 1888 F66
         |--Trachypatagus Pomel 1869 [incl. Leiopatagus Pomel 1869 (n. n.), Liopatagus (l. c.)] F66
         |    `--*T. oranensis Pomel 1869 F66
         |--Unifascia Cooke 1959 F66
         |    `--*U. carolinensis (Clark 1915) [=Macropneustes carolinensis] F66
         |--Anametalia Mortensen 1950 F66
         |    |--*A. sternaloides (Bolau 1874) [=Brissus sternaloides] F66
         |    `--A. regularis (Clark 1925) MG-H11
         |--Cyclaster Cotteau 1856 F66
         |    |--*C. declivus Cotteau 1856 F66
         |    `--C. recens F66
         |--Diplodetus Schlüter 1900 F66
         |    |--*D. (Diplodetus) brevistella Schlüter 1900 F66
         |    `--D. (Protobrissus Lambert 1907) F66
         |         `--D. (*P.) mortenseni (Lambert 1907) [=*Protobrissus mortenseni] F66
         |--Pharaonaster Lambert 1920 F66
         |    |--*P. (P.) ammon (Agassiz 1847) [=Macropneustes ammon] F66
         |    `--P. (Thebaster Checchia-Rispoli 1941) F66
         |         `--P. (*T.) fischeri (de Loriol 1881) [=Macropneustes fischeri] F66
         |--Brissus Gray 1825 (nom. cons.) (see below for synonymy) F66
         |    |--*B. unicolor (Leske 1778) [=Spatangus brissus unicolor, *Bryssus unicolor] F66
         |    |--B. agassizii Döderlein 1885 MG-H11 [=B. (*Allobrissus) agassizii F66]
         |    |--‘*Sandiegoaster’ durhami Sánchez Roig 1952 F66
         |    `--B. latecarinatus (Leske 1778) BM09
         |--Macropneustes Agassiz 1847 [incl. Plagiopneustes Fourtau 1905] F66
         |    |  i. s.: M. crassus [=*Plagiopneustes crassus] F66
         |    |--*M. (Macropneustes) deshayesi Agassiz 1847 F66
         |    `--M. (Deakia Pavay 1875) F66
         |         `--M. (*D.) rotundata (Pavay 1875) [=*Deakia rotundata] F66
         |--Plagiobrissus Pomel 1883 (see below for synonymy) F66
         |    |--*P. (Plagiobrissus) grandis (Gmelin 1788) (see below for synonymy) F66
         |    `--P. (Rhabdobrissus Cotteau 1889) [=Rhabdobryssus Meissner 1903; incl. Mortensaster Lambert 1922] F66
         |         |--P. (*R.) jullieni (Cotteau 1889) [=*Rhabdobrissus jullieni, *Rhabdobryssus jullieni] F66
         |         `--‘Metalia’ costae Gasco 1876 [=*Mortensaster costae] F66
         |--Gualtieria Agassiz 1847 [=Gaultieria (l. c.), Gualteria (l. c.), Gualtiera (l. c.)] F66
         |    |--*G. (Gualtieria) orbignyana Agassiz 1847 F66
         |    |--G. (Blaviaster Lambert 1920) [=Temnaster Lambert 1912 non Verrill 1894] F66
         |    |    `--G. (*B.) grossouvrei (Lambert 1912) [=Temnaster grossouvrei] F66
         |    `---G. (Granobrissoides Lambert 1920) F66
         |          `--G. (*G.) australiae Cotteau 1889 F66
         |--Brissopsis Agassiz in Agassiz & Desor 1847 (see below for synonymy) F66
         |    |--*B. lyrifera (Forbes 1841) [=Brissus lyrifer, *Bryssopsis lyrifera] F66
         |    |--B. cluciei Wright 1855 [=*Brissoma cluciei] F66
         |    |--B. elegans Agassiz 1847 [=*Toxobrissus elegans, *Toxobryssus elegans] F66
         |    |--B. elongata MD66
         |    |--B. lamberti Gauthier 1900 [=*Zeugaster lamberti] F66
         |    |--B. luzonicus (Gray 1851) MG-H11 [=*Kleinia luzonica F66]
         |    `--B. pacifica F66
         |--Metalia Gray 1855 (see below for synonymy) F66
         |    |--*M. sternalis (Lamarck 1816) [=Spatangus sternalis] F66
         |    |--M. dicrana Clark 1917 BM09
         |    |--‘*Xanthobrissus’ garreti Agassiz 1863 F66
         |    |--‘Echinus’ maculosus Gmelin 1788 [=*Metalopsis maculosus] F66
         |    |--‘Brissus’ robillardi de Loriol 1876 [=*Prometalia robillardi] F66
         |    |--M. spatagus (Linnaeus 1758) BM09
         |    `--‘*Eobrissus’ townsendi Bell 1904 F66
         `--Meoma Gray 1851 F66
              |--M. (Meoma) [incl. Hemibrissus Pomel 1869, Rhyssobrissus Agassiz 1863] F66
              |    |--*M. (M.) grandis Gray 1851 F66
              |    `--M. (M.) ventricosa (Lamarck 1816) [=Spatangus ventricosus, *Hemibrissus ventricosus] F66
              |--M. (Plethotaenia Clark 1917) F66
              |    `--M. (*P.) spatangoides (Agassiz 1883) [=Macropneustes spatangoides] F66
              `--M. (Schizobrissus Pomel 1869) [incl. Peripneustes Cotteau 1875] F66
                   |--M. (*S.) cruciata (Agassiz 1847) [=Brissus cruciatus] F66
                   `--M. (S.) antillarum (Cotteau 1875) [=*Peripneustes antillarum] F66

Brissopsis Agassiz in Agassiz & Desor 1847 [=Bryssopsis Meissner 1903; incl. Brissoma Pomel 1888, Kleinia Gray 1851, Toxobrissus Desor 1858, Toxobryssus Meissner 1903, Zeugaster Lambert 1907] F66

Brissus Gray 1825 (nom. cons.) [=Bryssus Martens 1869; incl. Allobrissus Mortensen 1950, Sandiegoaster Sánchez Roig 1952] F66

Metalia Gray 1855 [incl. Eobrissus Bell 1904, Metaliopsis Fourtau 1913, Prometalia Pomel 1883, Xanthobrissus Agassiz 1863] F66

Plagiobrissus Pomel 1883 [=Plagionotus Agassiz & Desor 1847 non Mulsant 1842, Plagiostomus d’Orbigny 1854 non Hermansen 1847] F66

*Plagiobrissus (Plagiobrissus) grandis (Gmelin 1788) [=Echinus grandis, *Plagionotus grandis, *Plagiostomus grandis] F66

*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.

[F66] Fischer, A. G. 1966. Spatangoids. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt U. Echinodermata 3 vol. 2 pp. U543–U628. The Geological Society of America, Inc., and The University of Kansas Press.

[K79] Kauffman, E. G. 1979. Cretaceous. In: Robison, R. A., & C. Teichert (eds) Treatise on Invertebrate Paleontology pt A. Introduction. Fossilisation (Taphonomy), Biogeography and Biostratigraphy pp. A418–A487. The Geological Society of America, Inc.: Boulder (Colorado), 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.

[MD66] Melville, R. V., & J. W. Durham. 1966. Echinoids—skeletal morphology. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt U. Echinodermata 3 vol. 1 pp. U220–U251. The Geological Society of America, Inc., and The University of Kansas Press.