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

Belongs within: Irregularia.

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 1966b): 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.

    |--Cardabia bullarensis Foster & Philip 1978SG93
    |--Calymne Thomson 1877 [Calymnidae]WD66b
    |    `--*C. relicta Thomson 1877WD66b
    |--Stenonaster Lambert 1922 [=Stenocorys Lambert & Thiéry 1917, Stenonia Desor 1858 non Gray 1853; Stenonasteridae]WD66b
    |    |--*S. tuberculata (Defrance 1816) [=Anachytes tuberculata, *Stenocorys tuberculata, *Stenonia tuberculata]WD66b
    |    |--S. douvillei Lambert 1928SG93
    |    `--S. morgani Gauthier 1902SG93
    |    |--Chelonechinus Bather 1934WD66b
    |    |    `--*C. suvae Bather 1934WD66b
    |    |--Pilematechinus Agassiz 1904WD66b
    |    |    `--*P. rathbuni (Agassiz 1898) [=Cystechinus rathbuni]WD66b
    |    |--Plexechinus Agassiz 1896WD66b
    |    |    |--*P. cinctus Agassiz 1896WD66b
    |    |    `--P. planus Mironov 1978SS05
    |    |--Sanchezaster Lambert in Sánchez Roig 1924WD66b
    |    |    `--*S. habanensis Lambert in Sánchez Roig 1924WD66b
    |    |--Sternopatagus de Maijere 1902WD66b
    |    |    `--*S. sibogae de Meijere 1902WD66b
    |    `--Urechinus Agassiz 1879 [incl. Cystechinus Agassiz 1879]WD66b
    |         |--*U. naresianus Agassiz 1879WD66b
    |         `--*Cystechinus’ wyvillei Agassiz 1879WD66b
    |    |--Ceratophysa Pomel 1883 [=Rodocystis Lambert & Thiéry 1924]WD66b
    |    |    `--*C. ceratopyga (Agassiz 1879) [=*Rodocystis ceratopyga]WD66b
    |    |--Cystocrepis Mortensen 1907WD66b
    |    |    `--*C. setigera (Agassiz 1898) [=Echinocrepis setigera]WD66b
    |    |--Echinocrepis Agassiz 1879WD66b
    |    |    `--*E. cuneata Agassiz 1879WD66b
    |    |--Echinosigra Mortensen 1907WD66b
    |    |    `--*E. phiale (Thomson 1873) [=Pourtalesia phiale]WD66b
    |    |--Helgocystis Mortensen 1907WD66b
    |    |    `--*H. carinata (Agassiz 1879) [=Pourtalesia carinata]WD66b
    |    |--Spatagocystis Agassiz 1879WD66b
    |    |    `--*S. challengeri Agassiz 1879WD66b
    |    `--Pourtalesia Agassiz 1869 [incl. Phyale Pomel 1883 non Koch 1847, Phyalopsis Pomel 1883]WD66b
    |         |--*P. miranda Agassiz 1869WD66b
    |         |--P. jeffreysi Thomson 1873 [=*Phyale jeffreysi]WD66b
    |         `--P. laguncula Agassiz 1879 [=*Phyalopsis laguncula]WD66b
         |--Aurelianaster Lambert & Thiéry 1925 (see below for synonymy)WD66b
         |    `--*A. valettei (Lambert in Vallette 1913) [=*Leiocorys valettei, *Vallettaster valettei]WD66b
         |--Basseaster Lambert 1936WD66b
         |    `--*B. rostratus Lambert 1936WD66b
         |--Cardiaster Forbes 1850WD66b
         |    `--C. granulosus (Goldfuss 1826) (see below for synonymy)WD66b
         |--Cardiotaxis Lambert 1917WD66b
         |    `--*C. peroni (Lambert 1887) [=Cardiaster peroni]WD66b
         |--Cibaster Pomel 1883WD66b
         |    `--*C. bourgeoisanus d’Orbigny 1853) [=Cardiaster bourgeoisanus]WD66b
         |--Duncaniaster Lambert 1896WD66b
         |    `--*D. australiae (Duncan 1887) [=Holaster australiae]WD66b
         |--Entomaster Gauthier 1888WD66b
         |    `--*E. rousseli Gautheir 1888WD66b
         |--Galeaster Seunes 1889WD66b
         |    `--*G. bertrandi Seunes 1889WD66b
         |--Ganbirretia Gauthier 1903WD66b
         |    `--*G. douvillei Gauthier 1903WD66b
         |--Garumnaster Lambert 1907WD66b
         |    `--*G. michaleti Lambert 1907WD66b
         |--Guettaria Gauthier 1888WD66b
         |    `--*G. angladei Gauthier 1888WD66b
         |--Hagenowia Duncan 1889 [=Hagenovia (l. c.), Martinosigra Nielson 1942]WD66b
         |    `--*H. rostrata (Forbes 1889) [=Cardiaster rostratus, *Martinosigra rostrata]WD66b
         |--Hemipneustes Agassiz 1836WD66b
         |    `--H. striatoradiatus (Leske 1778) (see below for synonymy)WD66b
         |--Infulaster Desor 1858WD66b
         |    `--I. excentricus (Woodward 1833) (see below for synonymy)WD66b
         |--Ismidaster Boehm 1927WD66b
         |    `--*I. toulai Boehm 1927WD66b
         |--Jeronia Seunes 1888WD66b
         |    `--*J. pyrenaica Seunes 1888WD66b
         |--Labrotaxis Casey 1960WD66b
         |    `--*L. cantianus (Casey 1960) [=Holaster (*Labrotaxis) cantianus]WD66b
         |--Lampadaster Cotteau 1889WD66b
         |    `--*L. gradidieri Cotteau 1889WD66b
         |--Lampadocorys Pomel 1883WD66b
         |    `--*L. sulcatus (Cotteau 1873) [=Holaster sulcatus]WD66b
         |--Messaoudia Lambert 1917WD66b
         |    `--*M. pyriformis (Peron & Gauthier 1878) [=Holaster pyriformis]WD66b
         |--Offaster Desor 1858WD66b
         |    `--*O. pilula (Lamarck 1816) [=Ananchytes pilula]WD66b
         |--Opisopneustes Gauthier 1889WD66b
         |    `--*O. cossoni Gauthier 1889WD66b
         |--Paronaster Airaghi 1906WD66b
         |    `--*P. cupuliformis Airaghi 1906WD66b
         |--Pseudholaster Pomel 1883WD66b
         |    `--*P. bicarinatus (Agassiz 1847) [=Holaster bicarinatus]WD66b
         |--Pseudoffaster Lambert 1924WD66b
         |    `--*P. caucasicus (Dru 1884) [=Holaster caucasicus]WD66b
         |--Rispolia Lambert 1917WD66b
         |    `--*R. subtrigonata (Catullo 1827) [=Nucleolites subtrigonatus]WD66b
         |--Scagliaster Munier-Chalmas 1891WD66b
         |    `--*S. concavus (Catullo 1827) [=Ananchytes concavus]WD66b
         |--Stereopneustes de Meijere 1902WD66b
         |    `--*S. relictus de Meijere 1902WD66b
         |--Taphraster Pomel 1883WD66b
         |    `--*T. campicheanus [=Holaster campicheanus]WD66b
         |--Tholaster Seunes 1890 [=Gibbaster Seunes 1889 non Gauthier 1887]WD66b
         |    `--*T. munieri (Seunes 1889) [=*Gibbaster munieri]WD66b
         |--Titanaster Szörényi 1929WD66b
         |    `--*T. labiostoma Szörényi 1929WD66b
         |--Zumoffenia Fourtau 1912WD66b
         |    `--*Z. ludovici Fourtau 1912WD66b
         |--Pseudananchys Pomel 1883 [incl. Craginaster Lambert 1903]WD66b
         |    |--*P. algiris (Coquand 1862) [=Ananchytes algiris]WD66b
         |    `--‘Holaster’ completa Cragin 1893 [=*Craginaster completa]WD66b
         |--Sternotaxis Lambert 1893 [incl. Plesiocorys Pomel 1883]WD66b
         |    |--*S. planus (Mantell 1822) [=Spatangus planus]WD66b
         |    `--‘Holaster’ placenta Agassiz in Agassiz & Desor 1847 [=*Plesiocorys placenta]WD66b
         |--Toxopatagus Pomel 1883 [incl. Heteropneustes Pomel 1883]WD66b
         |    |--*T. italicus (Manzoni 1878) [=Hemipneustes italicus]WD66b
         |    `--*Heteropneustes’ deletrei Pomel 1883WD66b
         |--Stegaster Pomel 1883 [incl. Seunaster Lambert in Blayac 1912, Synochitis Lambert 1917]WD66b
         |    |--*S. gillieroni (de Loriol 1873) [=Cardiaster gillieroni]WD66b
         |    |--‘Holaster’ bouillei Cotteau in de Bouille 1873 [=*Seunaster bouillei]WD66b
         |    `--S. cotteauiWD66b
         |--Echinocorys Leske 1778 (see below for synonymy)WD66b
         |    |--*E. scutatus Leske 1778 (see below for synonymy)WD66b
         |    |--‘Ananchites’ carinatus Risso 1826R26
         |    |--‘Anachytes’ corculum Goldfuss 1826 [=*Corculum corculum]WD66b
         |    |--*Oolaster’ mattseensis Laube 1869WD66b
         |    |--*Ananchytes’ ovatus Lamarck 1816WD66b
         |    |--*Galeola’ papillosa Quenstedt 1874WD66b
         |    |--‘Ananchites’ rotundatus Risso 1826R26
         |    `--‘Ananchites’ stella Risso 1826R26
         `--Holaster Agassiz 1836 [incl. Ananchothuria Fossa-Mancini 1919, Holasteropsis Elbert 1902]WD66b
              |--*H. nodulosus (Goldfuss 1829) [=Spatangus nodulosus]WD66b
              |--H. cordatus Dubois 1836SG93
              |--*Holasteropsis’ credneriana Elbert 1902WD66b
              |--H. grasanus d’Orbigny 1853SG93
              |--H. planusK79
              |--H. subglobosusK79
              |--*Ananchothuria’ tesselata Fossa-Mancini 1919WD66b
              `--H. valanginensis Lambert 1917SG93
Holasteroida incertae sedis:
  Corechinus Kongiel 1936WD66b
    `--*C. pulaviensis Kongiel 1936WD66b
  Habanaster Lambert in Sánchez Roig 1924WD66b
    `--*H. sanchezi Lambert in Sánchez Roig 1924WD66b
  Menuthiaster Lambert 1896WD66b
    `--*M. cotteaui Lambert 1896WD66b
  Nordenskjoeldaster Lambert 1910WD66b
    `--*N. antarcticus Lambert 1910WD66b
  Physaster Pomel 1883 [incl. Inflataster Anthula 1899]WD66b
    `--*P. inflatus (d’Orbigny 1854) [=Echinospatagus inflatus]WD66b
  Turanglaster Solovyev & Melikov 1963WD66b
    `--*T. mazkii Solovyev & Melikov 1963WD66b
  Galeraster Cotteau 1890SG93, WD66a
    `--*G. australiae Cotteau 1890WD66a
    |--G. bellissae Foster & Philip 1978MM22
    |--G. sulcatusMM22
    `--G. tertiarusMM22

Aurelianaster Lambert & Thiéry 1925 [=Leiocorys Lambert in Vallette 1913, Vallettaster Lambert in Vallette 1924]WD66b

Cardiaster granulosus (Goldfuss 1826) [=Spatangus granulosus; incl. S. cordiformis Woodward 1853, *Cardiaster cordiformis]WD66b

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]WD66b

*Echinocorys scutatus Leske 1778 [=*Echinocorytes scutatus, *Galea scutata, *Spatagoides scutatus; incl. E. vulgaris Breynius 1732 (pre-Linnean)]WD66b

Hemipneustes striatoradiatus (Leske 1778) [=Spatangus striatoradiatus; incl. S. radiatus Lamarck 1840, *Hemipneustes radiatus]WD66b

Infulaster excentricus (Woodward 1833) [=Spatangus excentricus; incl. Cardiaster hagenowi d’Orbigny 1853, *Infulaster hagenowi]WD66b

*Type species of generic name indicated


[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).

[MM22] McNamara, K. J., & S. Martin. 2022. Middle Eocene echinoids from the western Eucla Basin, Western Australia. Records of the Western Australian Museum 37: 31–56.

[R26] Risso, A. 1826. Histoire naturelle des principales productions de l’Europe méridionale et particulièrement de celles des environs de Nice et des Alpes maritimes vol. 5. F.-G. Levrault: Paris.

[SG93] Simms, M. J., A. S. Gale, P. Gilliland, E. P. F. Rose & G. D. Sevastopulo. 1993. Echinodermata. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 491–528. Chapman & Hall: London.

[SS05] Stockley, B., A. B. Smith, T. Littlewood, H. A. Lessios & J. A. Mackenzie-Dodds. 2005. Phylogenetic relationships of spatangoid sea urchins (Echinoidea): taxon sampling density and congruence between morphological and molecular estimates. Zoologica Scripta 34: 447–468.

[WD66a] Wagner, C. D., & J. W. Durham. 1966a. Holectypoids. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt U. Echinodermata 3 vol. 2 pp. U440–U450. The Geological Society of America, Inc., and The University of Kansas Press.

[WD66b] Wagner, C. D., & J. W. Durham. 1966b. 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.

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