Red heart urchin Meoma ventricosa, copyright John Miller.

Belongs within: Irregularia.
Contains: Brissopsis, LoveniidaeSpatangus, Maretiidae.

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).

The Spatangidae are a family of heart urchins with petaloid ambulacra known from the Eocene to the present (Fischer 1966).

Eupatagus is a cosmopolitan genus of ovoid urchins known from the Eocene to the present (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
    |--Micrasteridae [Cyclasteridae, Micrasterinae]SS05
    |    |  i. s.: Brissopneustes Cotteau 1887F66
    |    |           `--*B. villanovae Cotteau 1887F66
    |    |         Isopneustes Pomel 1883F66
    |    |           |--*I. bourgeoisi (Cotteau 1869) [=Cyclaster bourgeoisi]F66
    |    |           `--I. subquadratus (Desor 1857)SG93
    |    |         Ovulaster Cotteau 1884F66
    |    |           |--*O. gauthieri Cotteau 1884F66
    |    |           `--O. aubertiF66
    |    |--Micraster Agassiz 1836SS05, F66 [=Plesiaster Pomel 1883SG93, F66, Pycnaster Pomel 1883F66]
    |    |    |  i. s.: M. antiquus Cotteau 1887SG93
    |    |    |         M. borchardi Hagenow 1853SG93
    |    |    |         M. corbovis Forbes 1850SG93
    |    |    |         M. cortestudinariumK79
    |    |    |         M. peini Coquand 1862 [=*Plesiaster peini]F66
    |    |    |--*M. (Micraster) coranguinum (Leske 1778) [=Spatangus coranguinum, *Pycnaster coranguinum]F66
    |    |    `--M. (Gibbaster Gauthier 1887)F66
    |    |         `--M. (*G.) fastigatus Gauthier 1887F66
    |    `--+--Cyclaster Cotteau in Leymerie & Cotteau 1856SS05, MM22
    |       |    |--*C. declivus Cotteau in Leymerie & Cotteau 1856MM22
    |       |    |--C. archeri (Tenison Woods 1867)MM22
    |       |    |--C. aturicus (Seunes 1888)MM22
    |       |    |--C. jamiei McNamara & Martin 2022MM22
    |       |    |--C. recensF66
    |       |    |--C. regalisSS05
    |       |    |--C. ruegensis (Kutscher 1978)MM22
    |       |    `--C. suecicus (Schlüter 1897)MM22
    |       `--+--Isaster Desor 1858SS05, MM22
    |          |    `--*I. aquitanicus (Grateloup 1836) [=Spatangus aquitanicus]F66
    |          `--Isopatagus Mortensen 1948SS05, F66
    |               `--*I. obovatus Mortensen 1948F66
    `--Brissidea [Brissidae, Brissina, Prospatangidae, Spatangidae, Unifasciidae]SS05
         |  i. s.: Aguayoaster Sánchez Roig 1952F66
         |           `--*A. aguayoi Sánchez Roig 1952F66
         |         Brissopatagus Cotteau 1863 [=Brissospatagus Agassiz 1874, Brissospatangus Cotteau 1866]F66
         |           `--*B. caumonti Cotteau 1863 [=*Brissospatangus caumonti]F66
         |         Fernandezaster Sánchez Roig 1952F66
         |           `--*F. mortenseni Sánchez Roig 1952F66
         |         Fourtaunia Lambert 1902F66
         |           `--*F. santamariae (Gauthier in Fourtau 1900) [=Hypsospatangus santamariae]F66
         |         Herreraster Sánchez Roig 1951F66
         |           `--*H. herrerae Sánchez Roig 1951F66
         |         Hikelaster Lambert & Thiéry 1920 [=Troschelia Duncan & Sladen 1883 non Mörch 1876]F66
         |           `--*H. tuberculatus (Duncan & Sladen 1883) [=*Troschelia tuberculata]F66
         |         Idiobryssus Clark 1939F66
         |           `--*I. coelus Clark 1939F66
         |         Lajanaster Sánchez Roig 1926F66
         |           `--*L. jacksoni Sánchez Roig 1926F66
         |         Mariania Airaghi 1901 [=Airaghia Lambert 1910]F66
         |           `--*M. marmorae (Agassiz 1847) [=Macropneustes marmorae, *Airaghia marmorae]F66
         |         Mauritianaster Lambert 1920F66
         |           `--*M. gentili Lambert 1920F66
         |         Megapneustes Gauthier 1898F66
         |           `--*M. grandis Gauthier 1898F66
         |         Neopneustes Duncan 1889F66
         |           `--*N. micrasteroides (Agassiz 1878) [=Rhynobrissus micrasteroides]F66
         |         Radiobrissus Fourtau 1913F66
         |           `--*R. gneffensis Fourtau 1913F66
         |         Rojasia Sánchez Roig 1951F66
         |           `--*R. rojasi Sánchez Roig 1951F66
         |         Spatangomorpha Boehm 1882F66
         |           `--*S. eximia Boehm 1882F66
         |         Stomaporus Cotteau 1888F66
         |           `--*S. hispanicus Cotteau 1888F66
         |         Trachypatagus Pomel 1869 [incl. Leiopatagus Pomel 1869 (n. n.), Liopatagus (l. c.)]F66
         |           `--*T. oranensis Pomel 1869F66
         |         Unifascia Cooke 1959F66
         |           `--*U. carolinensis (Clark 1915) [=Macropneustes carolinensis]F66
         |         Pharaonaster Lambert 1920F66
         |           |--*P. (P.) ammon (Agassiz 1847) [=Macropneustes ammon]F66
         |           `--P. (Thebaster Checchia-Rispoli 1941)F66
         |                `--P. (*T.) fischeri (de Loriol 1881) [=Macropneustes fischeri]F66
         |         Diplodetus Schlüter 1900F66
         |           |  i. s.: D. recklinghausenensis Schlüter 1900SG93
         |           |--*D. (Diplodetus) brevistella Schlüter 1900F66
         |           `--D. (Protobrissus Lambert 1907)F66
         |                `--D. (*P.) mortenseni (Lambert 1907) [=*Protobrissus mortenseni]F66
         |         Laevipatagus Noetling 1885 [incl. Leiospatangus Mayer 1861 (n. n.)]F66
         |           `--*L. bigibbus (von Beyrich 1848) [=Spatangus (Micraster) bigibbus]F66
         |         Semipetalion Szörényi 1963F66
         |           `--*S. anomon (Szörényi 1963) [=Atelospatangus (*Semipetalion) anomon]F66
         |         Nacospatangus Agassiz 1873 [=Nacopatagus Agassiz 1881]F66
         |           |--*N. (Nacospatangus) gracilis Agassiz 1873 [=*Nacopatagus gracilis]F66
         |           `--N. (Pseudomaretia Koehler 1914) (see below for synonymy)F66
         |                |--N. (*P.) alta (Agassiz 1863) [=Maretia alta]F66
         |                |--*Lonchophorus’ interruptus Studer 1880F66
         |                `--*Gonimaretia’ tylota Clark 1917F66
         |--+--+--Plesiopatagus Pomel 1883SS05, F66 [=Plesiospatangus Cotteau 1886F66]
         |  |  |    `--*P. cotteaui (de Loriol 1880) [=Eupatagus cotteaui, *Plesiospatangus cotteaui]F66
         |  |  `--+--Allobrissus Mortensen 1950SS05, F66
         |  |     |    `--*A. agassizii Döderlein 1885F66, SS05 [=Brissus (*Allobrissus) agassiziiF66]
         |  |     |--Anabrissus Mortensen 1950SS05, F66
         |  |     |    `--*A. damesi (Agassiz 1881) [=Brissus damesi]F66
         |  |     `--Brissus Gray 1825 (nom. cons.)SS05, F66 [=Bryssus Martens 1869F66; incl. Sandiegoaster Sánchez Roig 1952F66]
         |  |          |--*B. unicolor (Leske 1778) [=Spatangus brissus unicolor, *Bryssus unicolor]F66
         |  |          |--*Sandiegoaster’ durhami Sánchez Roig 1952F66
         |  |          |--B. latecarinatus (Leske 1778)BM09
         |  |          `--B. obesus Verrill 1867SS05
         |  `--+--BrissopsisSS05
         |     `--+--Archaeopneustes Gregory 1892SS05, F66
         |        |    `--*Archaeopneustes hystrix Agassiz 1880F66, SS05 [=Palaeopneustes hystrixF66]
         |        `--Meoma Gray 1851SS05, F66
         |             |  i. s.: M. brevipetalataH79
         |             |         M. tuberculataH79
         |             |--M. (Meoma) [incl. Hemibrissus Pomel 1869, Rhyssobrissus Agassiz 1863]F66
         |             |    |--*M. (M.) grandis Gray 1851F66
         |             |    `--M. (M.) ventricosa (Lamarck 1816) [=Spatangus ventricosus, *Hemibrissus ventricosus]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
         `--+--Macropneustes Agassiz 1847SS05, F66 [incl. Plagiopneustes Fourtau 1905F66]
            |    |  i. s.: M. australisH79
            |    |         M. cordatusH79
            |    |         M. crassus [=*Plagiopneustes crassus]F66
            |    |         M. spatangiformisH79
            |    |--*M. (Macropneustes) deshayesi Agassiz 1847F66
            |    `--M. (Deakia Pavay 1875)F66
            |         `--M. (*D.) rotundata (Pavay 1875) [=*Deakia rotundata]F66
            `--+--+--Anametalia Mortensen 1950SS05, F66
               |  |    |--*A. sternaloides (Bolau 1874) [=Brissus sternaloides]F66
               |  |    `--A. regularis (Clark 1925)MG-H11
               |  `--+--Cionobrissus Agassiz 1879SS05, F66 [=Cionobryssus Meissner 1903F66]
               |     |    `--*C. revinctus Agassiz 1879 [=*Cionobryssus revinctus]F66
               |     `--+--+--Granobrissoides Lambert 1920SS05, F66
               |        |  |    `--*G. australiae (Cotteau 1889) [=Gualtieria (*Granobrissoides) australiae]F66
               |        |  `--Rhabdobrissus Cotteau 1889SS05, F66 (see below for synonymy)
               |        |       |--*R. jullieni Cotteau 1889 (see below for synonymy)F66
               |        |       `--‘Metalia’ costae Gasco 1876 [=*Mortensaster costae]F66
               |        `--+--Plagiobrissus Pomel 1883SS05, F66 (see below for synonymy)
               |           |    `--*P. grandis (Gmelin 1788) (see below for synonymy)F66
               |           `--+--Eobrissus Bell 1904SS05, F66
               |              |    `--*E. townsendi Bell 1904F66
               |              `--Metalia Gray 1855SS05, F66 (see below for synonymy)
               |                   |--*M. sternalis (Lamarck 1816) [=Spatangus sternalis]F66
               |                   |--M. dicrana Clark 1917BM09
               |                   |--*Xanthobrissus’ garreti Agassiz 1863F66
               |                   |--‘Echinus’ maculosus Gmelin 1788 [=*Metalopsis maculosus]F66
               |                   |--M. nobilis Verrill 1867–1871SS05
               |                   |--‘Brissus’ robillardi de Loriol 1876 [=*Prometalia robillardi]F66
               |                   `--M. spatagus (Linnaeus 1758)BM09
               `--+--Rhynobrissus Agassiz 1872SS05, F66 [=Rhinobrissus Quenstedt 1874F66, Rhinobryssus Meissner 1903F66]
                  |    |--*R. pyramidalis Agassiz 1872 [=*Rhinobrissus pyramidalis, *Rhinobryssus pyramidalis]F66
                  |    |--R. hemiasteroides Agassiz 1879SM15
                  |    `--R. tumulus McNamara 1982SM15
                     `--+--+--Migliorinia Checchia-Rispoli 1942SS05, F66
                        |  |    `--*M. migiurtina Checchia-Rispoli 1942F66
                        |  |--+--Pycnolampas Agassiz & Clark 1907SS05, F66
                        |  |  |    `--*P. oviformis Agassiz & Clark 1907F66
                        |  |  `--Arcaechinus Kier 1957SS05, F66
                        |  |       `--*A. auraduensis Kier 1957F66
                        |  `--+--Palaeobrissus Agassiz 1883SS05, F66 [=Palaeobryssus Meissner 1903F66]
                        |     |    `--*P. hilgardi Agassiz 1883 [=*Palaeobryssus hilgardi]F66
                        |     `--+--Paleotrema Koehler 1914SS05, F66 [=Palaeotrema (l. c.)F66]
                        |        |    `--*P. loveni (Agassiz 1881) [=Palaeotropus loveni]F66
                        |        `--+--Palaeotropus Lovén 1872SS05, F66
                        |           |    `--*P. josephinae Lovén 1872F66
                        |           `--ScrippsechinusSS05
                        `--+--Lissospatangus Mortensen 1950SS05, F66
                           |    `--*L. hirsutus Mortensen 1950F66
                           `--+--Spatagobrissus Clark 1923SS05, F66
                              |    `--*S. mirabilis Clark 1923F66
                                 |  `--+--Atelospatangus Koch 1885SS05, F66 (see below for synonymy)
                                 |     |    |--*A. transilvanicus Koch 1885F66
                                 |     |    |--A. giardinalei (Oppenheim 1899) (see below for synonymy)F66
                                 |     |    `--A. magnus Szörényi 1963SG93
                                 |     `--+--Hemipatagus Desor 1858SS05, F66
                                 |        |    |--*H. hoffmanni (Goldfuss 1826) [=Spatangus hoffmanni, Maretia hoffmanni]F66
                                 |        |    `--H. formosusH79
                                 |        `--+--Hemimaretia Mortensen 1950SS05, F66
                                 |           |    `--*H. elevata (Döderlein 1907) [=Maretia elevata]F66
                                 |           `--Pseudolovenia Agassiz & Clark 1907SS05, F66
                                 |                `--*P. hirsuta Agassiz & Clark 1907F66
                                    |  `--+--Gualtieria Agassiz 1847SS05, F66 (see below for synonymy)
                                    |     |    |--*G. (Gualtieria) orbignyana Agassiz 1847F66
                                    |     |    `--G. (Blaviaster Lambert 1920) [=Temnaster Lambert 1912 non Verrill 1894]F66
                                    |     |         `--G. (*B.) grossouvrei (Lambert 1912) [=Temnaster grossouvrei]F66
                                    |     `--+--SpatangusSS05
                                    |        `--+--Granopatagus Lambert 1914SS05, F66 (see below for synonymy)
                                    |           |    `--*G. lonchophorus (Meneghini in Desor 1858) (see below for synonymy)F66
                                    |           `--Plethotaenia Clark 1917SS05, F66
                                    |                `--*P. spatangoides (Agassiz 1883) (see below for synonymy)F66
                                    `--+--+--+--Mazzettia Lambert & Thiéry 1915SS05, F66 (see below for synonymy)
                                       |  |  |    `--*M. pareti (Manzoni 1878) [=Maretia pareti, *Manzonia pareti]F66
                                       |  |  `--Gymnopatagus Döderlein 1901SS05, F66
                                       |  |       |--*G. valdiviae Döderlein 1901 [=Eupatagus (*Gymnopatagus) valdiviae]F66
                                       |  |       `--G. antillarum [=Eupatagus (Gymnopatagus) antillarum]F66
                                       |  `--+--Elipneustes Koehler 1914SS05, F66 (see below for synonymy)
                                       |     |    `--*E. denudatus (Koehler 1914) [=Eurypneustes denudatus]F66
                                       |     `--Linopneustes Agassiz 1881SS05, F66
                                       |          |--*L. murrayi (Agassiz 1873) [=Palaeopneustes murrayi]F66
                                       |          `--L. longispinus Agassiz 1878SS05
                                       `--+--Eupatagus Agassiz 1847SS05, F66 (see below for synonymy)
                                          |    |--*E. valenciennesi Agassiz 1847 (see below for synonymy)F66
                                          |    |--*Pseudopatagus’ cruciatus Pomel 1885F66
                                          |    |--*Megapatagus’ franciscanus Sánchez Roig 1953F66
                                          |    |--E. greyiH79
                                          |    |--*Zanolettiaster’ herrerae Sánchez Roig 1952F66
                                          |    |--‘Macropneustes’ lefebvrei DeLoriol 1881 [=*Heterospatangus lefebvrei]F66
                                          |    |--‘Euspatangus’ libycus de Loriol 1881 [=*Perispatangus libycus]F66
                                          |    `--‘Metalia’ melitensis Gregory 1891 [=*Melitia melitensis]F66
                                             `--+--Platybrissus Grube 1865SS05, F66 [=Platybryssus Meissner 1903F66]
                                                |    `--*P. roemeri Grube 1865 [=*Platybryssus roemeri]F66
                                                `--+--Eurypatagus Mortensen 1948SS05, F66
                                                   |    `--*E. ovalis Mortensen 1948 [=Platybrissus (*Eurypatagus) ovalis]F66
                                                   `--Paramaretia Mortensen 1950SS05, F66
                                                        `--*P. multituberculata Mortensen 1950F66

Atelospatangus Koch 1885SS05, F66 [incl. Lambertia Oppenheim 1899 non Souverbie 1869F66, Oppenheimia Cossmann 1900F66]

Atelospatangus giardinalei (Oppenheim 1899) [=*Lambertia giardinalei, *Oppenheimia giardinalei]F66

Elipneustes Koehler 1914SS05, F66 [incl. Eurypneustes Koehler 1914 non Duncan & Sladen 1882F66]

Eupatagus Agassiz 1847SS05, F66 [=Euspatangus Cotteau 1869F66, Koilospatangus Lambert 1906F66; incl. Heterospatangus Fourtau 1905F66, Megapatagus Sánchez Roig 1953F66, Melitia Fourtau 1913F66, Perispatangus Fourtau 1905F66, Pseudospatagus Pomel 1885F66, Zanolettiaster Sánchez Roig 1952F66]

*Eupatagus valenciennesi Agassiz 1847 [=*Euspatangus valenciennesi, *Koilospatangus valenciennesi]F66

Granopatagus Lambert 1914 [incl. Concophorus Laube 1869, Conchophorus (l. c.), Lonchophorus Pomel 1883 nec Germar 1824 nec Studer 1880]F66

*Granopatagus lonchophorus (Meneghini in Desor 1858) [=Spatangus lonchophorus]F66

Gualtieria Agassiz 1847SS05, F66 [=Gaultieria (l. c.)F66, Gualteria (l. c.)F66, Gualtiera (l. c.)F66]

Mazzettia Lambert & Thiéry 1915SS05, F66 [=Manzonia Pomel 1883 nec Garov. 1866 (ICBN) nec Brusina 1870F66

Metalia Gray 1855SS05, F66 [incl. Metaliopsis Fourtau 1913F66, Prometalia Pomel 1883F66, Xanthobrissus Agassiz 1863F66]

Nacospatangus (Pseudomaretia Koehler 1914) [incl. Gonimaretia Clark 1917, Lonchophorus Studer 1880 non Germar 1824]F66

Plagiobrissus Pomel 1883SS05, F66 [=Plagionotus Agassiz & Desor 1847 non Mulsant 1842F66, Plagiostomus d’Orbigny 1854 non Hermansen 1847F66]

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

*Plethotaenia spatangoides (Agassiz 1883) [=Macropneustes spatangoides, Meoma (*Plethotaenia) spatangoides]F66

*Rhabdobrissus jullieni Cotteau 1889 [=Plagiobrissus (*Rhabdobrissus) jullieni, *Rhabdobryssus jullieni]F66

*Type species of generic name indicated


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[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.

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[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).

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[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.

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

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