ETHMOSPHAERIDAE Haeckel, 1862

Suzuki, Noritoshi, Caulet, Jean-Pierre & Dumitrica, Paulian, 2021, A new integrated morpho- and molecular systematic classification of Cenozoic radiolarians (Class Polycystinea) - suprageneric taxonomy and logical nomenclatorial acts, Geodiversitas 43 (15), pp. 405-573 : 430-431

Family ETHMOSPHAERIDAE Haeckel, 1862

Ethmosphaerida Haeckel, 1862: 238, 346 [as a family]; Haeckel 1882: 448 [as a tribe]; Haeckel 1887: 61 ( sensu emend. ) [as a subfamily]. — Mivart 1878: 179 [as a subsection]. — Dunikowski 1882: 184 [as a tribe]. — Wisniowski 1889: 682. — Schröder 1909: 5 [as a subfamily].

Monosphaerida Haeckel, 1862: 230 [nomen nudum, above Cladococcida]. — Stöhr 1880: 85. — Dreyer 1913: 5 [as a family].

Heliosphaerida Haeckel, 1862: 238, 348 [as a tribe]; Haeckel 1882: 450 [as a tribe].

Ethmosphaeriden – Haeckel 1865: 366 [as a family].

Etmosphaeridae – Claus 1876: 159. — Loeblich & Tappan 1961: 221. — Kozur & Mostler 1979: 23 ( sensu emend. ). — Dumitrica 1984: 95. — Takahashi 1991: 61. — Tan 1998: 95 [as a subfamily].

Monosphaeridae – Zittel 1876-1880: 119 [nomen nudum].

Triosphaeria Haeckel, 1882: 452 [nomen nudum, as a subfamily].

Monosphaeria – Haeckel 1882: 448 [nomen nudum, as a subfamily]. — Dunikowski 1882: 184.

Liosphaerida Haeckel, 1887: 59 [as a family] (not 1882: 449). — Wisniowski 1889: 682 [as a family]. — Rüst 1892: 133. — Bütschli 1889: 1948 [as a family]. — Cayeux 1894: 204 [as a family]. — Hill & Jukes-Browne 1895: 605. — Schröder 1909: 5 [as a family]. — Anderson 1983: 23 [as a family].

Cenodiscida Haeckel, 1887: 409-411 [nomen nudum, as a family]. — Bütschli 1889: 1958 [as a family]. — nec Rüst 1892: 161 [as a family]. — Schröder 1909: 3 [as a family]. — Anderson 1983: 24 [as a family].

Zonodiscida Haeckel, 1887: 411 [nomen nudum, as a subfamily. — Schröder 1909: 39 [as a subfamily].

Larcarida Haeckel, 1887: 604, 605-606 [nomen nudum, as a family]. — Bütschli 1889: 1965 [as a family]. — Schröder 1909: 4 [as a family]. — Anderson 1983: 24 [as a family].

Cenolarcida Haeckel, 1887: 606 [nomen nudum, as a subfamily]. — Schröder 1909: 52 [as a subfamily].

Coccolarcida Haeckel, 1887: 606, 610 [nomen nudum, as a subfamily]. — Schröder 1909: 52 [as a subfamily].

Liosphaeriden – Haecker 1907: 118 [as a family]. — Orlev 1959: 430.

Liosphaeridae – Popofsky 1908: 206; Popofsky 1912: 82. — Enriques 1932: 982. — Clark & Campbell 1942: 19; 1945: 5. — Campbell & Clark 1944a: 8. — Frizzell & Middour 1951: 9. — Campbell 1954: D48. — Chediya 1959: 68. — Orlev 1959: 430. — Hollande & Enjumet 1960: 70. — Dieci 1964: 184. — Nakaseko & Sugano 1976: 119. — Tan & Tchang 1976: 225. — Tan & Su 1982: 136. — Blueford 1988: 247. — Chen & Tan 1996: 150. — Tan 1998: 95. — Chen et al. 2017: 81 ( sensu emend. ).

Larcaridae – Popofsky 1908: 229 [nomen nudum]. — Enriques 1932: 985. — Clark & Campbell 1942: 50; 1945: 27. — Chediya 1959: 151.

Monosphaerinae – Mast 1910: 155. — Popofsky 1912: 95.

Larcariidae – Poche 1913: 209 [nomen dubium]. — Kozur & Mostler 1979: 45.

Cenodiscidae – Poche 1913: 209 [nomen dubium]. — Campbell 1954: D77. — Orlev 1959: 441. — Chediya 1959: 120. — Nakaseko & Sugano 1976: 125. — Amon 2000: 40.

Liosphaerinae – Clark & Campbell 1942: 19. — Frizzell & Middour 1951: 9. — Campbell 1954: D48.

Ethmosphaerinae – Clark & Campbell 1942: 19. — Campbell & Clark 1944a: 8. — Clark & Campbell 1945: 6. — Campbell 1954: D48. — Chediya 1959: 68. — Loeblich & Tappan 1961: 222. — Dieci 1964: 64. — Kozur & Mostler 1979: 25.

Coccolarcinae – Clark & Campbell 1942: 50 [nomen dubium]; Clark & Campbell 1945: 28. — Campbell 1954: D95. — Chediya 1959: 152.

Heliosphaerinae – Campbell 1954: D62. — Tan & Tchang 1976: 230.

Cenodiscinae – Campbell 1954: D77 [nomen dubium].

Zonodiscinae – Chediya 1959: 12 [nomen dubium].

Cenolarcinae – Chediya 1959: 151 [nomen dubium].

Macrosphaeridae Hollande & Enjumet, 1960: 48 , 69, 114, 120 [nomen nudum]. — Cachon & Cachon 1972c: 297. — Petrushevskaya 1975: 571. — Anderson 1983: 51, 66. — Cachon & Cachon 1985: 286 [as a superfamily]. — Suzuki & Sugiyama 2001: 138.

Cyrtidosphaeridae Cachon & Cachon, 1972c: 293 ; Cachon & Cachon 1985: 288.

Ethmosphaerini – Kozur & Mostler 1979: 25 [as a tribe].

Cenodiscini – Kozur & Mostler 1979: 26 [nomen nudum].

Cyrtidosphaerid-type – Anderson 1983: 169.

Liosphaeracea – Göke 1984: 38 [as a subfamily].

TYPE GENUS. — Ethmosphaera Haeckel, 1861a: 802 [type species by monotypy: Ethmosphaera siphonophora Haeckel, 1861a: 802].

INCLUDED GENERA. — Cyrtidosphaera Haeckel, 1861a: 803 . — Ethmosphaera Haeckel, 1861a: 802 (= Ethmosphaerella with the same type species; Monosphaera n. syn.). — Haplosphaera Hollande & Enjumet, 1960: 114 . — Heliosphaera Haeckel, 1861a: 803 (= Heliosphaerella with the same type species). — Liosphaera Haeckel, 1887: 76 (= Melitomma with the same type species; Craspedomma n. syn.).

NOMINA DUBIA. — Briorradiolites, Cenodiscus , Cenolarcus , Coccolarcus , Ethmosphaeromma , Heliosphaeromma , Larcarium , Phormosphaera , Rhodosphaera , Rhodosphaerella , Rhodosphaeromma , Zonodiscus .

DIAGNOSIS. — Cladococcoidea without any other ornaments such as radial spines, excepting by-spines. Endoplasm is variable from transparent ( Haplosphaera ) to milky-white ( Cyrtidosphaera ) Size of the endoplasm is different in very large ( Cyrtidosphaera ) or very small forms ( Haplosphaera and Heliosphaera ).

STRATIGRAPHIC OCCURRENCE. — Late Oligocene-Living.

REMARKS

This family used to be called Liosphaeridae , but the oldest senior synonym is Ethmosphaeridae . Since both Liosphaeridae and Ethmosphaeridae were used valid names , the correct valid name is Ethmosphaeridae . The Ethmosphaeridae are distinguishable from the Cladococcidae by virtue of their radial spines and other ornaments. Cenosphaera has long been characterized as the genus having a single latticed cortical shell without any internal structure, but the usage of Cenosphaera was also problematic due to the following conditions: 1) The genus name was applied to spherical radiolarians whose internal structure might have been absent due to poor preservation. More specifically, very large numbers of Cenosphaera species were described from thin sections, increasing the problem of nomina dubia species in this genus; 2) It is practically necessary to separate in a “provisional group” such specimens as a tentative genus; 3) As explained in detail in the remarks of Haliommidae , the name-bearing type specimen of Cenosphaera has three concentric shells. Thus, it does not reconcile with the practical use of the representative genus; 4) Following the Code, the real Cenosphaera is a junior synonym of Haliomma ; 5) The true genus having a single latticed cortical shell may or may not exist; 6) No appropriate genus has been proposed to inherit the widely accepted concept of Cenosphaera ; and 7) We previously thought to use Monosphaera as a replacement name of the single-shelled “ Cenosphaera ” but this would have created another problem as to the nomen dubium status of the type species of Monosphaera. Considering these reasons, the definition of Ethmosphaera is expanded so as to include the widely accepted concept of Cenosphaera in the catalogue. In accordance with the Code, it was not possible to keep Cenosphaera as is (see details in the Remarks for Haliommidae ). Classic Cenosphaera , classified as Ethmosphaera in this catalogue, may be misidentified as spherical radiolarians whose radial spines were broken off, Nanina (Hexacromyidae) , Hollandosphaera (Hollandosphaeridae) , Cyrtidosphaera (Ethmosphaeridae) , Haplosphaera (Ethmosphaeridae) , young forms of Liosphaera (Ethmosphaeridae) , young forms of Cromyosphaera (Haliommidae) , Haliomma (Haliommidae) , Haliommantha (Haliommidae) and Entapium (Entapiidae) when the internal structure is lost. Moreover, it is impossible to differentiate these genera with scanning electron photos or with light microscopic photos under very shallow focus depths. The genus member of Ethmosphaeridae is solely determined by the lack of a skeletal structure within the large cortical shell and has not been supported by any molecular phylogenetic data. It is fundamentally impossible to discard morphospecies with a retrograde development of internal structures from the “true” Ethmosphaeridae . A living image for Cyrtidosphaera was obtained ( Kurihara et al. 2006: figs 4.1, 4.2; Suzuki & Not 2015: fig. 8.10.7). Protoplasm and algal symbionts were documented by epi-fluorescent observation with DAPI dyeing for Cyrtidosphaera ( Zhang et al. 2018: 19, fig. 3). Fine protoplasmic structure was illustrated for “ Cenosphaera ” ( Hollande & Enjumet 1960: pl. 1, figs 1-6; pl. 3, figs 1-5, 8-14; pl. 6, figs 1-4, 6; pl. 31, figs 1-7; pl. 32, figs 1, 2; Cachon & Cachon 1972b: fig. 1) and Heliosphaera ( Hollande & Enjumet 1960: pl. 55, figs 4, 5), but there are concerns regarding whether they were correctly identified as the same genus due to the fact that the axoplast system is often quite different among their assigned species.

VALIDITY OF GENERA

Ethmosphaera

Genera with one cortical shell are synonymized herein. The oldest available name is selected.

Liosphaera

Craspedomma is characterized by irregular pores on both inner and outer cortical shells ( Campbell 1954: D48). Irregularity of pores is an intraspecies or intraspecies difference in such kinds of cortical shells in any families.