CLADOCOCCIDAE 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 : 431-433

Family CLADOCOCCIDAE Haeckel, 1862

Cladococcida Haeckel, 1862: 238, 364 [as a family]. — Mivart 1878: 177 [as a subdivision].

Arachnosphaerida Haeckel, 1862: 238, 354 [as a tribe]; Haeckel 1882: 454 [as a tribe]; Haeckel 1887: 208.

Cladococcidae – Pantanelli 1880: 46.

Diplosphaerida Stöhr, 1880: 86 [nomen dubium, as a family]. — Haeckel 1882: 451 [as a tribe]; Haeckel 1887: 208. — Schröder 1909: 16 [as a rank between subfamily and genus].

Lychnosphaerida Haeckel, 1882: 452 [as a tribe].

Astrosphaerida Haeckel, 1887: 55, 206 (not 1882: 449). — nec Wisniowski 1889: 684. — Bütschli 1889: 1952 [as a family]. — nec Rüst 1892: 133. — Cayeux 1894: 205. — nec Hill & Jukes- Browne 1895: 605-606. — Schröder 1909: 2 [as a family]. — Mast 1910: 155 [as a family]. — Anderson 1980: 3, 5, 19 [as a family]; Anderson 1981: 248, 351, 360, 368 [as a family]; Anderson 1983: 23 [as a family].

Caryommida Haeckel, 1887: 208, 265 [nomen dubium, as a subfamily]. — Schröder 1909: 17 [as a subfamily].

Astrosphaeridae – Haecker 1908: 435. — Popofsky 1908: 211; 1912: 93. — Poche 1913: 207. — Enriques 1932: 982. — Campbell & Clark 1944a: 15; 1944b: 11. — Clark & Campbell 1945: 16. — Deflandre 1953: 416. — Campbell 1954: D60. — Dogiel & Reshetnyak 1955: 32. — Chediya 1959: 94. — Orlev 1959: 437. — Hollande & Enjumet 1960: 72-73. — Mamedov 1973: 49. — Pessagno 1976: 42. — Tan & Tchang 1976: 228. — BjØrklund 1976: 119. — Dumitrica 1979: 20. — Tan & Su 1982: 146. — van de Paverd 1995: 77. — Chen & Tan 1996: 150. — Tan 1998: 146. — De Wever et al. 2001: 108. — Afanasieva et al. 2005, S275. — Afanasieva & Amon 2006: 111. — Matsuzaki et al. 2015: 13. — Chen et al. 2017: 106.

Arachnosphaerinae – Mast 1910: 173 ( sensu emend. ). — Popofsky 1912: 104. — Campbell 1954: D66. — Tan & Tchang 1976: 231. — Petrushevskaya 1979: 108.

Astrosphaerinae – Campbell 1954: D60. — Tan & Tchang 1976: 228.

Caryomminae – Chediya 1959: 100 [nomen dubium]. Arachnosphaeridae – Petrushevskaya 1979: 106. — Dumitrica 1984: 99.

Cladococcid type – Anderson 1983: 168.

Astrosphaerins [sic] – Casey 1993: 253.

TYPE GENUS. — Cladococcus Müller, 1856: 485 [type species by monotypy: Cladococcus arborescens Müller, 1856: 485 ].

INCLUDED GENERA. — Arachnosphaera Haeckel, 1861a: 804 (= Arachnosphaerella with the same type species). — Arachnospongus Mast, 1910: 56. — Cladococcus Müller, 1856: 485 (= Cladococcalis with the same type species; Anomalacantha n. syn., Cladococcodes synonymized by Mast 1910; 158; Cladococcurus n. syn., Porococcus n. syn.). — Diplosphaera Haeckel, 1861a: 804 (= Diplosphaeromma with the same type species; Astrosphaera , Diplosphaerella , Leptosphaera , Leptosphaerella , synonymized by Hollande & Enjumet 1960: 116; Astrosphaerella n. syn., Astrospongus , Drymosphaeromma n. syn., Leptosphaeromma n. syn.). — Haeckeliella Hollande & Enjumet, 1960: 119 . — Lychnosphaera Haeckel, 1882: 452 (= Rhizoplegmidium n. syn., Rhizospongus n. syn., Thalassoplegma n. syn.).

NOMINA DUBIA. — Acanthospongus, Arachnopegma , Arachnopila , Arachnosphaeromma , Astrosphaeromma , Caryomma , Cladococcinus , Drymosphaera, Drymosphaerella , Elaphococcus , Elaphococcinus , Elaphococculus, Hexacladus , Rhizoplegma , Rhizoplegmarium , Spongopila .

JUNIOR HOMONYM. — Heteracantha Mast, 1910 (= Anomalacantha ) nec Brullé, 1834.

DIAGNOSIS. — Cladococcoidea with radial spines and/or other ornaments. Endoplasm is distributed from the center to the outer side of the first spherical cortical shell. Tens to hundreds of brownish grains, but not algal symbionts, are found scattered around the endoplasm in some genera with innumerous pseudopodia radiate throughout. In the “ Elaphococcinus ”-form of Cladococcus , a large reddish to brown endoplasm and a surrounding milky-white endoplasm nearly fill the entirety of the shell’s area; such that, the distal ends of ramified radial spines are exposed. Algal symbionts also appear to be present.

The axopodial system is that of anaxoplastid-type; no axoplast and no bundles of axoneme are present. The intracapsular zone includes the nucleus, a thin endoplasmic reticulum, an empty area with axoneme strands and a main endoplasmic reticulum from the center to the outer part. The presence of an empty zone between the nucleus and the main endoplasmic reticulum zone is a significant protoplasmic character. The nucleus, as well as the empty zone with axoneme strands, is always encrusted in the innermost latticed shell.

STRATIGRAPHIC OCCURRENCE. — Early Middle Miocene-Living.

REMARKS

Cladococcidae is the oldest senior synonym of Astrosphaeridae . Classical papers considered the family Astrosphaeridae as a junior synonym of Actinommidae Haeckel 1882 , but this was obviously rejected by both morphological and molecular studies. Cladococcus and Haeckeliella are regularly found in Cenozoic sediments and rocks. Arachnosphaera , Diplosphaera , the “ Elaphococcus ” form of Cladococcus are often encountered in shallow surface plankton samples. Notwithstanding, all genera except “ Elaphococcus ” are not preserved, not even on surface sediments other than in exceptional cases. As the living Cladococcidae are large and have easily recognizable endoplasm, they are easily observed in seawater momentarily after plankton towing and at lower magnitudes of binocular microscopes. Owing to this facility, living specimens were analyzed in several papers. Living image were illustrated for Arachnosphaera (Anderson 1983: fig. 1.1.B?; Yuasa et al. 2009: fig. 1c; Suzuki & Aita 2011: fig. 4J), Diplosphaera ( Suzuki & Sugiyama 2001: figs 2.2- 2.4; Matsuoka 2007: fig. 2.e; 2017: figs 2.1, 2.2; Yuasa et al. 2009: fig. 1d; Suzuki & Aita 2011: fig. 4B-right) and the “ Elaphococculus ” form of Cladococcus ( Suzuki & Not 2015: fig. 8.8.23; Matsuoka 2017: figs 3.1, 3.2). Protoplasm and algal symbionts were documented by epi-fluorescent observation with DAPI dyeing or other dyeing methods for Arachnosphaera ( Ogane et al. 2014: pl. 1, figs 5-6), Diplosphaera ( Suzuki & Not 2015: fig. 8.8.19; Zhang et al. 2018: 19, fig. 4), and Cladococcus ( Zhang et al. 2018: 19, fig. 2). Fixed images with dyeing were published for Arachnosphaera ( Aita et al. 2009: pl. 6, fig. 4; pl. 19, fig. 4; pl. 21, fig. 2; pl. 22, fig. 3), Cladococcus ( Aita et al. 2009: pl. 8, 4), “ Elaphococculus ” of Cladococcus ( Aita et al. 2009: pl. 26, fig. 1) and Lychnosphaera ( Aita et al. 2009: pl. 7, fig. 1). Fine protoplasmic structures were illustrated for Arachnosphaera ( Hollande & Enjumet 1960: pl. 9, figs 11, 12; pl. 11, figs 1-7; pl. 12, figs 2-5; pl. 22, fig. 6; pl. 29, figs 1, 2), Diplosphaera ( Hollande & Enjumet 1960: pl. 12, fig. 6; pl. 23, fig. 2; pl. 26, fig. 2), Cladococcus ( Hollande & Enjumet 1960: pl. 6, fig. 12; pl. 12, fig. 1; pl. 15, figs 4-6; pl. 26, fig. 1), Haeckeliella ( Hollande & Enjumet 1960: pl. 15, figs 1-3) and Lychnosphaera ( Hollande & Enjumet 1960: pl. 30, figs 1, 2). Internal skeletal structures were illustrated for Arachnosphaera ( Cachon & Cachon 1972b: pl. 29, fig. b) and Haeckeliella ( Takahashi 1991: pl. 10, fig. 2; van de Paverd 1995: pl. 23, fig. 3). Although it is impossible to amend the taxonomy for the host, parasites were recognized in Cladococcus as Solenodinium ( Hollande & Enjumet 1955: fig. 8). The environmental RAD-III Clade of Not et al. (2007) collected from 200-400 m water depths in the tropical Pacific was specified as the Cladococcidae (originally Astrosphaeridae ) by Li & Endo (2020).

VALIDITY OF GENERA

Diplosphaera

The combinations of Astrosphaera and Astrosphaerella and of Leptosphaera and Leptosphaeromma have the same type species, respectively. Astrosphaera -specimens were repeatedly named Diplosphaera ( Hollande & Enjumet 1960: 116; Kozur & Mostler 1979: 12; Suzuki & Sugiyama 2001: 118). Observation of living representatives of Diplosphaera hexagonalis easily proves the distinguishing characteristics between Astrospongus , Diplosphaerella , Drymosphaeromma , Leptosphaera and Leptosphaeromma at intraspecies or species levels. The oldest available name is Diplosphaera among them.

Cladococcus

The translated description of Anomalacantha (originally Heteracantha ) by Mast (1910: 37) from German follows. “ Monosphaerids with main and secondary spines. Three-sided main spines ramified or not, secondary spines always dichotomized.Shell always very thick with funnel-shaped pores. ” Cladococcodes is characterized by ramified branches on the radial spine, regular pores with similar sizes on the cortical shell ( Campbell 1954: D63); and Cladococcurus by ramified branches on the radial spine, irregular pores with dissimilar sizes on the cortical shell ( Campbell 1954: D63). Regularity of pores and their sizes are not distinguishing features at generic level. Ramified and branched patterns of radial spines are too variable among specimens to be useful for determination of genus. As for the taxonomic value of funnel-shaped pores, we have never met such pores like that in Anomalacantha so far. The young Cladococcus -form often looks like the type species of Porococcus , and the latter genus is, thus, regarded as a young form. This genus may be also used as a collective group that is defined by an assemblage of species, or stages of organisms, that cannot be allocated with confidence to nominal genera (See the Glossary of ICZN 1999). If Porococcus is a collective group, this genus does not compete in priority with another genus-group (ICZN 1999: article 23.7.2); typification for this genus is not necessary (ICZN 1999: article 42.3.1) and a type species can be disregarded (ICZN 1999: article 67.14). The oldest available name is Cladococcus among synonyms.

Lychnosphaera

Lychnosphaera has an empty space just above the cortical shell. Thalassoplegma is also the case with a very narrow space. The oldest synonym is Leptosphaera ( Haeckel 1887: 452) .