ZONARIIDAE Haeckel, 1887

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 : 453-455

Family ZONARIIDAE Haeckel, 1887

sensu Dumitrica (1989)

Zonarida Haeckel, 1887: 604, 682-684 [as a family]. — Bütschli 1889: 1968 [as a family]. — Schröder 1909: 4 [as a family]. — Anderson 1983: 24 [as a family].

Pylonida Haeckel, 1882: 463 [as a family, nomen dubium]; 1884: 29 [as a family]; 1887: 604, 628-632 [as a family]. — Bütschli 1889: 1966 [as a family]. — nec Rüst 1892: 174. — Schröder 1909: 4 [as a family]. — Anderson 1983: 24 [as a family].

Streblemida [sic] Haeckel, 1887: 604 [nomen nudum] (= Streblonida) [as a family].

Streblonida Haeckel, 1887: 702-704 [nomen dubium, as a family]. — Bütschli 1889: 1969 [as a family]. — Schröder 1909: 4 [as a family]. — Anderson 1983: 25 [as a family].

Zonartidae [sic] – Popofsky 1912: 124 (= Zonariidae ).

Pylonidae [sic] – Popofsky 1912: 145-146 [nomen dubium] (= Pyloniidae ). — Chediya 1959: 154. — Tan & Tchang 1976: 259. — Tan & Su 1982: 159. — van de Paverd 1995: 184. — Chen & Tan 1996: 152. — Tan 1998: 249-252. — Tan & Chen 1999: 241-243.

Monozoniinae Campbell, 1954: D96 [nomen dubium].

Zonariidae – Poche 1913: 210. — Campbell 1954: D98.

Pyloniidae – Poche 1913: 210 [nomen dubium]. — Campbell 1954: D96. — Riedel 1967b: 295; 1971: 655. — Nakaseko et al. 1975: 171. — Nakaseko & Sugano 1976: 126. — Riedel & Sanfilippo 1977: 867. — Dumitrica 1979: 24; 1984: 101; Dumitrica 1989: 253, 258. — Petrushevskaya 1979: 110. — Kozur & Mostler 1979: 45-46. — Anderson 1983: 39. — Takahashi 1991: 90. — Hollis 1997: 43. — Boltovskoy 1998: 32. — Anderson et al. 2002: 1002- 1003. — De Wever et al. 2001: 148. — Afanasieva et al. 2005: S286. — Afanasieva & Amon 2006: 129. — Chen et al. 2017: 150.

Strebloniidae – Poche 1913: 210 [nomen dubium]. — Campbell 1954: D100. — Tan & Su 1982: 163. — Chen & Tan 1996: 152. — Tan 1998: 283. — Tan & Chen 1999: 267. — Chen et al. 2017: 162.

Pyloniinae – Campbell 1954: D96 [nomen dubium]. — De Wever et al. 2001: 150-151. — Afanasieva et al. 2005: S286. — Afanasieva & Amon 2006: 129.

Tetrapyloniinae Campbell, 1954: D97 [nomen dubium].

Strebloniinae – Campbell 1954: D100 [nomen dubium].

Zonaridae [sic] – Chediya 1959: 158 (= Zonariidae ).

Streblonidae [sic] – Chediya 1959: 160 [nomen dubium] (= Strebloniidae ).

TYPE GENUS. — Zonarium Haeckel, 1887: 684 [type species by subsequent designation ( Campbell 1954: D98): Zonarium octangulum Haeckel, 1887: 685 ] = junior subjective synonym of Tetrapyle Müller, 1859a: 154 [type species by monotypy: Tetrapyle octacantha Müller, 1859b: 33 ].

INCLUDED GENERA. — Larcidium Haeckel, 1887: 611 . — Pylozonium Haeckel, 1887: 659 . — Tetrapyle Müller, 1859a: 154 (= Tetrapylura with the same type species; Echinosphaera , Trizonium , Trizonaris , synonymized by Matsuzaki et al. 2015: 34;? Amphiaspis n. syn., Amphipylura n. syn., Dizonitis n. syn., Larnacantha n. syn., Octopylura n. syn., Pylonura n. syn., Schizomma, Tetrapylissa synonymized by Itaki 2009: 47, Zonarium n. syn., Zonidium n. syn., Zoniscus n. syn.).

NOMINA DUBIA. — Amphipylissa , Amphipyle , Amphipylonium , Dizonaris , Dizonium , Monozonaris , Monozonitis , Monozonium , Octopylissa , Octopyle , Pylonissa , Pylonium , Spongophorticium , Spongophortis , Streblonia , Stypophorticium , Tetrapylonium , Trizonitis .

INVALID NAME. — Stylophorticium.

NOMEN NUDUM. — Caryolithis.

DIAGNOSIS. — Larcospiroidea with systems of three elliptical girdles in 3 successive, perpendicular planes. First system (medullary shell) consists of a heteropolar microsphere with 12 pores, an antapical sagittal ring and two lateral arches. The following system repeats the first system previously described several times.

A Protoplasm is documented for Tetrapyle . The endoplasm occupies the shell and occasionally the external most girdle, depending on its growth stage. The central part of the endoplasm tends to be reddish in color and is surrounded by a light brown endoplasm. The nucleus is located inside the second pseudo-concentric shell in the sense of Suzuki & Zhang (2016). Several dozens of algal symbionts surround the endoplasm. Hundreds of pseudopodia radiate from the entire protoplasm. One axoflagellum, rarely two, extend on the side of the external most girdle’s polar region. Gelatinous material covers all skeletons.

STRATIGRAPHIC OCCURRENCE. — late Late Miocene-Living.

REMARKS

The family name “ Pyloniidae ” has been widely used, but it is impossible to retain this common family name as the “ Pyloniidae ” is based on an unillustrated type species. The oldest available name, “Zonarida”, was automatically selected as a valid family name. Tetrapyle is often confused with Phorticium (Phorticiidae) in practical work regardless of their fundamental differences at the superfamily level. According to Zhang & Suzuki (2017: 42) , Phorticium tends to possess numerous pillar beams between the pseudo-concentric shells. Otherwise, the presence or absence of the S1a-girdle is the only way to differentiate these two genera. Internal skeletal structure was illustrated for Tetrapyle ( Dumitrica 1989: pl. 15, figs 12, 13; Sugiyama et al. 1992: pl. 11, figs 1-4). Algal symbionts of Tetrapyle were identified as Brandtodinium nutricula ( Probert et al. 2014) . Living and protoplasmic images were captured for Tetrapyle ( Matsuoka et al. 2001: pl. 1, fig. 2; Suzuki & Aita 2011: fig. 4Q; Probert et al. 2014: S1, Vil 231; Suzuki & Not 2015: fig. 8.8.25; Matsuoka et al. 2017: appendix A; Zhang & Suzuki 2017 : figs 7.1-7.8). Fine protoplasmic structure for Tetrapyle was illustrated ( Hollande & Enjumet 1960: pl. 24, fig. 3).

VALIDITY OF GENERA

Tetrapyle

These synonymized genera with Tetrapyle are considered to be erected for different ontogenetic growth stages and different appearances differently oriented as such as the Amphitholidae and Pylodiscidae (Ogane & Suzuki 2009 : fig. 3; Zhang & Suzuki 2017 : fig. 3). The morphological terminology follows Zhang & Suzuki (2017) as briefly explained in the “Validity of genera” for the Amphitholidae . First it is necessary to determine the number of pseudo-concentric shells referred to the type-illustration in Haeckel (1887), the size of the specimens and supporting images for these available names . Amphiaspis looks to have two pseudo-concentric shells. Schizomma looks similar to Amphiapis; but the shell size is twice in Schizomma than in Amphiaspis , having three pseudo-concentric shells. The genera with three pseudo-concentric shells are Amphipylura , Larnacantha , Octopylura , Pylonura , Tetrapyle , Tetrapylissa , Trizonaris , Zonarium , Zonidium and Zoniscus . The typeillustration of Dizonites is very ambiguous but it presumably possess three pseudo-concentric shell in consideration of its size. Echinosphaera is the largest among the synonymized genera here but it is difficult to specify the number of its pseudo-concentric shells. According to Zhang & Suzuki (2017: fig. 3), the morphotype with three pseudo-concentric shells has nine possibilities by the outermost girdle (G1, G2 or G3) and the anatomical orientation under the absolute Cartesian coordinates (Lt-, Pl- and Sg-views). From the Lt-view, the opening (gate) encircled by the first girdle (S1a-girdle) directly attached on the microsphere (S1a) is visible; from the Pl-view, the microsphere and the first girdle look to be overlapped; and from the Sg-view, the body of the first girdle (girdle itself) is visible. Referred to Zhang & Suzuki (2017) , Amphiaspis is the Lt-view of the two pseudo-concentric shells with G3-girdle and Zonarium is the Sg-view of the four concentric shells with the G2-girdle. The remaining genera have three pseudo-concentric shells but the different view under the absolute Cartesian coordinates. Amphipylura , Trizonium , Octopylura and Tetrapylissa are the Pl-views. The former two genera have an incomplete G2-girdle whereas the latter two genera develop the complete G2-girdle. Pylonura , Larnacantha , Zoniscus and Zonidium ( Haeckel 1887: pl. 50, fig. 12) have also the Pl-view. The first one has an incomplete G3-girdle and the remaining three genera have the complete G3-girdle. Tetrapyle , Dizonitis and Schizomma are the Sg-view. The first two genera have the complete G1-girdle and the last genus has an incomplete G2-girdle. In consideration of the specimen’s orientations and their growth stages, the type-illustrations of these genera are derived from several limited species within the same genus. The oldest available name among them is Tetrapyle . Amphiaspis is possible to be regarded as a collective name for the Zonariidae for practical usage.

Phylogenetic Molecular Lineage indet.

REMARKS

In contrast to the clear results for the subdivision of Lineages in Sandin et al. (2021), it was nearly impossible to clearly define the morphological “commonalities” in each Lineage. This is due to the fact that the superfamilies and families categorized here cannot be classified into known Lineages.