PANARTIDAE 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 : 422-424

Family PANARTIDAE Haeckel, 1887

Panartida Haeckel, 1887: 288, 375 [as a family]. — Bütschli 1889: 1957 [as a family]. — nec Rüst 1892: 161. — Schröder 1909: 3 [as a family]. — Anderson 1983: 23.

Artiscida Haeckel, 1882: 462 [nomen dubium, as a subfamily]; 1887: 288, 354 [as a family]. — Bütschli 1889: 1956 [as a family]; Schröder 1909: 3 [as a family]. — Anderson 1983: 23.

Cyphinida Haeckel, 1882: 462 [nomen dubium, as a subfamily]; 1887: 288, 359-360 [as a family]. — Bütschli 1889: 1956 [as a family]. — nec Rüst 1892: 160.— Schröder 1909: 3 [as a family]. — Anderson 1983: 23.

Zygartida Haeckel, 1882: 462 [nomen dubium, as a family]; 1884: 29 [as a family]; 1887: 288, 391-392 [as a family]. — Bütschli 1889: 1958 [as a family]. — Schröder 1909: 3 [as a family]. — Anderson 1983: 23.

Artisoida – Haeckel 1887: 288 [nomen dubium, as a family].

Druppulida Haeckel, 1887: 288, 306 [nomen dubium, as a family]. — Bütschli 1889: 1955 [as a family]. — Schröder 1909: 3 [as a family]. — Anderson 1983: 23 [as a family].

Zygocampida Haeckel, 1887: 392 [nomen dubium, as a subfamily].

Desmocampida Haeckel, 1887: 392 [nomen dubium, as a subfamily].

Phacopylida Dreyer, 1889: 28 [nomen dubium, as a subfamily].

Druppuliden – Haecker 1907: 119 [nomen dubium].

Druppulidae – Popofsky 1908: 219 [nomen dubium]; 1912: 114; Clark & Campbell 1942: 32; 1945: 19. — Campbell & Clark 1944a: 18. — Frizzell & Middour 1951: 20. — Campbell 1954: D69. — Orlev 1959: 440. — Chediya 1959: 108. — Nakaseko & Sugano 1976: 122. — Tan & Su 1982: 149. — Blueford 1988: 248. — Chen & Tan 1996: 151. — Tan 1998: 189. — Tan & Chen 1999: 195. — Chen et al. 2017: 134.

Cyphinidae – Popofsky 1908: 220-221 [nomen dubium]. — Chediya 1959: 116. — Tan & Tchang 1976: 237. — Tan & Su 1982: 150. — Chen & Tan 1996: 151. — Tan 1998: 198. — Tan & Chen 1999: 201. — Chen et al. 2017: 137.

Panartidae – Popofsky 1908: 221; 1912: 121. — Campbell, 1954: D75. — Chediya 1959: 118. — Tan & Tchang 1976: 238. — Tan 1998: 200.

Pipettarida Schröder, 1909: 37 [as a subfamily].

Artiscidae – Poche 1913: 209 [nomen dubium]. — Deflandre 1953: 421 ( sensu emend. ). — Campbell 1954: D74. — Chediya 1959: 115. — Petrushevskaya 1975: 577. — Dumitrica 1979: 22. — Petrushevskaya 1979: 114-115.

Zygartidae – Campbell & Clark 1944a: 23 [nomen dubium]. — Campbell 1954: D76. — Chediya 1959: 119. — Chen & Tan 1996: 151.

Zygartinae – Campbell 1954: D76 [nomen dubium].

Desmocampinae – Campbell 1954: D76. — Chediya 1959: 119 [nomen dubium].

Cyphantidae Campbell, 1954: D74 [junior homonym].

Zygocampinae – Chediya 1959: 120 [nomen dubium].

Cyphantellidae – Loeblich &Tappan 1961: 223 [junior homonym].

Artiscinae – Riedel 1967b: 294 ( sensu emend. ) [nomen dubium]; 1971: 652. — Riedel & Sanfilippo 1971: 1587. — Petrushevskaya & Kozlova 1972: 521. — Nakaseko et al. 1975: 169. — Riedel & Sanfilippo 1977: 863. — Sakai 1980: 705. — Sanfilippo & Riedel 1980: 1009. — Anderson 1983: 37-38. — Dumitrica 1984: 97. — Sanfilippo et al. 1985: 655. — Takahashi 1991: 79. — De Wever et al. 2001: 123. — Afanasieva et al. 2005: S288. — Afanasieva & Amon 2006: 131.

Artistidae [sic] – Nakaseko & Sugano 1976: 122 [nomen dubium] (= Artiscidae ).

TYPE GENUS. — Panartus Haeckel, 1887: 376 [type species by subsequent designation ( Campbell 1954: D76): Panartus tetrathalamus Haeckel, 1887: 378 ] = junior subjective synonym of Didymocyrtis Haeckel, 1862: 445 [type species by absolute tautonomy: Haliomma didymocyrtis Haeckel, 1861a: 816 ].

INCLUDED GENERA. — Cannartus Haeckel, 1882: 462 (= Cannartidissa , Cannartiscus , Pipetta , Pipettaria , Pipettella synonymized by Riedel 1971: 652; Druppula n. syn., Druppuletta n. syn.). — Diartus Sanfilippo & Riedel, 1980: 1010 . — Didymocyrtis Haeckel, 1862: 445 (= Artidium n. syn., Cyphinura n. syn., Cyphocolpus n. syn., Desmartus n. syn., Ommatocampula n. syn., Panaromium n. syn., Panartus, Panartella synonymized by Sanfilippo & Riedel 1980: 1010, Panartidium n. syn., Panartissa n. syn., Panartura n. syn., Peripanartium n. syn., Peripanartula n. syn., Peripanartus n. syn., Peripanicula n. syn.). — Spongolivella Dumitrica, 2021: 2 .

INVALID NAME. — Artocarpium.

NOMINA DUBIA. — Artiscium , Artiscus , Cannartidella , Cannartidium , Caryodruppula , Cladospyris , Cromyocarpus , Cromyodruppa , Cromyodruppium , Cypassis , Cyphantella , Cyphantissa , Cyphinidium , Cyphinidoma , Cyphinidura , Cyphinoma , Cyphinus , Cyphonium , Desmocampe, Didymospyris, Diplellipsis , Druppocarpetta , Druppocarpissa , Druppocarpus , Haeckelocyphanta , Ommatacantha , Ommatartus , Ommatocorona , Ommatocyrtis , Ommatospyris , Panarelium , Panarium , Panartoma , Panicidium , Panicium , Peripanarium , Peripanicea , Peripanicium, Phacopyle , Prunocarpetta , Prunocarpilla , Prunocarpus , Prunulissa , Spongoliva , Spongolivetta , Spongolivina , Stylartella , Stylartura , Stylartus , Zygartus , Zygocampe .

NOMEN NUDUM. — Ommatocoryne.

JUNIOR HOMONYMS. — Cyphanta Haeckel, 1887 nec Walker, 1865 (= Cyphantella ).

DIAGNOSIS. — Lithocyclioidea with twin oval balloon-shaped latticed shells. The equatorial plane of the flattened double medullary shell is vertical to the longest axis of the fully-grown shell. Balloonshaped latticed shells, nearly all equal in size, are disposed along the equatorial plane. Radial beams emanating from the flattened double medullary shell extend across the equatorial plane to connect with the latticed shell.

A reddish endoplasm occupies the inner part of shell and a yellow-

ish red endoplasm encases them within the outermost shell. Algal symbionts are scattered inside and outside of the outermost shell. Nucleus is located inside the medullary shell. A long, robust axoflagellum extends from the longest axis of the fully-grown shell. The axoflagellum is perpendicular to the equatorial plane of the flattened double medullary shell.

STRATIGRAPHIC OCCURRENCE. — Early Oligocene-Living.

REMARKS

This family is easily distinguishable from any other of the spumellarian families by the virtue of its morphology closely resembling that of a drum. This family was once called either as “ Artiscinae ” or “Artiscinidae”, but this familiar name is unacceptable as it is based on an unillustrated type species. The genus Panartus and the family name Panartidae were used in several Chinese radiolarian monographs (e.g., Tan & Tchang 1976; Tan 1998) and there is no longer logical reason to retain the family name “ Artiscinae ” or “Artiscinidae.” Considering the genus Actinomma was significantly separated from the family Panartidae by molecular phylogeny analyses ( KrabberØd et al. 2011; Sandin et al. 2021). The Panartidae is presumed to be quite different from the Lithocycliidae ( Riedel & Sanfilippo 1981: fig. 12-6). The morphological changes of Panartidae genera were continually traced over the early Miocene at a morphospecies level ( Riedel & Sanfilippo 1971: pls 1C, 2C; Sanfilippo & Riedel 1980: text-fig. 1; Sanfilippo et al. 1985: 656). Their morphological changes were analyzed with quantitative methods ( Sachs & Hasson 1979; Kellogg 1980) and using more sophisticated mathematical methods ( Yoshino et al. 2019). The Panartidae are polycystines of particular importance to high resolution age determination so the criteria for identifying at specific level should be standardized ( Sakai 1980; Sanfilippo et al. 1985). However, the Panartidae have been carefully identified in mid-latitudes samples due to there being several undescribed species in the mid-latitudes of the North Pacific (e.g., “ Cannartus lineage” and “ Ommatartus lineage” in Sakai 1980), of the Southern Ocean (e.g., Lazarus 1992) and of the North Atlantic (Nishimura A. 1987). This suggests that the evolutionary history of Cannartus , Diartus and Didymocyrtis shown in Sanfilippo & Riedel (1980) never included other panartid species.

Many biological studies were carried out on living Didymocyrtis as they are commonly collected in plankton sampling.The relationship between their cytological structures was examined in detail ( Sugiyama & Anderson 1998a). Illustration of living forms were given for Didymocyrtis ( Matsuoka 1993b: pl. 3, figs 5, 6; 2017: figs 9.1, 9.2; Sugiyama & Anderson 1998a; text-figs 1-7, Takahashi et al. 2003: figs 3, 4; Suzuki & Aita 2011: fig. 4K; Probert et al. 2014: S1, SES 19; Suzuki & Not 2015: fig. 8.4.1, 8.8.5; Matsuoka et al. 2017: appendix A) and its internal skeletal structure was illustrated ( Anderson et al. 1986a: pl. 1, figs 3, 4; Sugiyama et al. 1992: pl. 4, fig. 7; Matsuoka 2009: fig. 3.20-3.24). Algal symbionts were documented by epi-fluorescent observation with DAPI dyeing for Didymocyrtis ( Zhang et al. 2018: 11, fig. 8). Algal symbionts of Didymocyrtis were identified as Brandtodinium nutricula by Probert et al. (2014).

VALIDITY OF GENERA

Cannartus

In addition to the synonymy published byRiedel (1971: 652), Druppula and Druppuletta are synonymized with Cannartus because the young form of Cannartus lacks the polar tubes as shown by the supporting images for these two genera. Cannartus was established in 1882 and all the other available genera listed here were published in 1887.

Didymocyrtis

All type species listed in the genera synonymy fall in the species conception of the Didymocyrtis -lineages of Sanfilippo et al. (1985: 656-660). Didymocyrtis , the oldest available name was published in 1862. All the other available names were published in 1882 and later.