THEOPERIDAE Haeckel, 1882

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 : 525-527

Family THEOPERIDAE Haeckel, 1882

sensu Suzuki emend. herein

Theoperida Haeckel, 1882: 435 [as a tribe]; 1887: 1313, 1325, 1354 [as a subfamily].

Rhopalocanida Haeckel, 1882: 437 [below tribe].

Artoperida Haeckel, 1882: 438 [as a tribe].

Theophaenida Haeckel, 1887: 1313, 1366, 1393 [as a subfamily].

Theoperinae – Clark & Campbell 1942: 81. — Campbell & Clark 1944b: 30. — Campbell 1954: D130, D132. — Chediya 1959: 216. — Petrushevskaya 1981: 231.

Theophaeninae – Chediya 1959: 219.

Theoperidae – Riedel 1967b: 296 ( sensu emend. ); 1971: 656. — Nakaseko & Sugano 1976: 130. — Riedel & Sanfilippo 1977: 870. — Anderson 1983: 40. — Sanfilippo et al. 1985: 666. — Blueford 1988: 244. — Takemura 1986: 43. — Takahashi 1991: 113. — Dumitrica 1995: 29. — Boltovskoy 1998: 33. — Cordey 1998: 118. —Sugiyama 1998: 233-234. — Takemura & Ling 1998: 159. — Kiessling 1999: 55. — Anderson et al. 2002: 1017. — De Wever et al. 2001: 282-283. — Suzuki H. et al. 2002: 180. — Suzuki & Gawlick 2003: 176. — Afanasieva et al. 2005: S297. — Afanasieva & Amon 2006: 146. — nec Suzuki H. et al. 2002: 105.

TYPE GENUS. — Theopera Haeckel, 1882: 436 [type species by subsequent designation ( Campbell 1954: D130): Theopera prismatica Haeckel, 1887: 1357 ] = junior subjective synonym of Rhopalocanium Ehrenberg, 1846: 385 [type species by subsequent monotypy: Rhopalocanium ornatum Ehrenberg, 1847: 55 ].

INCLUDED GENERA. — Clathropyrgus Haeckel, 1882: 439 . — Cyrtopera Haeckel, 1882: 439 (= Artopera Haeckel, 1882 nec Artopera Haeckel, 1887 , with the same type species).— Eusyringium Haeckel, 1882: 437 (= Eusyringartus with the same type species; Pterosyringium synonymized by Petrushevskaya 1981: 218). — Rhopalocanium Ehrenberg, 1846: 385 (= Artoperina n. syn., Dictyatractus n. syn., Rhopalatractus n. syn., Rhopalocyrtis n. syn., Theophaena n. syn.; Theopera synonymized by Petrushevskaya 1981: 232). — Stichopilidium Haeckel, 1887: 1438 .

INVALID NAME. — Sethornithium.

NOMINA DUBIA. — Hexalatractus , Sestrornithium .

JUNIOR HOMONYMS. — Artopera Haeckel, 1887 (= Artoperina ) nec Haeckel, 1882; Pteropilium Haeckel, 1887 (= Rhopalocyrtis) nec Haeckel, 1882.

DIAGNOSIS. — Theoperidae consist of three segmented Pterocorythoidea with a thick-walled spherical cephalis, a truncated conical or globular thorax, and a well-defined abdomen and/or conical tube.Two or three undulations rarely develop on the abdomen.Three wings or feet extend from the D- and double L-rods between the upper part of the thorax and the thoracic-abdominal boundary. The wings are rarely absent.

The cephalic initial spicular system consists of MB, A-, V-, D-, and double L-rods. The basal ring directly connects to the apical side end of MB, double L- and V-rods, forming four collar pores. In some members, an additional twin pore connected to D- and double Lrods is present on the apical side of the basal ring. The basal ring is largely free from the shell wall or is attached to the shell with D- and double L-rods. The basal ring sharply bends along the line with double L-rods. An A-rod is free in the cephalic cavity and forms an apical horn on the cephalis. A free A-rod in cephalic cavity is connected to an apical horn. A free D-rod, close to the MB, merges to the shell test. Little to nothing is known regarding the protoplasm.

STRATIGRAPHIC OCCURRENCE. — Early Paleocene-early Middle Miocene.

REMARKS

The concept of the Theoperidae differs considerably among publications. The most widely applied concept was defined by Riedel (1967b: 296). In the Theoperidae family, he included nassellarians with simple cephalic structures enclosed in a small spherical cephalis, and with a “cyrtid” shell differentiated along a heteropolar axis. The “probable polyphyletic” character, noticed by Riedel (1967b) for the Theoperidae , has been against regarded as monophyletic (e.g., Sugiyama 1998: 233-234). Due to the nomenclature act following the Code, the taxonomic name Theoperidae was transformed into the concept of the superfamily Eucyrtidioidea ( Petrushevskaya 1981: 200-202) . However, the “probable polyphyletic” possibility appears to have been lost. Nishimura (1990: 125-126) insisted on using Podocyrtidae instead of Theoperidae as Theopera is a junior synonym of Lithornithium . This explanation is not only nonsensical but also an incorrect nomenclatural act due to the nomen nudum status of Lithornithium . The concept of “ Podocyrtidae ” mixes one family of Lineage I ( Eucyrtidiidae in our catalogue) and five families of Lineage IV ( Lithochytrididae , Lophocyrtiidae , Pterocorythidae , Theocotylidae and Theoperidae in our catalogue). Thus, the concept of “ Podocyrtidae ” must be rejected. Besides a very broad concept of the Theoperidae ( Riedel 1967b; Nishimura 1990; Sugiyama 1998), the concept of “Theoperinae” adopted by Petrushevskaya (1981: 231) and of “ Theoperidae ” adopted by De Wever et al. (2001: 282- 283) is further limited by the morphological commonality to the type genus Theopera . However, their concepts are slightly different in so far as to include Pterocyrtidium (Rhopalosyringiidae) and Lamprotripus (Bekomidae) from Petrushevskaya (1981). Conversely, Lychnocanissa (originally Lychnocanoma ) and Lychnocanium (Lithochytrididae) are included in De Wever et al. (2001: 282). The improbable placement of these genera in the Theoperidae was explained in the remarks for Lamprotripus . Several species generally classified in Lychnocanissa (originally Lychnocanoma ) ( O’Connor 1997a: pl. 9, figs 9-12; 1; 1999: pl. 4, figs 11, 15) have a cephalic structure identical to that of the Theoperidae ; this is marked by the presence of many rods around the cephalic base that join the basal ring and shell test. For O’Connor (1997a; 1999) these species develop a tube-like porous skirt, which is common to the thorax of the Theoperidae genera. Lychnocanissa is a very large group. Nonetheless, this does not suggest that all Lychnocanissa belong to the Theoperidae .

The cephalic initial spicular system was reported for Cyrtopera ( Nishimura 1992: pl. 8, figs 2, 12; O’Connor 1999: pl. 4, figs 16-21), for some Lychnocanissa members ( O’Connor 1997a: pl. 9, figs 9-12; 1999: pl. 4, figs 11, 15), Rhopalocanium ( Nishimura 1990: fig. 27.10), and the “ Lychnocanissa ”-form of Rhopalocanium ( Nishimura 1992: pl. 9, figs 12 16; O’Connor 1999: pl. 4, figs 1-15). The aforementioned explanations have a common cephalic structure with a few exceptions ( Nishimura 1992: pl. 8, fig. 8).

VALIDITY OF GENERA

Rhopalocanium

Rhopalocyrtis and Pteropilium have the same type species. The genera listed here have common characters including two to three segments, inverted conical final segment, and wing-like lateral appendages that are directly connected with D- and L-rods. Apart from the final segment, the largest segment is cupola-shaped. Skeleton robust. In ignorance of the final segment, these genera may be subdivided into a two-segmented group ( Rhopalocanium , Artoperina , Dictyatractus , Rhopalatractus , Theopera , and Theophaena ) and a three-segmented group (Rhopalocyrtis). Apical horn on cephalis is variable, from very short ( Rhopalatractus , Theopera , and Theophaena ) to upwardly thickening hyaline cephalic wall ( Artoperina ), to very robust and long ( Rhopalocanium and Dictyatractus ). These continuous changes are insufficient for distinction at the genus level. The robustness and length of wing-like lateral appendages are also variable, but in a different way from the apical horn. Winglike lateral appendages form ridges on the test, and its distal part is free from the test. Free lateral appendages sometimes develop poreless or latticed web from the test. The following differences are summarized in the Atlas for each genus based on type and supporting images: thoracic ribs and base of free lateral appendages on upper abdomen in Dictyatractus and Rhopalocanium ; thoracic or thoracic to abdominal ribs and free lateral appendages with poreless web on upper abdomen or distal margin of abdomen in Rhopalatractus ; thoracic to abdominal ribs as base of free lateral appendages in Artoperina ; extended thoracic to abdominal ribs with very short lateral appendages from distal margin of abdomen in Theopera ; and ribs vertically passing through thorax and free lateral appendages originating from abdomen in Theophaena . As the base and free parts of the lateral appendages overlap among these genera, these characters are inappropriate for genus classification. All genera have three lateral appendages, except Theophaena . No real specimens have been reported for Theopera . The final segments can be divided into three types: slender, inverted, conical in shape and a straight extending terminal tube in Dictyatractus , Rhopalatractus , and Rhopalocanium ; inverted, conical in shape without opening in Theopera and Theophaena ; short, truncated, inverted, conical in shape with fenestrated aperture at end of final segment in Artoperina ; and short, truncated, slightly inverted, conical in shape with large aperture at end of final segment in Dictyatractus and Rhopalatractus . As described above, Dictyatractus and Rhopalatractus may have different final segments, in which case detailed differences in the final segment may represent intraspecific or infraspecific variation rather than genus differences.We have discarded all differences among all two-segmented genera described in previous studies ( Haeckel 1882, 1887; Petrushevskaya 1981; Kozlova 1999) from the genus criteria. The remaining genus is Rhopalocyrtis, which was defined by Bütschli (1882: 526) and translated as follows: “ Four segments. Short apical horn.Well-developed abdomen and lateral appendages oriented slightly downward and to posterior, such that they originate from continuous second and third segments, which are strongly connected. ” We consider this description to be similar to that of two-segmented genera, except for the number of segments in Rhopalocyrtis. Among these groups, the oldest available name is Rhopalocanium .

Incertae familiae nassellarians

INCLUDED GENUS. — Aspis Nishimura, 1992: 358 .

STRATIGRAPHIC OCCURRENCE. — Middle Paleocene-Late Paleocene.

REMARKS

The cephalic structure of Aspis was only shown in Hollis (2002: pl. 5, fig. 11a). This genus is characterized by the absence of internal dividers and has a very complex cephalic initial spicular system. Dumitrica (1973b: pl. 3, figs 2-4) identified this genus as Ceratocyrtis at the time.

Orphaned nassellarian family ranks

Calodictya Ehrenberg, 1847: 54 [invalid name, as a family]; 1876: 156. — Schomburgk 1847: 124, 126 [as a family].

Cyrtiida Haeckel, 1862: 237-238, 272-290 [invalid name, as a family]. — Zittel 1876-1880: 120 [as a group].

Dicyrtida Haeckel, 1862: 238, 280, 296 [invalid name, as a tribe]; 1887: 1192 [as a section between suborder and family]. — Zittel 1876-1880: 121 [rank unknown]. — Stöhr 1880: 99 [as a family]. — Poche 1913: 220 [as a super-superfamily]. — Popofsky 1913: 333 [as a section between suborder and family]. — Schröder 1914: 91, 100 [as a group between suborder and family]. — Chediya 1959: 199 [as a group between superfamily and family].

Polycyrtida Haeckel, 1862: 238, 280, 341 [invalid name, as a tribe]. — Zittel 1876-1880: 123 [rank unknown].

Zygocyrtida Haeckel, 1862: 238, 280, 291 [invalid name, as a tribe]. — Zittel 1876-1880: 121 [rank unknown]. — Stöhr 1880: 97 [as a family].

Cyrlidae [sic] – Claus 1876: 159 [invalid name] (= Cyrtidae ).

Cyrtiden – Hertwig 1879: 202-214 [invalid name, as a family].

Plectida Haeckel, 1882: 423 [invalid name, as a family]. — Lankester 1885: 850 [as a family].

Cyrtida – Haeckel 1882: 425 [invalid name, as a family]; 1884: 31 [as a family]. — Lankester 1885: 850 [as a family].

Dyocyrtida Haeckel, 1882: 430 [invalid name, as a subfamily].

Triocyrtida Haeckel, 1882: 434 [nomen nudum, as a subfamily]; 1887: 1313 [as a section between suborder and family].

Lophocorida Haeckel, 1882: 434 [invalid name, below a tribe].

Artocorida Haeckel, 1882: 437 [nomen nudum, as a tribe].

Tetracyrtida Haeckel, 1882: 437 [invalid name, as a subfamily].

Theophatnida Haeckel, 1882: 437 [nomen nudum, as a tribe].

Dyospyrida Haeckel, 1882: 441 [nomen nudum, as a subfamily].

Pentaspyrida Haeckel, 1882: 442 [invalid name, as a subfamily]; 1887: 1024, 1052 [as a subfamily].

Pleurospyrida Haeckel, 1882: 444 [nomen nudum, as a subfamily].

Stephida Haeckel, 1882: 444-445 [invalid name, as a family]. — Lankester 1885: 850 [as a family].

Dyostephanida Haeckel, 1882: 446 [nomen nudum, as a subfamily].

Dyostephida Haeckel, 1882: 446 [nomen nudum, as a subfamily].

Parastephida Haeckel, 1882: 446 [nomen nudum, as a subfamily].

Plectoida Haeckel, 1884: 30 [invalid name, as a family].

Cyrtoidea – Haecker 1908: 448 [invalid name, as a rank between suborder and family]. — Chediya 1959: 188 [as a superfamily]. — Pessagno 1977b: 933 [as a superfamily]. — Cachon & Cachon 1985: 294 [as a superfamily].

Pentaspyrinae – Clark & Campbell 1942: 57 [invalid name]; 1945: 32. — Campbell & Clark 1944a: 35. — Chediya 1959: 179.

Dicyrtoidea – Clark & Campbell 1942: 65 [invalid name, as a section above a family]; 1945: 36. — Campbell & Clark 1944a: 40 [as a section above a family]; 1944b: 23 [as a section].

Enneaplagiinae Campbell, 1954: D104 [nomen nudum].

Pentaspyridinae – Campbell 1954: D112 [invalid name].

Theophatninae Campbell, 1954: D134 [nomen nudum].

Cyrtoidae – Orlev 1959: 454 [invalid name, as a family].

Dicyrtoidae – Cachon & Cachon 1985: 294 [invalid name].

REMARKS

Families with no assigned species that can tentatively be placed in Nassellaria and are “ nomina nuda ” without any taxonomic information are simply listed herein. This list does not include higher ranks than the family-rank (e.g., suborder Cyrtida ).