PTEROCORYTHIDAE 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 : 520-523

Family PTEROCORYTHIDAE Haeckel, 1882

Pterocorida Haeckel, 1882: 435 [below a tribe].

Sethocorida Haeckel, 1882: 430 [as a tribe]; 1887: 1192, 1289 [as a subfamily].

Calocyclida Haeckel, 1882: 434 [below tribe].

Podocyrtida Haeckel, 1882: 435 [below tribe]; 1887: 1313, 1314- 1315 [as a family]. — Bütschli 1889: 1990 [as a family]. — nec Rüst 1892: 183. — Nigrini 1967: 65 [as a family].

Sethocyrtida Haeckel, 1887: 1192, 1288-1289 [as a family]. — Bütschli 1889: 1989 [as a family]. — Rüst 1892: 182 [as a family]. — nec Cayeux 1894: 208.

Phormocyrtida Haeckel, 1887: 1313, 1365-1366 [as a family]. — Bütschli 1889: 1992 [as a family].— Nigrini 1967: 65-66 [as a family].

Theocyrtida Haeckel, 1887: 1313, 1395-1396 [as a family]. — Wisniowski 1889: 689 [as a family]. — Bütschli 1889: 1992 [as a family]. — Rüst 1892: 183 [as a family]. — nec Cayeux 1894: 209. — Nigrini 1967: 66 [as a family].

Phormocampida Haeckel, 1887: 1435 [as a family]; 1887: 1453- 1454 [as a family]. — Bütschli 1889: 1994 [as a family].

Phormocampiden – Haecker 1907: 126.

Phormocyrtiden – Haecker 1907: 126 [as a family].

Lamprocycladidae [sic] – Haecker 1908: 452-454 (= Lamprocyclidae).

Sethocyrtidae [sic] – Popofsky 1908: 287 (= Sethocyrtididae);1913: 372. — Schröder 1914: 100, 113. — Clark & Campbell 1942: 75; 1945: 40. — Campbell & Clark 1944a: 43; 1944b: 26. — Chediya 1959: 208. — Tan & Tchang 1976: 282. — Tan & Su 1982: 175; 2003: 113, 170. — Chen & Tan 1996: 153. — Chen et al. 2017: 202.

Podocyrtidae [sic] – Popofsky 1908: 288 (= Podocyrtididae); 1913: 373. — Schröder 1914: 119. — Clark & Campbell 1942: 80. — Campbell & Clark 1944a: 46; 1944b: 29 ( sensu emend. ). — Dogiel & Reshetnyak 1955: 47. — Chediya 1959: 213. — Tan & Tchang 1976: 283. — Tan & Su 1982: 176; 2003: 113, 174-175. — Nishimura 1990: 125-126 ( sensu emend. ). — van de Paverd 1995: 238. — Chen & Tan 1996: 153. — Tan & Chen 1999: 323. — Chen et al. 2017: 207.

Phormocyrtidae [sic] – Popofsky 1908: 289 (= Phormocyrtididae); 1913: 395. — Schröder 1914: 127. — Clark & Campbell 1942: 81; 1945: 43. — Campbell & Clark 1944a: 47; 1944b: 31. — Chediya 1959: 217. — Chen & Tan 1996: 153. — Tan & Chen 1999: 336. — Tan & Su 2003: 113, 192. — Chen et al. 2017: 212.

Phormocampidae – Poche 1913: 221. — nec Khabakov 1937: 110. — Campbell & Clark 1944b: 37. — Chediya 1959: 228. — Tan & Su 2003: 113, 214.

Sethocyrtididae – Poche 1913: 221.

Theocyrtidae [sic] – Popofsky 1913: 397 (= Theocyrtididae). — Schröder 1914: 129. — Clark & Campbell 1942: 89; 1945: 47. — Campbell & Clark 1944a: 49; 1944b: 32. — Chediya 1959:

220. — Tan & Tchang 1976: 289. — Tan & Su 1982: 177; 2003: 113, 200. — Chen & Tan 1996: 154. — Chen et al. 2017: 218.

Sethocorinae [sic] – Clark & Campbell 1942: 75 (= Sethocorythinae); 1945: 40. — Campbell & Clark 1944a: 43; 1944b: 27. — Chediya 1959: 208.

Sethocyrtinae [sic] – Orlev 1959: 455-456 (= Sethocyrtididae).

Theocyrtinae [sic] – Orlev 1959: 457 (= Theocyrtidinae).

Phormocampinae – Orlev 1959: 459.

Pterocoryidae [sic] – Riedel 1967b: 296 (= Pterocorythidae ) ( sensu emend. ); 1971: 657. — Riedel & Sanfilippo 1971: 1598. — Petrushevskaya & Kozlova 1972: 543. — Nakaseko et al. 1975: 174. — Nakaseko & Sugano 1976: 130. — Petrushevskaya 1981: 274-276. — Kozlova 1999: 144. — Amon 2000: 65-66. — Afanasieva et al. 2005: S300. — Afanasieva & Amon 2006: 149.

Pterocorydinae [sic] – Petrushevskaya 1971a: 230-231 (= Pterocorythinae); 1971b: 986.

Pterocorythidae – Moore 1972: 147 ( sensu emend. ). — Riedel & Sanfilippo 1977: 876. — Dumitrica 1979: 34. — Anderson 1983: 43. — Sanfilippo et al. 1985: 691. — Caulet & Nigrini 1988: 223. — Nigrini & Caulet 1988: 342. — Takahashi 1991: 123. — Hollis 1997: 65. — O’Connor 1997b: 108 ( sensu emend. ). — Boltovskoy 1998: 33. — Sugiyama 1998: 233. — Anderson et al. 2002: 1018. — De Wever et al. 2001: 258.

Calocyclinae [sic] – Petrushevskaya 1981: 226-227 (= Calocyclidinae). — Amon 2000: 64. — Afanasieva et al. 2005: S298. — Afanasieva & Amon 2006: 147.

Podocyrtiinae [sic] – Petrushevskaya 1981: 276 (= Podocyrtididae). — Afanasieva et al. 2005: S300. — Afanasieva & Amon 2006: 149.

Sethocorynae [sic] – Petrushevskaya 1981: 280 (= Sethocorythinae).

Pterocoryinae [sic] – Petrushevskaya 1981: 283 (= Pterocorythinae). — Afanasieva et al. 2005: S300. — Afanasieva & Amon 2006: 149.

Sethoconidae Nishimura, 1990: 124. — van de Paverd 1995: 229.

Lamprocyclidae – Tochilina 1997: 11-12.

Podocyrtinae [sic] – Amon 2000: 66 (= Podocyrtididae).

Sethocoryinae [sic] – Amon 2000: 67 (= Sethocorythinae). — Afanasieva et al. 2005: S300. — Afanasieva & Amon 2006: 149.

TYPE GENUS. — Pterocorys Haeckel, 1882: 435 [type species by subsequent designation ( Campbell 1954: D130): Pterocorys campanula Haeckel, 1887: 1316 ].

INCLUDED GENERA. — Albatrossidium Sanfilippo & Riedel, 1992: 16 . — Anthocyrtidium Haeckel, 1882: 430 (= Anthocyrtissa , Sethocyrtis synonymized by Caulet 1974: 239; Anthocyrtura synonymized by Petrushevskaya & Kozlova 1972: 545; Phormocampe synonymized by Petrushevskaya 1981: 282; Sethocanium n. syn.; Sethocorys synonymized by Caulet 1979: 132). — Calocyclas Ehrenberg, 1847: 54 (= Calocyclissa with the same type species; Anthocyrtonium synonymized by Petrushevskaya 1981: 280; Calocycletta n. syn., Calocyclior n. syn., Calocyclissima n. syn., Calocyclopsis n. syn.). — Calocycloma Haeckel, 1887: 1384 . — Lamprocyclas Haeckel, 1882: 434 (= Lamprocyclia with the same type species; Androcyclas synonymized by Petrushevskaya 1971a: 117; Craterocyclas, Hexalodus, Theocorbis synonymized by Petrushevskaya & Kozlova 1972: 544). — Lamprocyrtis Kling, 1973: 638 . — Lampterium Haeckel, 1882: 434 (= Alacorys , Tetralacorys ,? Lamptidium synonymized by Petrushevskaya & Kozlova 1972: 543). — Phormocyrtis Haeckel, 1887: 1368 . — Podocyrtis Ehrenberg, 1846: 385 (= Podocyrtidium with the same type species; Podocyrtoges n. syn., Podocyrtonium n. syn., Podocyrtopsis n. syn.). — Pterocorys Haeckel, 1882: 435 (= Sethoconus with the same type species, Conarachnium n. syn.; Lithopilium , Theoconus , Theocorax , synonymized by Petrushevskaya 1971a: 232). — Tetracorethra Haeckel, 1882: 429 (= Hexacorethra synonymized by Petrushevskaya 1971a: 234). — Theocorythium Haeckel, 1887: 1416 (= Theocapsilla synonymized by Petrushevskaya 1981: 286; Theocapsura synonymized by Petrushevskaya 1981: 287;? Theocorypha ,? Theocyrtis n. syn.). —? Anthocyrtoma Haeckel, 1887: 1268 . —? Calocyclura Haeckel, 1887: 1384 [errata 1764]. —? Theocorusca Haeckel, 1887: 1407 .

NOMINA DUBIA. — Cyrtocorys, Ennealacorys , Lamprocycloma , Lamptonium , Phrenocodon .

JUNIOR HOMONYM. — Cyrtocoris Haeckel, 1882 (= Cyrtocorys) nec White, 1842.

DIAGNOSIS. — Pterocorythidae consist of a small cephalis, a truncated, wide conical thorax, and a large abdomen. The aperture is always open and is associated with numerous feet, a circular rim on aperture, or a very coarse frame instead of abdomen. The cephalis varies from an elongated shape with a long apical horn to a thick-walled spherical shape with a robust apical horn. A pore, or pore-like depression, between the cephalis and thorax (sutural pore) is present ( Anthocyrtidium , Calocyclas, Theocorythium ) or absent ( Lamprocyclas , Lamprocyrtis ) as a stable character at the genus level. A sutural pore is always connected to the A-rod. In forms with an elongated cephalis, a free A-rod merges with the cephalic wall, constituting a rim, or part of a blade, along the apical horn. An alignment of several pores is visible on both sides of the A-rod on the cephalic surface. This alignment is found along the A-rod in Calocyclas and Lamprocyrtis . The elongated cephalis is divided into one larger unpaired lobe and two small- er lateral paired lobes separated by two directed arches AL that are obliquely oriented downward. The D-rod and double L-rod extend downward to merge with the thoracic wall but are never connected to the feet.

The cephalic initial spicular system consists of MB, A-rod, V-rod, double L-rod, and a double AL-arch. The A-rod is free near the basal ring and merges with the cephalic wall, becoming an apical horn. The V-rod is also free in the cephalic cavity and rarely penetrates the cephalic wall. Both the double l-rod and the distinguishing Ax-rod are present or absent. Three types of basal ring are recognized as an infra-species variation in most genera: (Type A) The basal ring is directly connected with the apical side end of MB, double L-rod, and V-rod to form four collar pores; (Type B) Differing from Type A, the basal ring is directly connected to the double l-rod, instead of the MB, forming four collar pores; and (Type C) a basal ring with six collar pores made of D-, V-, double l- and double L-rods. In the case of Type B, an additional very small double pore on the shell wall is formed by the D-rod, double l-rod and double Dl-arch. In the case of Type C, the double pores on the apical side of the basal ring are particularly very small.

A transparent endoplasm fills the cephalic cavity. Endoplasmic lobes of even sizes are present. Their size among specimens is variable but they never extend beyond the thorax. Algal symbionts are present and surround the endoplasmic lobes on the inner side of the shell test. Pseudopodia radiate throughout the test; a conical bundle of pseudopodia and a long robust thick pseudopodium (axial projection) extend outward from the aperture of the test. Tetracorethra has a degraded shell test. Living forms of Tetracorethra are also characterized by well-developed endoplasmic lobes and the absence of algal symbionts. The growth and development of pseudopodia remains still unknown in Tetracorethra .

STRATIGRAPHIC OCCURRENCE. — Late Paleocene-Living.

REMARKS

The type genus of this family is Pterocorys and the genitive form is Pterocorythos, thus the stem is Pterocoryth-, and the family name should be Pterocorythidae (seeMoore 1972: 147). Some species of Pterocorythidae with an absent lobe-like cephalis are occasionally misidentified as the Theocotylidae . This misidentification occurs regardless of the fundamental differences in the combination stability of the cephalic structure. The A-rod of the Pterocorythidae merges with the cephalic wall in most of the species, whereas the A-rod in Theocotylidae is free in the cephalic cavity and extends vertically to attach itself to the top of the cephalic cavity.The Pterocorythidae are generally distinguished by the presence of an elongated cephalis with an A-rod running along the cephalic wall (e.g., De Wever et al. 2001). However, this diagnosis cannot be rigidly applied for some genera such as Calocyclas , due to the variability of its cephalis which may be spherical ( Calocyclas sensu stricto) to elongated (the Calocycletta -form of Calocyclas ) or of an intermediate form ( Calocyclior , Calocyclissima and the Calocyclopsis -form of Calocyclas ). The genus Calocyclas ( sensu stricto) differs from other Pterocorythidae by the A-rod position, which is variable and free in the cephalic cavity ( Ogane et al. 2009b: pl. 94, fig. 7c). Other species assigned commonly to synonymies of Calocyclas ( Calocycletta , Calocyclior , Calocyclissima and Calocyclopsis -forms) have an A-rod that merges with the cephalic wall ( Moore 1972: pl. 1, fig. 1; pl. 2, fig. 5).

The cephalic structure has been described in Anthocyrtidium ( Caulet 1974: pl. 9, figs 1-2; Nishimura & Yamauchi 1984: pl. 37, fig. 3; Nishimura 1990: figs 31.6, 31.9;Sugiyama et al. 1992: pl. 26, fig. 2; O’Connor 1997b: pl. 5, figs 9-13), Calocyclas ( O’Connor 1997b: pl. 8, figs 2, 3;1999: pl. 3, figs 5-11; Sugiyama & Furutani 1992: pl. 19, fig. 7),? Calocycloma ( Nishimura 1990: figs 30.3-30.5), Calocyclura (Sugiyama et al. 1992: pl. 20, figs 5, 6), Lamprocyclas ( Nishimura & Yamauchi 1984: pl. 37, fig. 11; Nishimura 1990: figs 31.1, 31.2, 39.1; Sugiyama & Furutani 1992: pl. 18, fig. 5;Sugiyama et al. 1992: pl. 26, figs 5, 6; Tochilina 1997: pl. 1, fig. 2; pl. 2, figs 1-6; pl. 4, figs 2, 3; pl. 14, figs 14, 15; O’Connor 1999: pl. 4, figs 28-32), Lamprocyrtis ( Caulet 1971: pls 3, 4; 1974: pl. 10, figs 5, 6; Nishimura 1990: fig. 38.1; Sugiyama et al. 1992: pl. 27, fig. 4), Lampterium ( Nishimura 1990: figs 30.1, 30.2), Pterocorys ( Nishimura & Yamauchi 1984: pl. 38, fig. 6), and Theocorythium ( Nishimura & Yamauchi 1984: pl. 38, fig. 10; Sugiyama & Furutani 1992: pl. 19, fig. 1). The position of the A-rod, merged to the cephalic wall, is visible in the light microscopic image of Phormocyrtis ( Ogane et al. 2009b: pl. 58, fig. 3f) and Podocyrtis ( Ogane et al. 2009b: pl. 95, fig. 4b). The presence or absence of double l-rods varies at the species level, but not at the genus-, nor family-level. A good review of this difference was well illustrated inTochilina (1997) and according to her, Lamprocyclas has basal rings of Type A ( Tochilina 1997: pl. 4, fig. 5), Type B ( Tochilina 1997: pl. 4, fig. 2), and Type C ( Tochilina 1997: pl. 1, fig. 2; pl. 5, fig. 15).

The genus composition of the Pterocorythidae differs considerably among Petrushevskaya (1981: 275-276), De Wever et al. (2001: 258) and this catalogue. These divergences result from detailed investigation of the cephalic structure in 1990s. Additionally, our explanation is based on the first comprehensive integrated study of the cephalic structure anatomy and the molecular phylogeny. The overall appearance of Tetracorethra resembles the Plagiacanthoidea type, but the cephalic initial spicular system and protoplasmic structure of the former are identical to that of Pterocorys ( Petrushevskaya 1971a: 234; 1981: 291). This opinion was supported by the molecular phylogenetic data of Sandin et al. (2019).

The three questionably assigned genera ( Anthocyrtoma , Calocyclura and Theocorusca ) were not treated in De Wever et al. (2001). Petrushevskaya (1981) placed Anthocyrtoma and Calocyclura in subfamilies “Lapmpromitrinae” and “Theocotylinae” ( Petrushevskaya 1981: 104-105, 222), respectively.The taxonomic position of Theocorusca was not clearly mentioned in Petrushevskaya (1981: 316) but appears included in the Cannobotrydidae as the figure of this genus was placed with those of Botryocylinder and Rhopalosyringium . Herein, Anthocyrtoma is tentatively included in the Pterocorythidae . The complete form of Anthocyrtoma ( Riedel & Sanfilippo 1973: pl. 3, fig. 5) is observed with a very large appendage consisting of many feet appearing below a giant thorax. Partially incomplete specimens of Anthocyrtoma ( Riedel & Sanfilippo 1973: pl. 6, fig. 4) clearly display a free A-rod in the cephalic cavity and a probable V-rod along the left side of the cephalic cavity that resembles a dark line. This cephalic structure is more likely similar to that of the Theocotylidae than that of the Pterocorythidae . Other forms of Anthocyrtoma ( Riedel & Sanfilippo 1973: pl. 6, figs 2, 3) appear further as synonyms of the genera Clistophatna Haeckel 1882 and Clistophaena Haeckel 1887 ( Theocotylidae ). If this observation is correct, the valid genus name for Anthocyrtoma is Clistophatna , and thus becomes a member of Theocotylidae . The taxonomic position of Calocyclura may also be placed in Theocotylidae due to the six collar pores in the basal ring, which is not directly adjoined to the D-rod. In addition, Calocyclura has a free D-rod near the MB, a vertical extending free A-rod, and has not sutural pore (Sugiyama et al. 1992: pl. 20, figs 5, 6). The appropriate position of Theocorusca is unclear.

The ultrafine protoplasmic structure of Pterocorys was documented by Sugiyama & Anderson (1997b: pl. 1, figs 2, 3, 5; pls 4, 5) through transmitted scanning microscope (TEM) images. A normal optical image of living specimens was given for Anthocyrtidium ( Suzuki & Not 2015: fig. 8.11.17), Pterocorys ( Sashida & Kurihara 1999: fig. 11.15; Matsuoka 2007: figs 4b, 5b; 2017: figs 26, 27; Matsuoka et al. 2017: Appendix b), Tetracorethra ( Zhang et al. 2018: 15, fig. 24, p. 21, fig. 30) and Theocorythium (Matsuoka 2017: fig. 28).

The evolution of the Pterocorythidae at the family, genus and species levels was studied on the basis of a continuous stratigraphic distribution and a detailed geographic distribution. The evolution has been well documented at the family level ( Sanfilippo & Riedel 1992), illustrated at genus level for the Podocyrtis -Lampterium lineage ( Riedel & Sanfilippo 1981: fig. 12.7; Sanfilippo & Riedel 1990), and explained at species level for Anthocyrtidium ( Nigrini & Caulet 1988) and Calocyclas (originally Calocycletta in Moore 1972: text-fig. 1; Riedel & Sanfilippo 1981: fig. 12.11).

The morphologic changes between Podocyrtis sinuosa and Lampromitra mitra were examined by landmark, outline semi-landmark and landmark-constrained outline analysis ( Danelian & MacLeod 2019). Lamprocyclas may be infected with Marine Alveolata Group I ( Ikenoue et al. 2016).

VALIDITY OF GENERA

Calocyclas

Calocyclissa has the same type species as Calocyclas . The validity of Calocyclas was complicated by a circumvention of the ICZN rules about the type species of Cycladophora . Campbell (1954: D132) incorrectly designated Cycladophora stiligera as the type species of Cycladophora . Riedel & Sanfilippo (1970: 529) followed this designation and also synonymized Cycladophora stiligera with Calocyclas turris , which is the type species of Calocyclas ; thus, Calocyclas was considered a junior synonym of Cycladophora sensu Riedel & Sanfilippo (1970) . Once the type species of Cycladophora was corrected to Cycladophora davisiana by Lombari & Lazarus (1988), Calocyclas was no longer considered a synonym of Cycladophora .

Sanfilippo & Riedel (1992) established Calocyclior , Calocyclissima , and Calocyclopsis as subgenera of Calocycletta ; they are all monotypic subgenera with the following characters. Calocyclissima differs from the other subgenera in possessing longitudinal ribs between rows of thoracic pores, and very short cylindrical horns; Calocyclopsis is distinguished by its few broad shovel-shaped feet ( Sanfilippo & Riedel 1992: 30); Calocyclior is defined by a larger and more inflated thorax, more delicate abdomen with a longer porous part, and short termination with triangular teeth ( Sanfilippo & Riedel 1992: 31). However, there is no apparent need for the separation of these species into subgenera.

Petrushevskaya (1981: 279) revised the definition of Calocycletta , translated as follows: “ A helmet-shaped , typically pterocorythid cephalis basally narrowing. Cephalis height larger than its basal width. Collar area distinct. ‘Neck’ and lateral lobes well differentiated. Upper part of the shell cupolashaped; lower part cylindrical. External constriction between the thorax and abdomen almost unmarked. Pores similar in size and shape. Shell wall of the thorax thicker than on the abdomen. Except for ribs on the thorax, all thickened parts mamillated. Wall of the abdomen hyaline near the aperture, but may be composed of long, flat, ribbon-shaped teeth, surrounding the wide-open aperture. Porous part of the abdomen not longer than the thorax, but with teeth, the abdomen may be much longer than the thorax. ” This revised definition covers the characters of Calocyclas . Anthocyrtonium was previously synonymized with Calocyclas by Petrushevskaya (1981: 280). Among these groups, the oldest available name is Calocyclas .

Lampterium

Lampterium was designated as a subgenus of Podocyrtis ( Sanfilippo & Riedel 1992) , but it is convenient to regard it as a genus for disambiguation from true Podocyrtis . Tetralacorys has the same type species as Alacorys . Alacorys and Lamptidium were previously synonymized by Petrushevskaya & Kozlova (1972: 543); thus, Tetralacorys was automatically synonymized with Lampterium. Lampterium and Tetralacorys were simultaneously published in Haeckel (1882: 434 for the former and 436 for the latter). Lampterium has generally been preferred to Tetralacorys ; therefore, the former is selected as a valid name.

Podocyrtis

Podocyrtidium has the same type species as Podocyrtis . Podocyrtoges and Podocyrtopsis were established as subgenera of Podocyrtis by Sanfilippo & Riedel (1992). Podocyrtopsis is distinguished from Podocyrtoges by its larger thorax, lack of feet, and by abdominal pores being irregular in size and arrangement ( Sanfilippo & Riedel 1992: 14). Podocyrtonium differs from Podocyrtis by having a larger and wider abdomen ( Petrushevskaya 1981: 218). Podocyrtoges differs from Podocyrtis in having a larger abdomen than thorax. The thorax of Podocyrtopsis is larger than that of Podocyrtoges ( Sanfilippo & Riedel 1992: 14) . Sanfilippo & Riedel (1992) stressed that comparisons of single factors are much less satisfactory than considering entire subgeneric lineages, some of which contain diverse forms that cannot be briefly characterized. Thus, these groups should be considered a single genus, without subgenera; Podocyrtis is the oldest available name among them.

Pterocorys

Sethoconus and Conarachnium have the same type species, as do Theoconus and Theocorax . Theoconus was previously synonymized with Pterocorys ; thus, Theocorax is also automatically synonymized with Pterocorys . Eucyrtidium trochus was examined by Ehrenberg himself, and designated the type species of Conarachnium and Sethoconus , based on specimens in the Ehrenberg collection ( Suzuki et al. 2009c: pl. 55, figs 12a-c); the lectotype is a dorsal or ventral view of a young specimen of Pterocorys zancleus . In conclusion, these genera are synonymous. Pterocorys and Conarachnium were simultaneously published in Haeckel (1882: 430 for Conarachnium and 435 for Pterocorys ). As the type species of Pterocorys is a better specimen than that of Conarachnium , the former is selected as the valid name.

Theocorythium

Theocorypha has the same type species as Theocyrtis . The original specimen of Eucyrtidium barbadense which was examined by Ehrenberg himself was found in the Ehrenberg collection ( Ogane et al. 2009b, pl. 85, figs 7a-c) and was designated as the lectotype. This species is the type species of Theocyrtis . As shown in the supporting image for Theocyrtis in the Atlas , the real specimen is in very poor condition. Theocorythium is known from the late early Miocene, but the lectotype of Theocyrtis was dated to about the late Eocene. We tentatively synonymize Theocyrtis and Theocorypha with Theocorythium , but it should be considered nomen dubium. The genus name Theocyrtis has been used for important biostratigraphic marker species such as Theocyrtis tuberosa , but a new genus should be established for taxonomic stability.