Revision of the family Tetrentactiniidae Kozur et Mostler, 1979 (radiolarians of the Late Paleozoic). Part 1. Family, subfamilies

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Abstract

The family Tetrentactiniidae Kozur et Mostler, 1979, is revised and its diagnosis and generic composition are emended. The subfamily Tetrentactiniinae Kozur et Mostler, 1979, stat. nov., is revised, and its rank is emended. The new subfamily Uralitininae Afanasieva et Gainullina, subfam. nov. is established. A diagnosis of a new genus and new species Uralitina megalospina gen. et sp. nov. is given. The validus status is restored for the genus Triaenoentactinosphaera Wang, 1997.

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About the authors

M. S. Afanasieva

Borissiak Paleontological Institute, Russian Academy of Sciences

Author for correspondence.
Email: afanasieva@paleo.ru
Russian Federation, Moscow, 117647

E. A. Gainullina

Borissiak Paleontological Institute, Russian Academy of Sciences; Lomonosov Moscow State University

Email: elika10@bk.ru
Russian Federation, Moscow, 117647; Moscow, 119991

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2. Fig. 1. Family Tetrentactiniidae Kozur et Mostler, 1979, sensu Dumitrica, 2011: a – Tetrentactinia barysphaera Foreman, 1963, Upper Devonian, Famennian Stage, Huron Formation, Ohio, USA (Foreman, 1963, p. 282, pl. 7, fig. 9); b – Triaenosphaera sicarius Deflandre, 1973, Lower Carboniferous, Visean Stage, Montaigne Noire Mountains, France (Deflandre, 1973, p. 1150, pl. 2, fig. 3); c – Tetraspongoactinia holdsworthi Won, 1998, Lower Carboniferous, upper Tournaisian substage, Rhenish Slate Mountains, Germany (Won, 1998, p. 257, pl. 4, fig. 10); d – Multisphaera impersepta Nazarov et Afanasieva in Afanasieva, 2000, Lower Permian, Artinskian stage, Ural River, Donskoye village, Southern Urals, Russia (Afanasyeva, 2000, pl. 81, fig. 7); d – Tetratormentum narthecium Nazarov et Ormiston, 1985, Lower Permian, Artinskian stage, Ural River, Donskoye village, Southern Urals, Russia (Nazarov, Ormiston, 1985, p. 42, pl. 5, fig. 8); e – Tetragregnon sycamorensis Ormiston et Lane, 1976, Lower Carboniferous, Visean Stage, Sycmore Formation, Arbuckle Mountains, Oklahoma, USA (Ormiston, Lane, 1976, p. 167, pl. 2, fig. 6); f – Ellipsostigma australe Hinde, 1899, Middle Devonian, Givetian Stage, Yarrimi Formation, New South Wales, Australia (Hinde, 1899, p. 51, pl. 9, fig. 5); h – Staurentactinia nazarovi Schwartzapfel et Holdsworth, 1996, Upper Devonian, Upper Famennian, Woodford Formation, Criner Hills and Arbuckle Mountains, Oklahoma, USA (Schwartzapfel, Holdsworth, 1996, p. 202, pl. 9, fig. 8). Scale bar: a, g–f, h – 100 µm; b, c, g – 50 µm.

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3. Fig. 2. Possible variant of formation of the spicule and main needles: a – silicon-oxygen tetrahedron [SiO4], silicon atom is highlighted in black (from: Afanasieva et al., 2005, Fig. 3, a); b – hypothetical four-beam spicule (after: Popofsky, 1913, Fig. 1); c – empirical model of formation of the main needles (needles are shown with arrows, from: Afanasieva et al., 2005, Fig. 11, B).

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4. Fig. 3. Four-rayed spicule: a–g – Longibelona spiroacus (Afanasieva, 2023), preserved fragments of spicule rays (from: Afanasieva, 2023b, Fig. 6, a, b, d, f); d–g – Longibelona alia Afanasieva et Gainullina, gen. et sp. nov. (MS): d – fragment of spicule ray (from: Afanasieva, 2023b, pl. II, fig. 1), f, g – spicule ray with preserved bases of three more rays; h – Triaenosphaera fortunatovae Afanasieva, 2023, preserved two rays of the spicule, on one of which the bases of two more rays are visible (from: Afanasieva, 2023b, pl. II, fig. 3); i, k – Longibelona neglecta Afanasieva et Gainullina, gen. et sp. nov. (MS), fragment of a spicule ray. Designations: arrows indicate the preserved rays of the four-ray spicule (a, d, i, k) and the rays of the spicule with the preserved bases of three more rays of the four-ray spicule (g, f–h). Scale bar: g – 10 µm; a – 20 µm; g, j – 30 µm; c, d, f, h – 50 µm; b, i – 100 µm.

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5. Fig. 4. Microsphere in representatives of the subfamily Uralitininae, subfam. nov.: a, b – Triaenoentactinosphaera, microsphere and primary rod-shaped spines: a – T. sicarius (Deflandre, 1973 sensu Won, 1998) (restored from: Won, 1998, pl. 7, fig. 15); b – T. regularia Wang, 1997 (restored from: Wang, 1997, pl. 4, fig. 8); c–d – Kashiwara magna Sashida et Tonishi, 1985, microsphere and trilobed primary needles: c, d – porous outer and inner shells of the microsphere (restored from: Feng et al., 2007, pl. 6, figs. 17, 18); d – porous outer shell of the microsphere (restored from: Sashida, Tonishi, 1985, pl. 5, fig. 7); e, g – Kashiwara roadensis Nestell et Nestell, 2020, complex structure of the microsphere with a porous outer and mesh-like inner shell (adapted with modifications from: Nestell, Nestell, 2020, pl. 8, figs. 7a and 7b): e – arrow points to the three-lobed primary spine, g – fragment showing the double shell of the microsphere with fragments of the spicule rays (shown by small arrows), the large arrow points to the rod-shaped primary spine; h, i – Uralitina megalospina Afanasieva et Gainullina, gen. et sp. nov., specimen PIN, no. 0729: h – general view, i – fragment, arrows point to the primary rod-shaped spine and part of the spicule ray enclosed in the microsphere. Scale bar: a–c, d, f, h – 100 µm, g, f – 50 µm, i – 30 µm.

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6. Fig. 5. Examples of the structure of the skeletal shells of Tetrenta-ctiniidae Kozur et Mostler, 1979, rev. et emend., herein: a – one porous skeletal shell with spines, Longivelona neglecta Afanasieva et Gainullina, sp. nov, specimen PIN, No. 0209; b – two layers: porous → fine-mesh veil, Uralitina megalospina Afanasieva et Gai-nullina, gen. et sp. nov., the arrow shows the supporting spines of the porous skeletal shell; c – three layers: porous → mesh → fine-mesh veil, Longibelona globosa Afanasieva et Gainullina, gen. et sp. nov. (MS). Scale bar 30 µm.

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7. Fig. 6. Scheme of formation of the main needles: a – microsphere and primary spicule, b – primary needle, c – fragment of the outer sphere and large pores at the base of the main needle, d – main three-lobed needle.

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8. Fig. 7. The shape of the needles in representatives of the family Tetrentactiniidae Kozur et Mostler, 1979, rev. et emend., herein: a – needles covered with spongy skeletal tissue: Tetrentactinia barysphaera Foreman, 1963 (from: Afanaseva, 2000, pl. 84, fig. 5); b–d – three-lobed needles with subtriangular edges: b – Kashiwara magna Sashida et Tonishi, 1985 (restored from: Sashida, Tonishi, 1985, pl. 5, fig. 6), c – Uralitina megalospina Afanasieva et Gainullina, gen. et sp. nov., holotype PIN, no. 1108, g – Triaenosphaera fortunatovae Afanasieva, 2023 (from: Afanaseva, 2023b, pl. II, fig. 9); d–g – three-lobed needles with subrectangular needle edges, i.e. practically the same width of needles almost along their entire length: d – Longibelona neglecta Afanasieva et Gainullina, gen. et sp. nov. (MS) demonstrates spines on a porous sphere (from: Afanaseva, 2023b, pl. II, fig. 4), e – Longibelona globosa Afanasieva et Gainullina, gen. et sp. nov. (MS), the reticular layer and large pores at the base of the needle are clearly visible (shown by the arrow), g – Longibelona megacantha (Feng in Feng et al., 2007) (restored from: Feng et al., 2007, pl. 4, fig. 15); h–j – three-lobed twisted needles: h, i – Longibelona spiroacus (Afanasieva, 2023) (from: Afanasieva, 2023b, fig. 6, a, c), k – Tetraedrella tetraedros Afanasieva et Gainullina, gen. et sp. nov. (MS). Scale bar 100 µm.

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9. Fig. 8. Reconstruction of the distribution of genera of the family Tetrentactiniidae Kozur et Mostler, 1979, rev. et emend., herein. Genera and species: Tetrentactinia barysphaera Foreman, 1963, Russia, Timan-Pechora Basin (Afanasyeva, 2000, pl. 84, fig. 5); Triaenosphaera sicarius Deflandre, 1973, Germany, Northern Bavaria, Frankenwald (Kiessling, Tragelehn, 1994, pl. 5, fig. 21); Longibelona globosa Afanasieva et Gainullina, gen. et sp. nov. (MS); Tetraedrella tetraedros Afanasieva et Gainullina, gen. et sp. nov. (MS); Triaenoentactinosphaera regularia Wang, 1997 (Wang, 1997, pl. 4, fig. 8); Kashiwara magna Sashida et Tonishi, 1985 (Sashida, Tonishi, 1985, pl. 5, fig. 6); Uralitina megalospina gen. et sp. nov. Distribution of genera. Upper Devonian: Frasnian, Kazakhstan, Mugodzhary (Nazarov, 1973); Famennian: Russia, Kolymskoe Uplift (Nazarov et al., 1981), Timan-Pechora Basin, Polar Urals (Afanasyeva, 2000, 2020; Afanasieva, Amon, 2011; Afanasieva, 2020); Belarus, Pripyat Trough (Nazarov, 1988; Afanasieva, 2020; Afanasieva, 2020); USA, Ohio (Foreman, 1963; Maletz, 2011); Germany, Frankenwald (Kiessling, Tragelehn, 1994); northwestern China, Xinjiang Province (Wang, 1997). Lower Carboniferous: Tournaisian: Germany, Rhenish Slate Mountains (Braun, 1989; Won, 1998); France, Hautes-Pyrenees (Gourmelon, 1986), Montaigne-Noire Mountains (Gourmelon, 1987); USA, Utah, South Lakeside Mountains (Park, Won 2012); lower Tournaisian, Russia, Volga-Ural Basin, well. Melekesskaya-1 (Afanasieva, 2023b; Afanasieva et al., 2023); Visean: France, Montaigne-Noire Mountains (Deflandre, 1973); Germany, Rhenish Slate Mountains (Won, 1983); Northern Westphalia (Won, Seo, 2010). Lower Permian, Russia, Southern Urals: Asselian and Sakmarian stages, Sakmara River, Kondurovsky section (Afanasieva, 2021); Artinskian stage, Ural River, Donskoye village (this work). Middle Permian, Roadian and Capitanian stages, USA, Texas, Guadalupe Mountains (Maldonado, Noble, 2010; Nestell, Nestell, 2020). Upper Permian, Japan (Kuwahara, Yao, 1998, 2001; Hori, 2005; Sano et al., 2010); Philippines, central Busuanga Island (Tumanda, 1991, 1994; Tumanda et al., 1990); Upper Permian, Changsingian: central and southwestern Japan (Sashida, Tonishi, 1985, 1988; Feng et al., 2007); Eastern Thailand, Klaeng (Sashida et al., 2000); Southern China, Guangxi Province (Feng et al., 2007). Stage designations: fr – Frasnian, fm – Famennian, t – Tournaisian, v – Visean, s – Serpukhovian, b – Bashkir, ms – Moscow, ks – Kasimovian, gz – Gzhelian, as – Asselian, sk – Sakmarian, ar – Artinskian, kn – Kungurian, rod – Roadian, wor – Wordian, cap – Captainian, wch – Uchapingian, chn – Changhsingian. Scale bar 100 µm.

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