Cyrtospirifer Vjacheslavi, a New Cyrtospiriferid Species (Cyrtospiriferidae, Brachiopoda) from the Middle Frasnian of the East European Platform1 E

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ISSN 0031-0301, Paleontological Journal, 2013, Vol. 47, No. 6, pp. 581–587. © Pleiades Publishing, Ltd., 2013. Original Russian Text © E.V. Sokiran, 2013, published in Paleontologicheskii Zhurnal, 2013, No. 6, pp. 24–28.

Cyrtospirifer vjacheslavi, a New Cyrtospiriferid Species (Cyrtospiriferidae, Brachiopoda) from the Middle Frasnian of the East European Platform1 E. V. Sokiran All-Russia Research Institute of Oil and Geological Prospecting, Liteinyi pr. 39, St. Petersburg, 192104 Russia e-mail: [email protected] Received July 21, 2011; in final form, February 1, 2013

Abstract—A new spiriferid brachiopod species, Cyrtospirifer vjacheslavi, from the Middle Frasnian of the East European Platform is described. These spiriferide shells from the Middle Frasnian of central Euro- pean Russia and Middle Timan assigned traditionally to Cyrtospirifer disjunctus for more then a century are not conspecific to British species and are here reassigned to Cyrtospirifer vjacheslavi sp. nov.

Keywords: Brachiopoda, Spiriferida, new species, Devonian, Middle Frasnian, East European Platform, Russia DOI: 10.1134/S0031030113060099

1 Spiriferide brachiopods of the genus Cyrtospirifer Cyrtospirifer. No attempts were made, however, to are widespread and abundant in the Upper Devonian compare Russian shells directly with the actual types deposits in Europe, Middle East and North America. from Britain and they are completely missing in the However, a number of widely recognized species of latest publications (e.g., Nalivkin 1947; Ljaschenko this genus, mostly established in the 19th and early 1959). Identification of British and Russian shells were 20th century remain poorly understood or misunder- a consequence of their lack of full study, because of the stood because of insufficient knowledge of the type very poor state of preservation of British shells material, absence of documentation of their precise described and illustrated by J. Sowerby (Sedgwick and stratigraphic origin, and insufficient information on Murchison, 1840, pl. 53, fig. 8; pl. 54, figs. 12, 13) and the morphology available from the original publica- later by J. Phillips (1841, pl. 29, figs. 128f–128h, 129; tions (Sartenaer 1999; Ma and Day 1999). In particu- pl. 30, fig. 129), there was no reliable data on the exter- lar, such a brachiopods as Cyrtospirifer disjunctus nal and internal morphology of representatives by this (Sowerby, 1840) is widespread in the Frasnian deposits species. Therefore, the main features of S. disjunctus of the East European Platform and even used as the and later Cyrtospirifer disjunctus in understanding of index species of the Cyrtospirifer disjunctus—Stenom- E. Verneuil (in Murchison et al., 1845), F.N. Cherny- etoporhynchus pavlovi Biozone (Rzhonsnitskaya, shev (1887), S.V. Obruchev (1915), D.V. Nalivkin 2000). (1930, 1947) and A.I. Ljaschenko (1959) was sub- The type material of Spirifer disjunctus comes from quadrate shells and relatively low interarea with nearly the Devonian deposits of the outskirts of Barnstaple and parallel edges. However the typical samples from the Petherwin (Cornwall, United Kingdom), from which Petherwin Formation differ from the Russian species has been described by J. Sowerby (Sowerby in Sedgwick not only stratigraphically but also morphologically. and Murchison, 1840). E. Verneuil was the first who According to L.E. Popov and M. Basset (personal reported Spirifer disjunctus from the Late Devonian of communication) the originals of Spirifer disjunctus central European Russia (Verneuil in Murchison et al., from the J. Sowerby collection were lost. Currently, 1845). Shells described by E. Verneuil were sampled in the collection includes a single specimen identified by outskirts of Petino villages (Don river basin) and there is J. Sowerby; however, it is poorly preserved; its photo- no doubt that they came from the rocks subsequently graphs were kindly provided by L.E. Popov and assigned to the Upper Semiluki subhorizon. M. Bassett (Pl. 7, fig. 2), as well as photographs Spiri- fer disjunctus from the collection of Phillips (Pl. 7, In 1930 D.V. Nalivkin placed Spirifer disjunctus figs. 1, 3–6). described by Verneuil from central European Russia to There were at least three working quarry in South 1 The article was translated by the authors. Petherwin is the main source of spiriferide shells in the

581 582 SOKIRAN area where J. Sowerby and J. Phillips collected Spirifer Order Spiriferida Waagen, 1883 disjunctus. According to Steward (1981) a single lime- Suborder Spiriferidina Waagen, 1883 stone bed within the unit contains conodonts indicative S u p e r f a m i l y Cyrtospiriferoidea Termier of the Famennian upper Scaphignathus velifer to lower et Termier, 1949 Polygnathus styriacus zones. Unfortunately currently poor preservation of shells of Spirifer disjunctus poses Family Cyrtospiriferidae Termier et Termier, 1949 problems with the study of their internal structure and Subfamily Cyrtospiriferinae Termier et Termier, 1949 therefore their exact generic identification. Genus Cyrtospirifer Nalivkin in Frederiks, 1924 In Russia the name Cyrtospirifer disjunctus is attrib- Cyrtospirifer vjacheslavi Sokiran, sp. nov. uted to the spiriferide shells spread widely in the Plate 6, figs. 1–4 deposits corresponding to the Semiluki Horizon. In Spirifer disjunctus Sowerby: Murchison et al., 1845: p. 157, fact these shells have an excellent preservations and pl.4: fig. 4; Chernyschev, 1884: pl. 2, fig. 7; Obrouchev, 1916, are different in the morphology and age from British p. 46, pl. 2, figs. 1, 2. Spirifer disjunctus and they are regarded here as a sep- Spirifer Archiaci Murchison: Venyukov 1886, p. 66, pl. 3, fig. 2. arate new species Cyrtospirifer vjacheslavi sp. nov. Spirifer Verneuili Murchison: Venyukov 1886: p. 66, pl. 3, fig. 4. Shells of the new species considered in the present Cyrtospirifer disjunctus (Sowerby): Nalivkin 1947, p. 113, study come from two main sources: (1) the Semiluki pl. 27, fig. 4; Ljaschenko 1959, p. 176, pl. 52, figs. 1–3. Horizon of the Don River Basin in the Voronezh Etymology. In honor of late Professor Region (also known in Russian geological literature as Vjacheslav G. Khalymbadzha. the “Central Devonian Field”); and (2) deposits cor- H o l o t y p e: CNIGR 2332/2858, complete shell, responding to the Kraipol Formation of Middle central regions of East European platform, right bank Timan. In the Central Devonian Field Cyrtospirifer of the Don River, Semiluki village, Upper Devonian, vjacheslavi sp. nov. usually co-occurs with the brachi- Middle Frasnian, Upper Semilukian subhorizon. opods Stenometoporhynchus pavlovi (Nalivkin, 1930), D e s c r i p t i o n (Fig. 1). The shells are medium- Nervostrofia gassanovae (Ljaschenko, 1959), Hypothy- sized slightly ventribiconvex, subquadrate to trans- ridina semilukiana Nallivkin, 1930, Pseudoatrypa verse trapezoidal in outline with the maximum width uralica (Nalivkin, 1930), Cyrtina demarlii Bouchard, at the hinge line, cardinal extremities are acute and 1949, Anathyris helmersenii (Buch, 1840). According alate. The anterior commissure is uniplicate. The ven- conodonts this level corresponds to the Polygnathus tral valve is strongly and evenly convex in lateral profile efimovae Zone. with a beak projecting strongly beyond the hinge line In Middle Timan Cyrtospirifer vjacheslavi sp. nov. and curved dorsally. The lateral slopes are very slightly occurs in the Kraipol Formation from the natural out- convex in the lateral profile, becoming concave mar- crops exposed in the Tsylma, Myla and Pizhma ginally in strongly alate specimens. The ventral inter- Pechorskaya river basins. Conodont assemblage area is curved, strongly apsacline, with the outer mar- reported by Kuzmin (2001) suggesting the Middle to gins almost parallel to the hinge line, declining Upper Po. asymmetricus Zone. The deposits of the abruptly near cardinal extremities. The delthyrium is upper part of the Kraipol Formation are tentatively large, triangular, up to 5 mm high and varying from 3.1 assigned to the Ancyrognathus triangularis Zone to 6.7 mm in width in the largest specimens, covering (Kuzmin 2001). Associated brachiopod fauna from up to half its height by the secondary shell deposits the Kraipol Formation includes such taxa as Stenom- under the pedicle callist. The ventral sulcus is deep and etoporhynchus pavlovi (Nalivkin, 1930), ‘Atrypa’ narrow, originating at the umbo and terminated into a koloschka Nalivkin, 1941, ‘Atrypa’ svinordi (Venyukov, low, rounded tongue. The dorsal valve is moderately 1886) Spinatrypa (Spinatrypa) semilukiana Ljas- and evenly convex with a low and narrow median fold chenko, 1959, Cyrtospirifer rudkinensis Ljacshenko, rounded in cross section and originating at the umbo. 1959, C. mylaensis Sokiran, 2006, C. tenticulum (Ver- The lateral slopes are gently convex in transverse pro- neuil 1845), Cyrtina demarlii (Bouchard, 1949), Ilme- file. The dorsal interarea is low, nearly orthocline. nia perlevis (Nalivkin, 1930), I. altovae Nalivkin, Radial ornament is costate with flattened ribs on 1941, Anathyris svinordensis Nalivkin, 1941, Anathyris flanks, bifurcating mainly in the ventral sulcus and helmersenii (Buch, 1840). dorsal fold. The number of ribs varies from 11 to 16 in

Explanation of Plate 6 Figs. 1–4. Cyrtospirifer vjacheslavi sp. nov.; 1—CNIGR 2332/2858, holotype, complete shell, ventral valve (1a) (×1.5), dorsal valve (1b) (×1.5), posterior view (1c) (×1.5), lateral view (1d) (×1.5) of conjoined valves., East European Platform, Voronezh region, village of Semiluki, right bank of the Don River, Upper Devonian, Middle Frasnian, Upper Semilukian subhorizon. 2— CNIGR 2332/2858, complete shell, 2a—ventral valve (×1.5), dorsal valve (2b) (×1.5), posterior view (2c) (×1.5) of conjoined valves., Middle Timan, left bank of the Myla River, Upper Devonian, Middle Frasnian, Kraipol Formation. 3—CNIGR 1/13124, radial ornament in ventral sulcus showing pustules (×5), 4—CNIGR 2/13124, radial ornament and superimposed concentric fila (×5), all Middle Timan, left bank of the Myla River, Upper Devonian, Middle Frasnian, Kraipol Formation.

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Plate 6

1a 1d 2a

1b 2e 2b

1c 2c

3 4

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(a) (b) (c) (d) (e) (f) (g)

(h) (i) (j)

(k) (l) (m)

(n) (o) (p)

(q) (r) (s) 5 mm

(t) (u)

Fig. 1. Cyrtospirifer vjacheslavi sp. nov.: a–x CNIGR 4/13124—transverse serial sections of the shell (×2), East European Platform, Voronezh region, Semiluki village, right bank of the Don River, Upper Devonian, Middle Frasnian, Upper Semilukian subhorizon. the dorsal median fold and ventral sulcus, and from 17 preserved. The surface of the ventral interarea is also to 27 on flanks of both valves. Most of the costae bifur- covered with fine vertical capillae usually obscured by cate from the median rib and from a pair of ribs flank- concentric ornament of strong, ridgelike, evenly ing the fold and sulcus. Superimposed radial capillae spaced fila up to 27 per 5 mm. Fine pustules are are very fine, about 50 per 5 mm, and usually poorly

Explanation of Plate 7 Figs. 1–6. Spirifer disjunctus (Sowerby); 1—NHM BC 58481, latex cast of ventral internal mould (×1.5), 2—NHM Bb 350, internal mould of conjoined valves (×1.5), 2a—ventral and 2b—dorsal views, 3—NHM BC 58482: 3a—latex cast of ventral internal mould (×1.5), 3b—posterior view of ventral internal mould (×1.5), 4—NHM BC 58485, posterior view (×1.5), 5—NHM B43939/a, latex cast of ventral internal mould (×1.5), 6—NHM B43939/b; 6a—latex cast of ventral valve (×1.5), 6b—radial ornament in ventral valve (×4), United Kingdom, Cornwall, South Petherwin, Landlake quarry, Upper Devonian, Middle-Upper Famennian, Petherwin Formation.

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Plate 7

12a

4 6а

56b

PALEONTOLOGICAL JOURNAL Vol. 47 No. 6 2013 586 SOKIRAN present, but poorly preserved in the most of observed pl. 5, figs. 1–9); however, this species differs from shells. C. vjacheslavi in the well-developed dorsal median The ventral interior has a prominent pedicle callist, fold and the winged shell shape. Perhaps, Cyrtospirifer strong teeth and long, diverging, extrasinal dental lonsdalii (Murchison, 1840, pl. 2, figs. 1–2; Grabau, plates with adminicules. The dorsal interior has a wide 1931, p. 213, pl. 27, figs. 1–6) from the Frasnian of ctenophoridium clearly subdivided into two lateral Ferques is synonym Cyrtospirifer grabaui Paeckelmann parts, dorsolaterally directed crura, and a fine median (Paeckelmann, 1942, p. 111, pl. 4, figs. 1–3; Vander- ridge bisecting the adductor muscle field. The spiralia cammen, 1959, pl. 5, figs. 1–9). have up to 12 whorls. Remarks. Specimens of Spirifer disjunctus Dimensions in mm and ratios: (Pl. 7, figs. 6a, 6b) show numerous rib bifurcations on flanks of both valves, whereas in Specimen no. LWL/WTHrvv Cyrtospirifer radial Holotype 2332 29.20 37.30 0.78 16.6 4.0 ribs are simple on flanks and bifurcating only in the 2604 27.60 37.60 0.73 20.60 4.70 ventral sulcus and dorsal fold. However, imperfect 2603 30.90 37.90 0.81 23.50 2.0 preservation of shells in the type locality makes very 2843 30.10 36.10 0.83 26.80 1.90 difficult to prove it without a general revision of the 2611 29.70 36.90 0.80 28.40 3.00 taxa assigned to the subfamily Cyrtospiriferinae, 2585 29.50 40.60 0.73 23.40 3.90 which is far beyond the scope of the present paper. 2614 25.20 34.80 0.72 20.70 3.10 O c c u r r e n c e. Upper Devonian, Middle Fras- 2855 27.90 33.30 0.84 32.30 2.50 nian, Upper Semiluki Subhorizon of the central 2614 26.70 35.0 0.76 20.50 2.90 regions of European Russia; Kraipol Formation of the 2848 25.60 34.80 0.74 18.70 Middle Timan. M a t e r i a l. Ten complete shells from Comparison and remarks. Shells of D.V. Nalivkin’s collection (CNIGR no. 2858) and Spirifer disjunctus deposited in the Natural History 109 specimens from M.M. Tolstichina collections Museum, London revealled that they differ signifi- (CNIGR no. 13020) all specimens come from the cantly from Cyrtospirifer vjacheslavi sp. nov. in spite of Upper Semiluki Subhorizon (villages of Petino, Semi- sometimes significant distortion of the British shells. luki, Devitsa, and Ust’e). 118 complete shells and Spirifer disjunctus has a broader triangular interarea 49 disarticulated valves from F.N. Chernyshev collec- and ribs bifurcations on flanks of both valves. The new tions (CNIGR no. 1226) from the Kraipol Formation species is morphologically similar to C. mylaensis of the Tsylma, Myla, and Pizhma Pecherskaya rivers in Sokiran 2006, however, the latter is characterized by a Middle Timan. more developed wide trapezoidal shell, a relatively Abbreviations. L—length of shell, W—width of high triangular ventral interarea, fewer ribs in the sul- shell. L/W—the ratio of length to width of shell, T— cus and fold, which is extremely poorly developed, in thickness of shell, Hrvv—height of interarea of pedi- the middle of which there is a slight depression. Lower cle valve. Semilukian Cyrtospirifer rudkinensis Ljaschenko, 1959 differs in its larger shell, triangular ventral interarea, The brachiopods described in this paper are housed higher and coarser ribs. Some representatives of at the CNIGR Museum in St. Petersburg. Cyrtospirifer schelonicus Nalivkin, 1941 from Lower Semilukian Subhorizon, which is rather variable, is ACKNOWLEDGMENTS similar to Cyrtospirifer vjacheslavi sp. nov. However C. schelonicus has a wide trapezoidal shell with a high I am grateful to Dr. M. Basset and Dr. L.E. Popov triangular interarea and well-developed dorsal median (Department of Geology, National Museum of Wales, fold. Cyrtospirifer vjacheslavi also has a distinct affinity Cardiff, United Kingdom) for assistance in the study to Early Frasnian Cyrtospirifer malaisi (Gosselet, of British material and valuable advice in writing the 1894) from the Dinant Basin, which Sartenaer (1982) article. National Museum of Wales provided fundidng considered as the type species of the genus Subquadri- for E.V. Sokiran during her visit to Cardiff in 2000. The angulispirifer, however, Carter et al. (1994) regarded study was supported by Silesian University at Sosnow- this genus as a junior synonym of Cyrtospirifer. Our iec (Poland), the Polish State Committee for Scien- shells differ from Cyrtospirifer malaisi in having a tific Research (KBN grant 3 P04D 040 22, headed by deeper sulcus and rounded lateral shell margins. Peetz Dr. G. Racki). (1901) proposed that Cyrtospirifer lonsdalii (Murchi- son 1840) is a synonym of Spirifer disjunctus (=Cyrtospirifer vjacheslavi sp. nov.), because it also has REFERENCES a narrow interarea with outer edges subparallel to the Carter, J.L., Johnson, J.G., Gourvennec, R., and hinge line. In fact, a narrow interarea with parallel Hou, H.F., A revised classification of the spiriferid brachio- edges is observed in other species of the genus, such as pods, Ann. Carnegie Mus., 1994, vol. 63, pp. 327–374. Cyrtospirifer grabaui Paeckelmann (Paeckelmann, Chernyshev, F.N., Materials to the study of the Devonian of 1942, p. 111, pl. 4, figs. 1–3; Vandercammen, 1959, Russia, Tr. Geol. Kom., 1884, vol. 1, no. 3, pp. 1–81.

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