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Karpinskiosaurus Ultimus (Seymouriamorpha, Parareptilia) from the Upper Permian of European Russia V

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Karpinskiosaurus Ultimus (Seymouriamorpha, Parareptilia) from the Upper Permian of European Russia V Paleontological Journal, Vol. 36, No. 1, 2002, pp. 72–79. Translated from Paleontologicheskii Zhurnal, No. 1, 2002, pp. 77–84. Original Russian Text Copyright © 2002 by Bulanov. English Translation Copyright © 2002 by åÄIä “Nauka /Interperiodica” (Russia). Karpinskiosaurus ultimus (Seymouriamorpha, Parareptilia) from the Upper Permian of European Russia V. V. Bulanov Paleontological Institute, Russian Academy of Sciences, Profsoyuznaya ul. 123, Moscow, 117997 Russia e-mail: [email protected] Received April 14, 1999 Abstract—The genus Raphanodon Ivachnenko, 1987 (Leptorophidae, Seymouriamorpha) is not a valid taxon, since its type species Nycteroleter ultimus Tchudinov et Vjuschkov, 1956 is assigned to the genus Karpinskio- saurus. The cranial anatomy of K. ultimus is described in the present paper with newly collected material from the Babintsevo locality. The genus Karpinskiosaurus is assigned to the subfamily Karpinskiosaurinae, which is affiliated to the Lower Permian Discosauriscinae (Discosauriscus and Ariekanerpeton) to form the family Karpinskiosauridae. INTRODUCTION the skull structure of this form will be published in the future. In 1956, Tchudinov and Vjuschkov described a new species of the genus Nycteroleter, N. ultimus based on The genus Karpinskiosaurus was established by jaw material from the Pron’kino locality (Orenburg Sushkin (1925) based on two specimens (an incomplete Region) (Tchudinov and Vjuschkov, 1956). Later, Iva- skeleton and a skull) described by Amalitsky (1921) as khnenko separated this form from the nycteroleters and Kotlassia secunda from the excavations at the Malaya placed it in the family Leptorophidae (Seymouriamor- Severnaya Dvina River (Sokolki locality). Sushkin pha, Parareptilia) under a new name, Raphanodon. This ranked this genus as a new family, Karpinskiosauridae, paper included reconstructions of the skull roof and based on its distinctions from Kotlassia proper (Kotlas- palatal complex of R. ultimus. However, these were sia prima) such as a different number of presacral and only based on the material from the Babintsevo locality, sacral vertebrae, the shape of the neural processes, and which yielded several incomplete skulls together with the relative depth of the otic notches. the jaws morphologically similar to those previously Later, Sushkin (1926, p. 339) mentioned another known from the Pron’kino locality (Ivakhnenko, 1987). species of Karpinskiosaurus, K. neglectus, which was established on the basis of a skull from the same local- Revision of the material demonstrated that a part of ity. This skull differs from that of K. secundus by specimens from Babintsevo referred to as R. ultimus slightly different proportions, the shape of postparietals and all of the jaw material from Pron’kino, including (dermosupraoccipitals), and by the pattern of the der- the holotype Nycteroleter (=Raphanodon) ultimus, mal ornament. Unfortunately, there were no figure belong to the genus Karpinskiosaurus. This is sup- image, complete description, and collection number of ported by the presence of such peculiar features of the the specimen in this paper; thus, this specimen is uni- genus Karpinskiosaurus as numerous jaw teeth, deep dentifiable. Currently, only one skull of Karpinskiosau- otic notches, the retention of premaxillary fontanels, rus, belonging to holotype K. secundus from the the pitted nature of dermal ornament, and a number of Sokolki locality, is housed at the Paleontological Insti- other characters. This assemblage of features strongly tute of the Russian Academy of Sciences (PIN). The distinguishes this form from the other Raphanodon specimen named K. neglectus is probably presented in species, R. tverdochlebovae, described on the basis of the paper by Watson (1954, text-figs. 31 and 32); how- cranial material from the Donguz-6 locality (Ivakh- ever, schematic sketches do not make possible the nenko, 1987). As far as Nycteroleter ultimus being a establishment of distinctions between the two species type species of the genus Raphanodon, the latter cannot of Karpinskiosaurus, including the characters men- be regarded as a valid taxon. The new generic name tioned by Sushkin. Thus, the validity of K. neglectus as Raphaniscus for the form from the Donguz-6 locality a separate species remains doubtful. was established earlier (Bulanov, 2000). Bystrow (1944) regarded Karpinskiosaurus speci- A significant part of the collection from Babintsevo mens as juveniles of Kotlassia prima and presented actually belongs to a seymouriamorph that is morpho- mixed description of both forms. Later, his reconstruc- logically close to Raphaniscus tverdochlebovae and tions combining the characters of Kotlassia and obviously belongs to the same genus. Information on Karpinskiosaurus were used by the majority of authors 72 KARPINSKIOSAURUS ULTIMUS (SEYMOURIAMORPHA, PARAREPTILIA) 73 who developed seymouriamorph systematics. Only Type species. Kotlassia secunda Amalitzky, Watson (1954), who owned one of Karpinskiosaurus 1921; Upper Permian, Upper Tatarian Substage, Vyat- skulls (K. neglectus?), insisted on the taxonomic auton- kian Regional Stage, Scutosaurus karpinskii Zone; omy of this genus (Watson, 1954). Russia, Arkhangelsk Region, Kotlas District, Sokolki The genus Karpinskiosaurus was only recently locality. introduced anew in the seymouriamorph systematics as Diagnosis. The genus forms monotypical sub- a valid taxon (Ivakhnenko, 1987; Ivakhnenko et al., family. 1997); however, it was placed in the same family as Species composition. The genus includes Kotlassia. The latter appeared to have much in common two species: K. ultimus (Tchudinov and Vjuschkov, with Raphaniscus in skull structure (the depth of the 1956) and K. secundus (Amalitzky, 1921); Upper Per- otic notches, the presence of well-developed postchoa- mian, Upper Tatarian Substage, North Dvinian and nal rows of large palatine teeth, the shape and pattern of Vyatkian Regional stages; European Russia, Orenburg, parasphenoid dentition, the form and size of the squa- Arkhangelsk and Vladimir regions. mosal, etc.). In this connection, it is appropriate to regard this genus as a leptorophid, placing it in a sepa- rate subfamily together with Raphaniscus. Karpinskiosaurus ultimus (Tchudinov et Vjuschkov, 1956) Nycteroleter ultimus: Tchudinov and Vjuschkov, 1956, p. 547; The description of Karpinskiosaurus ultimus using Kalandadze et al., 1968, p. 81. new material from the Babintsevo locality collected by Raphanodon ultimus: Ivakhnenko, 1987, p. 41 (partim); Ivakh- the author during the 1997–1998 field seasons is pre- nenko et al., 1997, p. 15 (partim). sented below. Holotype. PIN, no. 521/104, dentary; Orenburg Region, Sorochinsk District, Pron’kino locality; Upper SYSTEMATIC PALEONTOLOGY Permian, Upper Tatarian Substage, Vyatkian Regional Stage, Sarma Formation. Family Karpinskiosauridae Sushkin, 1925 Diagnosis. Intertemporal large and elongated. Subfamily Karpinskiosaurinae Sushkin, 1925 Ascending lamina of maxilla not developed. The mar- Karpinskiosauridae: Sushkin, 1925, p. 179; Ivakhnenko et al., ginal tooth apices only slightly curved lingually. 1997, p. 15. Description (Figs. 1–5). The skulls studied Type genus. Karpinskiosaurus Sushkin, 1925. range from 15 mm to 40 mm of length. In the speci- Diagnosis. Skull of adults more than 70 mm mens of K. ultimus, at the stage of the 30- to 40-mm- long. Canals of seismosensory system absent and long skull (specimens PIN, no. 4617/158; SGU, endocranium well ossified in adults. Dorsal processes nos. 104V/2008 and 104V/2009), the orbits are dis- of premaxillae high; fontanel between them retained. placed slightly anteriorly (Figs. 1a, 1b, 2a, and 2b). The Base of jaw teeth elongated and slightly folded. Postor- parietals are small; a tiny rounded pineal foramen is bital region of jugal enlarged. Cellular surface sculptur- located anterior to the ossification centers of these ing rapidly formed in ontogeny. bones, close to the suture with the frontals. The frontals Comparison. The subfamily Karpinskiosauri- are narrow; their length is approximately equal to the nae differs from the Discosauriscinae by its larger size, orbital length; the nasals are of the same width but the absence of the seismosensory grooves at the late slightly shorter. The prefrontal is longer than the post- developmental stages, well-ossified endocranium in frontal. The lachrymal is deep in the postnasal region; adults, longer dorsal processes of the premaxillae, the it expands far posteriorly below the orbit. The lachry- retention of a fontanel between them, the elongation monasal duct begins like a narrow funnel in the anterior and slightly folded pattern of the jaw tooth bases, the wall of the orbit and opens on the anteroventral edge of larger extension of the postorbital division of the jugals, the lachrymal. and a rapid formation of cellular ornament in the course A spoon-shaped septomaxilla is displaced inside the of ontogeny. nasal cavity; the septomaxillary opening is apparently closed. Genus Karpinskiosaurus Sushkin, 1925 The postorbital is triangular. The intertemporal is large and elongated, larger than the supratemporal. In Kotlassia: Amalitsky, 1921, p. 1 (partim); Efremov, 1940, p. 379 (partim); Bystrow, 1944, p. 380 (partim); Efremov and Vjuschkov, all available skulls, the postparietals and tabulars are 1955, p. 18 (partim); Konzhukova, 1964, p. 141 (partim); Tatarinov, lost; however, in specimens PIN, no. 4617/158 and 1972, p. 71 (partim); Tverdokhlebova and Ivakhnenko, 1994, p. 124; SGU, no. 104V/2009, the posterolateral
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