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Revision of the Late Permian Chroniosuchians (Amphibia, Anthracosauromorpha) from Eastern Europe V

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Revision of the Late Permian Chroniosuchians (Amphibia, Anthracosauromorpha) from Eastern Europe V Paleontological Journal, Vol. 32, No. 4, 1998, pp. 390–401. Translated from Paleontologicheskii Zhurnal, No. 4, 1998, pp. 68–77. Original Russian Text Copyright © 1998 by Golubev. English Translation Copyright © 1998 by åÄàä ç‡Û͇ /Interperiodica Publishing (Russia). Revision of the Late Permian Chroniosuchians (Amphibia, Anthracosauromorpha) from Eastern Europe V. K. Golubev Paleontological Institute, Russian Academy of Sciences, ul. Profsoyuznaya 123, Moscow, 117647 Russia Received March 31, 1997 Abstract—The family Chroniosuchidae is revised. The systematic importance of various morphological fea- tures is evaluated. The family systematic structure is presented and the generic and specific diagnoses are refor- mulated. A new genus Jarilinus and a new species Chroniosaurus levis are established. Evolutionary peculiar- ities of chroniosuchids are discussed. A zonal biostratigraphic chart of the Upper Tatarian deposits of eastern Europe based on tetrapod data is suggested. INTRODUCTION. All published chroniosuchian data were revised and new material was described by Ivachnenko and Tver- Chroniosuchians are one of the most characteristic dochlebova (1980). These authors regarded the chro- elements of the Late Tatarian tetrapod faunas of Euro- niosuchians as a suborder of the order Anthracosauro- pean Russia. This group of relict anthracosaurs morpha. Two families: the Chroniosuchidae and the replaced the archegosauroid labyrinthodonts that dom- Bystrowianidae were recognized within the suborder. inated the aquatic tetrapod communities of the Kaza- The family Chroniosuchidae includes the small-sized nian and Early Tatarian in this territory. Chroniosu- Chroniosuchus paradoxus and Chroniosaurus don- chians include two families: the Chroniosuchidae Vjus- gusensis, as well as the large Chroniosuchus uralensis chkov, 1957 and the Bystrowianidae Vjuschkov, 1957 Tverdochlebova originally described in that paper from (Golubev, 1998). The bystrowianids were widely dis- the locality Blumental-3, (Orenburg Region). The tributed mainly in the Triassic, only the earliest mem- name Jugosuchus is considered to be valid again. Apart bers of this family emerge at the end of the Permian from the type species J. licharevi (=Chroniosuchus (Shishkin and Novikov, 1992). The chroniosuchids, in vjuschkovi), J. mirabilis and two new species: J. boreus contrast, are known only from the Permian. Ivachnenko (Mikulino, Vologda Region) and J. hart- The first known chroniosuchids: Chroniosuchus manni Ivachnenko (Syomin Ovrag, Tatarstan), were paradoxus and C. mirabilis were described by Vjusch- referred to this genus. The genera Jugosuchus and kov (1957). A fragment of the vertebral column from Bystrowiana were united into the family Bystrowian- Zavrazhye (Arkhangelsk Region) specimen PIN, idae. Later Ivachnenko and Shishkin (Shishkin and no. 2353/6 was described as Chroniosuchus sp. in that Novikov, 1992) suggested, that Jugosuchus should paper.1 At the end of the 1930s Riabinin in a manuscript belong to the Chroniosuchidae. I did not accept this described another chroniosuchid Jugosuchus licharevi point of view (Golubev, 1998). The paper included a from fragments of the lower jaws from Savvatii description of a chroniosuchid Uralerpeton tverdochle- (Arkhangelsk Region). However, this description was bovae, distinguished from other chroniosuchids by published by Shishkin thirty years later (Riabinin and considerably narrowed scutes. Shishkin, 1962). Tverdochlebova (1967) supplied the Thus, the Chroniosuchidae currently include four name Chroniosuchus vjuschkovi to a chroniosuchian genera and eight species: Chroniosuchus (C. para- from Zavrazhye described by Vjuschkov. Tverdochle- doxus and C. uralensis), Jugosuchus (J. licharevi, bova (1968) pointed to a significant similarity of the J. mirabilis, J. boreus, and J. hartmanni), Chroniosaurus present form to the genus Chroniosuchus and condition- (C. dongusensis) and Uralerpeton (U. tverdochlebovae). ally regarded it as Chroniosuchus licharevi, since J. licharevi is represented by poorly diagnostic remains. The chroniosuchids are very uniform morphologi- She also suggested, that C. vjuschkovi may turn out to cally. The largest variability range is exemplified by the be a junior synonym of C. licharevi. Tverdochlebova dermal ornament of the skull and dorsal armor scutes. (1972) erected a new chroniosuchid genus and species Several scute types are distinguished in the chronio- Chroniosaurus dongusensis. suchids based upon sculpturing features that may be arranged in a single morphological row: pustular, pec- 2 1 PIN—Paleontological Institute, Russian Academy of Sciences, tinate, cristate and pitted. The pustular sculpturing SGU—Saratov State University, TsNIGR—Central Research Geological–Prospecting Museum (St. Petersburg). SGU collec- 2 The terms pectinate and cristate to denote the dermal ornamenta- tions used here are housed in PIN. tion types are introduced here for the first time. 390 REVISION OF THE LATE PERMIAN CHRONIOSUCHIANS 391 ‡ 1 cm b 1 cm c d 1 cm e 1 cm fg Fig. 1. Scute dermal sculpture types in the family Chroniosuchidae: (a) pustular, Chroniosaurus dongusensis, topotype PIN, no. 3585/119; (b) pectinate, Chroniosaurus levis sp. nov., holotype SGU, no. 104B/1102; (c) cristate, Jarilinus mirabilis gen. nov., topotype PIN, no. 523/20; (d) pitted, Chroniosuchus licharevi, specimen PIN, no. 2005/2578; (e) pitted, Chroniosuchus paradoxus, topotype PIN, no. 521/3; (f) and (g) pitted, Uralerpeton tverdochlebovae, paratype PIN, no. 1100/94 and holotype PIN, no. 1100/8 correspondingly. type formed of isolated tubercles evenly distributed of regular isometric (Figs. 1e–1f), or irregular shape over the surface is the initial form (Fig. 1a; Pl. 4, figs. 1 (Figs. 1d and 1g; Pl. 5, figs. 2–6). The distinct longitu- and 2). This term was suggested by Shishkin (1987). dinal cristae, characteristic of the pectinate and cristate The pectinate type is characterized by fusion of the types are less expressed in this case and sometimes are neighboring tubercle bases that results in the formation absent. Numerous intermediate ornamentation types of short low vermiform crests bearing several promi- are observable between the main ones. nent knobs (pectens). The pectens are disposed mainly A certain scute ornamentation type, independent of transversely. The other characteristic feature of this its size, is characteristic for the scutes from particular scute type is the presence of two–five distinctly exhib- localities. The observable individual variation range ited longitudinal swollen crests (cristae). The cristae does not exceed the extent of distinctions between the may occupy the area on the scute body along the medial main types considered above, and includes intermedi- line, along the scute body and borders of the wings (the ate sculpture forms from those typical for the given most frequently occurring position) and directly on the locality and the neighboring ones. For example, the wing surface (Fig. 1b; Pl. 4, figs. 3–5). The cristate type pustular sculpture is characteristic of the “Jugosuchus is distinguished by further fusion of tubercles resulting boreus” scutes from Poteryakha-2 locality (Vologda in the formation of several distinct transverse crests of Region). Nevertheless, some samples demonstrate the second order (Fig. 1c; Pl. 5, fig. 1) between the lon- sculpturing transitive towards the pectinate type. The gitudinal crests of the first order. Occurrence of numer- extreme deviation type is displayed in the specimen ous anastomoses between the transverse cristae results PIN, no. 3713/57. The tubercle bases of the dorsal surface in the formation of pitted sculpturing. The pits may be of the scute body fuse in this specimen forming short PALEONTOLOGICAL JOURNAL Vol. 32 No. 4 1998 392 GOLUBEV Upper Tatarian correlation chart from the terrestrial vertebrate fauna Provincial zones by tetrapods Faunistics Zonal assemblage Assem- Zone Subzone Subassemblage blage System Division Stage Substage Regional Stage Archosaurus, Archosau- Uralerpeton, Bystrowiana, rus Dvinosaurus egregius, D. purlensis, Vyazniki rossicus Whaitsiidae, Dicynodon, Karpinskiosauridae Chroniosuchus Scutosaurus, Inostrancevia, Vyatkian Scutosau- paradoxus Annatherapsidus, Chroniosuchus, rus karpin- Jarilinus, Sokolki skii Jarilinus Karpinskiosauridae, Dicynodon, mirabilis Dvinosaurus primus, Cynodontia Upper Upper Tatarian Permian Chroniosaurus Proelginia, Chroniosaurus, levis Oudenodon, Sauroctonus, Sokolki Proelginia Dvinosaurus primus, permiana Ilyinskoye Chroniosaurus Leptorophidae, Galeopidae dongusensis Deltavjatia, Tropidostoma, Deltavjatia Galeopidae, Ictidosuchidae, Severodvinian vjatkensis Nycteroleteridae, Leptorophidae, Kotel'nich Chroniosuchidae gen. et sp. nov. transverse vermiform tuberculated crests characteristic of Sokolki subassemblage structure (table). The chronio- the pectinate type scutes. Longitudinal crests are always suchid scute distribution pattern in the Tatarian section present. However, in this case they are essentially pectens of the Sukhona River is even more distinctly repre- and do not look like swollen ridges typical for pectinate sented (Fig. 2). The scutes of the pustular type are scutes. On the other hand, the “J. boreus” scutes from found in the Ustye Strelny locality (Poldarsa Forma- Mutovino (Vologda Region) belong to the pectinate type. tion, the upper part of the Strel’na unit). The pustular- However, there are also the forms varying towards the pustular (specimen SGU no. 104B/1109) or cristate pectinate
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