Tetrapod localities from the Triassic of the SE of European Russia

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Abstract

Fossil tetrapods (amphibians and reptiles) have been discovered at 206 localities in the Lower and Middle Triassic of the southern Urals area of European Russia. The first sites were found in the 1940s, and subsequent surveys, from the 1960s to the present day, have revealed many more. Broad-scale stratigraphic schemes have been published, but full documentation of the rich tetrapod faunas has not been presented before.

The area of richest deposits covers some 900,000 km2 of territory between Samara on the River Volga in the NW, and Orenburg and Sakmara in the SW. Continental sedimentary deposits, consisting of mudstones, siltstones, sandstones, and conglomerates deposited by rivers flowing off the Ural Mountain chain, span much of the Lower and Middle Triassic (Induan, Olenekian, Anisian, Ladinian). The succession is divided into seven successive svitas, or assemblages: Kopanskaya (Induan), Staritskaya, Kzylsaiskaya, Gostevskaya, and Petropavlovskaya (all Olenekian), Donguz (Anisian), and Bukobay (Ladinian).

This succession, comprising up to 3.5 km of fluvial and lacustrine sediments, documents major climatic changes. At the beginning of the Early Triassic, arid-zone facies were widely developed, aeolian, piedmont and proluvium. These were replaced by fluvial facies, with some features indicating aridity. At the end of the Middle Triassic, deltaic and lacustrine-marsh formations were dominant, indicating more humid conditions.

The succession of Early to Mid Triassic tetrapod faunas documents the recovery of life after the end-Permian mass extinction. The earliest faunas consist only of small, aquatic tetrapods, in low-diversity, low-abundance assemblages. Climbing the succession through the Early Triassic, more terrestrially adapted tetrapods appear, and larger herbivorous and carnivorous reptiles come to dominate in the Mid Triassic as ecosystems were rebuilt.

Introduction

Triassic deposits are best developed in the south of European Russia, where they follow a long succession of Permian sediments (Newell et al., 1999). The Triassic is exposed along the valleys of the Ural, Sakmara, and Samara rivers and their tributaries, as well as along the left-bank (eastern) tributaries of the Volga, from the foreland fold belt of the Urals in the east (100 km to the east of Orenburg) to the Samara meridian in the west (Fig. 1).

This area is remarkable for the concentration of Permo-Triassic tetrapod localities. The terrestrial vertebrate faunas provide a basis for the division and correlation of continental red-bed formations from Eastern Europe. The history of attempts to refine the stratigraphic division of the continental Triassic in the SE of European Russia demonstrates that increases in the number of tetrapod localities has made stratigraphic division more reliable and more refined.

An early scheme for dividing the Permian–Triassic according to tetrapods was proposed by Efremov (1937) and Efremov and V'yushkov (1955). When the scheme was initiated, the number of localities was relatively small, only 13. Later research by Garyainov and Ochev (1962) revealed 32 more, resulting in changes to the biostratigraphic scheme. The scheme was further refined after Blom (1968) discovered 30 tetrapod localities in the Samara Region. In the course of detailed geological survey, a group of geologists from the Geology Research Institute, the University of Saratov, led by V.P. Tverdokhlebov, discovered over 130 localities and revised those of Blom. Tverdokhlebov, 1966, Tverdokhlebov, 1970, etc.) has combined the data on terrestrial vertebrates and the results of facies-cyclic analyses from numerous sections to lay the foundations of a Triassic regional scheme, which is presented here. During the course of this work, substantial new materials of tetrapods were acquired.

Tetrapod remains were identified and described, and the most significant locations excavated by V.G. Ochev (University of Saratov) and M.A. Shishkin (Paleontological Institute, Moscow). In addition, tetrapod faunas were described by L.P. Tatarinov, A.G. Sennikov, N.N. Kalandadze, I.V. Novikov (Paleontological Institute, Moscow); fish remains were studied by M.G. Minikh and A.V. Minikh (Geology Institute, the University of Saratov).

Ochev and Shishkin reviewed the tetrapod remains from the southeastern part of European Russia, and elaborated and revised Efremov's biostratigraphic scheme Shishkin and Ochev, 1967, Shishkin and Ochev, 1985, Ochev and Shishkin, 1985. They established the Triassic of the Southern Cis-Urals as a standard for global correlation of coeval continental deposits Ochev and Shishkin, 1989, Novikov, 1991b, Shishkin and Ochev, 1985. The history of fossil collecting in the Permo-Triassic of the Urals is described by Ochev and Surkov (2000), Triassic tetrapod biostratigraphy in Russia is reviewed by Shishkin et al. (2000), and the amphibian and reptile faunas by Sennikov (1995b). The amphibian and reptile groups are reviewed by various authors in Benton et al. (2000).

The information presented here has not been collected together in print, in any language. Limited information of this kind is available in Efremov and V'yushkov (1955) and Blom (1968), but these authors list only a quarter or so of the localities that are now known. Subsequent publications, in Russian, offer limited glimpses of the new data, but they are far from comprehensive, and nothing is available in English. The present paper is a comprehensive catalogue of all known vertebrate-bearing localities from the famous South Urals region, giving basic information on geographic location, sedimentology, fossil finds, and age. This is fundamental for an understanding of the sequence of recovery after the end-Permian mass extinction, and for comparison with the Karroo faunas from South Africa.

A note on names. We have attempted to transliterate all Russian place names and author names according to a standard Anglo-Russian scheme Kielan-Jaworowska, 1993, Benton, 2000. Here and there, older ‘Germanic’ transliterations of author names survive by convention, for example in the names of authors of taxa—compare Ochev (Anglo-Russian transliteration) and Otschev (Germanic transliteration).

Section snippets

Geological structure

The SE of European Russia is geologically heterogeneous. The eastern part of the region (to the east of the Orenburg meridian) belongs to the Cis-Ural Marginal Trough (South Cis-Urals), and the western part constitutes the southeastern slope of the Volga-Ural Anteclise, forming a part of the East European Platform (Fig. 1).

The Cis-Ural Trough runs north–south. It opens broadly into the Peri-Caspian Depression to the south of the Ural River valley. The Trough is as wide as 110 km there,

Stratigraphy

In European Russia, the Triassic is represented by all three divisions, Lower, Middle, and Upper (Fig. 2). Locations of tetrapod-remains provide a basis for biostratigraphic division of the Lower and Middle Triassic deposits; unfortunately none are found in the Upper Triassic, so no scheme for its division is presented.

List of tetrapod stratigraphic distributions

Most specimens of fossil amphibians and reptiles are located in the collections of the Paleontological Institute, Russian Academy of Sciences, Moscow (PIN); Saratov State University, Saratov (SGU), and the Centralny Nauchno-Issledovatelskii Geologo-Razvedochny Muzei, Saint Petersburg (CNIGR).

List of tetrapod localities

The localities are numbered 1–206, according to a long-standing catalogue held at the Geological Institute in Saratov. Their distribution is indicated in Fig. 17.

Kopanskaya Svita

(12) Astrakhanovka I; (25) Gryaznushka; (27) Blyumental; (39) Nikolskoe; (53) Perevolotzkoe; (59) Yaprintzevo; (60) Krasnaya Pavlovka; (61) Radovka; (63) Aleksandrovka; (64) Kozlovka; (87) Kamenny Yar III; (88) Kamenny Yar IV; (93) Fedorovka I; (94) Fedorovka II; (96) Krasnaya Yaruga I; (98) Rodnikoviy Dol; (104) Kamenny II; (105) Pavlovka–Antonovka; (111) Volchiy II; (112) Volchiy III; (113) Kazanka I; (125) Staraya Terlovka; (126) Buzulukskoe; (129) Michurin; (130) Zhuravlevka; (131)

Acknowledgements

We thank the Royal Society and the Russian Academy of Sciences for funding joint fieldwork between the Saratov Institute of Geology and the University of Bristol in 1995 and 1996, and NATO and the Royal Society for a postdoctoral award to MVS for a year in the Department of Earth Sciences, University of Bristol.

Valentin Petrovich Tverdokhlebov is head of the mapping teams and field expeditions branch of the Geological Institute at the University of Saratov, where he has worked since 1956. He graduated from the University of Saratov, and has specialized since in mapping the Permo-Triassic successions of the South Urals around the city of Orenburg. He completed his PhD in 1967 on the sedimentology and stratigraphy of the Permo-Triassic, focusing in particular on the arid formations. From 1970 to 1973,

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    Valentin Petrovich Tverdokhlebov is head of the mapping teams and field expeditions branch of the Geological Institute at the University of Saratov, where he has worked since 1956. He graduated from the University of Saratov, and has specialized since in mapping the Permo-Triassic successions of the South Urals around the city of Orenburg. He completed his PhD in 1967 on the sedimentology and stratigraphy of the Permo-Triassic, focusing in particular on the arid formations. From 1970 to 1973, he participated, as sedimentologist, in a number of expeditions to study the Early Cretaceous dinosaur beds of Mongolia. He was made Doctor of Science in 1996. He has published 100 scientific papers during his career.

    Galina Ivanovna Tverdokhlebova graduated from the Geology Department of the University of Saratov in 1953 and became a permanent researcher in the Geological Institute, where she worked on geological mapping of the Cis-Ural region. She has specialized in the Upper Permian tetrapods of the region, particularly the amphibians, on which she completed her doctoral thesis. She has since participated in numerous expeditions to parts of Russia (basins of rivers Mezen, Sukhona, Malaya Severnaya Dvina), as well as to Turkmenistan, Uzbekistan, and Tadzhikistan. She has published more than 40 scientific papers.

    Mikhail Viktorovich Surkov studied in the Geology Department of the University of Saratov, and presented his PhD thesis on ‘Triassic anomodonts from the eastern part of European Russia and their stratigraphic significance’ in 1999. He has worked as a lecturer and scientific researcher in that Department since 1995. In 1998 he received a Soros postgraduate fellowship award, in 2001 a NATO/ Royal Society Postdoctoral Fellowship to work in Bristol, and in 2002 awards from INTAS and from the Palaeontological Association to continue his work on the phylogeny and palaeobiology of Triassic dicynodonts. He has published 20 scientific papers.

    Michael Benton is Professor of Vertebrate Palaeontology, and Head of Department, at the Department of Earth Sciences in Bristol. He has long had interests in the Permo-Triassic transition and in the fossil reptiles of those times. He graduated with a PhD from the University of Newcastle-upon-Tyne in 1981, and has since worked at the Universities of Oxford, Belfast, and Bristol. He was coordinator of a major British–Russian collaborative project from 1993 to 1996, funded by the Royal Society and the Russian Academy of Sciences, which operated exchanges between the two countries and led to a number of joint expeditions and joint publications (e.g. Benton et al., 2000).

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