Earth-Science Reviews 69 (2005) 27–77 www.elsevier.com/locate/earscirev Upper Permian vertebrates and their sedimentological context in the South Urals, Russia Valentin P. Tverdokhlebova, Galina I. Tverdokhlebovaa, Alla V. Minikha, Mikhail V. Surkova, Michael J. Bentonb,* aGeological Institute of Saratov University, Ulitsa Moskovskaya, 161, Saratov 410075, Russia bDepartment of Earth Sciences, University of Bristol, Bristol, BS8 1RJ, UK Received 22 April 2004; accepted 7 July 2004 Abstract Fossil fishes and tetrapods (amphibians and reptiles) have been discovered at 81 localities in the Upper Permian of the Southern Urals area of European Russia. The first sites were found in the 1940s, and subsequent surveys 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. A continental succession, some 3 km thick, of mudstones, siltstones, and sandstones, deposited on mudflats and in small rivers flowing off the Ural Mountain chain, span the last two stages of the Permian (Kazanian, Tatarian). The succession is divided into seven successive units of Kazanian (Kalinovskaya, Osinovskaya, and Belebey svitas, in succession) and Tatarian age, which is further subdivided into the early Tatatian Urzhumian Gorizont (Bolshekinelskaya and Amanakskaya svitas, in succession), and the late Tatarian Severodvinian (Vyazovskaya and Malokinelskaya svitas, of equivalent age) and Vyatkian gorizonts (Kulchumovskaya and Kutulukskaya svitas, of equivalent age). This succession documents major climatic changes, with increasing aridity through the Late Permian. The climate changes are manifested in changing sedimentation and the spread of dryland plants, and peak aridity was achieved right at the Permo–Triassic (PTr) boundary, coincident with global warming. Uplift of the Urals and extinction of land plants led to stripping of soils and massive run-off from the mountains; these phenomena have been identified at the PTr boundary elsewhere (South Africa, Australia) and this may be a key part of the end-Permian mass extinction. The succession of Late Permian fish and tetrapod faunas in Russia documents their richness and diversity before the mass extinction. The terminal Permian Kulchomovskaya and Kutulukskaya svitas have yielded respectively some 6 and 13 species of fishes (sharks, bony fishes, lungfishes) and 11 and 14 species of tetrapods (aquatic amphibians, herbivorous and carnivorous reptiles of all sizes up to the hippo-sized pareiasaurs and sabre-toothed gorgonopsians). Immediately following * Corresponding author. Tel.: +44 117 9545400; fax: +44 117 9253385. E-mail address: [email protected] (M.J. Benton). 0012-8252/$ - see front matter D 2004 Elsevier B.V. All rights reserved. doi:10.1016/j.earscirev.2004.07.003 28 V.P. Tverdokhlebov et al. / Earth-Science Reviews 69 (2005) 27–77 the end-Permian environmental catastrophe, earliest Triassic faunas consisted only of a few fish taxa and small, aquatic tetrapods, in low-diversity, low-abundance assemblages. D 2004 Elsevier B.V. All rights reserved. Keywords: Permian; Triassic; Mass extinction; Tetrapod; Amphibian; Reptile; Russia; Urals; Stratigraphy 1. Introduction dominantly from lakes and small rivers and deltas, with some marine and brackish units in the early The end-Permian crisis was the largest mass Kazanian (Newell et al., 1999). extinction ever, with estimates of the loss of as many Bones of fossil reptiles were reported from the as 80–90% of all species (Benton, 2003). The impact Copper Sandstones of Perm’ Province and the of the crisis on land is less well understood than in Orenburg area in the late 18th century, and the first marine settings, partly because of dating and correla- descriptions were published in the 1830s and 1840s. tion problems, but partly also because there are few At first, these were thought to have been the bones of areas where extensive faunas and floras are known deceased miners, or at least of mammals. Studies by from both the Late Permian and Early Triassic. Russian, English, and German palaeontologists during Indeed, there are only two regions in the world where the 19th century revealed a fauna of dinocephalian tetrapod faunas may be tracked across the boundary: synapsids and temnospondyls that demonstrated links the Karoo Basin in South Africa, and the Russian with the new discoveries from the Karoo of South Permo–Triassic successions, especially in the zone Africa (Ochev and Surkov, 2000). Intensive collecting from the River Volga to the plain south of the Ural by Russian palaeontologists through the 20th century, Mountains. Fossil amphibians and reptiles have been revealed a succession of faunas of fishes, amphibians, known from the Russian successions for more than and reptiles throughout the Late Permian from S to N 200 years, and many summaries have been published, of the Ural Mountain chain. in Russian and in other languages. However, the last We focus here on the Southern Urals area. Late comprehensive overview (Efremov and V’yushkov, Permian tetrapods are known also from the Severnaya 1955) lacked a modern stratigraphic and sedimento- Dvina River, Inta, and the Pechora Basin in the far N, logical context, and new finds have immeasurably as well as from some locations around the towns of expanded knowledge of the faunas. This paper Kotlas, Kirov, Kazan’, and Perm in the central area of presents the Late Permian tetrapod faunas of the S European Russia. The Inta fauna, of possible Ufimian Urals and their context, and it is a companion to Tverdokhlebov et al. (2003), which covered the Early and Mid Triassic tetrapod faunas. Upper Permian deposits are common throughout the whole of the E of European Russia, from the Barents Sea coast in the N, to the Pre-Caspian Depression in the S. The richest sites for fossil vertebrates lie in the S Urals region, around the town of Orenburg, extending W towards Samara and E to the Autonomous Republic of Bashkortostan (Fig. 1). The best exposures are seen close to the major rivers, 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. The Fig. 1. Geographical setting of the South Urals region. Permian tetrapod-bearing localities are located in the grid area (shown in whole area lay some 308N of the Permian palaeo- more detail in Fig. 23). For major zones: I, the Cis-Ural Trough; II, equator, and extensive red-bed sediments accumu- the southeastern slope of the Volga–Ural Anteclise; III, the Peri- lated, comprising some 3 km of deposits pre- Caspian Depression; IV, the Ural Mountain fold system. V.P. Tverdokhlebov et al. / Earth-Science Reviews 69 (2005) 27–77 29 age, consists of eryopoid temnospondyls, a captorhi- (1996, 1998, 1999, 2000), Ivakhnenko et al. (1997), nid, and a bolosaurid. The Ezhovo (Ocher) fauna from and Bulanov (1999, 2000, 2002a,b). Ezhovov, near Perm, of late Kazanian and possibly The first attempt to synthesize the data on Permian earliest Tatarian age, comprises various temnospond- and Triassic terrestrial vertebrates was made by yls, estemmenosuchid dinocephalians, and biarmosu- Efremov and V’yushkov (1955). Information on the chians (Chudinov, 1983; Modesto and Rybczysnki, S Cis-Urals was supplemented by Garyainov and 2000). These faunas are not further discussed here. Ochev (1962), who were able to list only seven Upper The stratigraphic subdivision of the Russian Permian tetrapod localities. A later catalogue by continental Upper Permian has been difficult because Kalandadze et al. (1968) gave more localities. The of the general scarcity of fossils. As for the overlying most extensive overview was provided by Tverdokh- Lower and Middle Triassic succession (Tverdokhle- lebova (1976), who was able to include all the new bov et al., 2003), most stratigraphic schemes have sites discovered by the geological surveying teams relied on the sporadically abundant vertebrate fossils from the SGU. Her catalogue was reproduced by and the long-term and substantial faunal changes. The Ivakhnenko et al. (1997). earliest scheme for subdivision of the Upper Permian All the Upper Permian localities in the E of and Triassic of European Russia based on tetrapod European Russia that have yielded tetrapod and fish remains was presented by Efremov (1937, 1950) and remains are in Kazanian and Tatarian deposits Efremov and V’yushkov (1955). This tetrapod bio- (Tverdokhlebova et al., 1989). Here, we review stratigraphy was confirmed and refined through the current understanding of the stratigraphic divisions 1960s and 1970s, and more recently an attempt has in the S Cis-Urals (the Cis-Ural Marginal Trough) and been made to develop a biostratigraphy based on in the SE slope of the Volga–Ural Anteclise, and ostracod remains determined by Molostovskaya outline the geological structure and history of the (1993, 1997, 1999, 2000). region. A broad overview is given of the sedimentary In the 1950s, only a limited number of Permian and environmental history of the region, as well as of tetrapod localities were known, and only from the the evolution of vertebrate ecosystems. The key focus Northern sections; only occasional finds had been of the paper is the sites themselves, documenting for reported from the SE of the Russian Plate. the first time the geographic location, sedimentology, Extensive geological surveying in the 1960s and fossil finds, and age of each site. This documentation 1970s increased the number of tetrapods identified. is a crucial first step in attempting to use the extensive About 80 new localities for tetrapod and fish Russian data to document the effects of the end- remains were discovered in the S Cis-Urals and Permian mass extinction, and to compare them with on the SE slope of the Volga–Ural Anteclise, and the Kazanian and Tatarian successions in the Karoo these were studied by a group of geologists and Basin of S Africa.