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The First Find of Permian Ceratodontids (Dipnoi, Osteichthyes) in Russia O

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The First Find of Permian Ceratodontids (Dipnoi, Osteichthyes) in Russia O ISSN 00310301, Paleontological Journal, 2015, Vol. 49, No. 10, pp. 1112–1124. © Pleiades Publishing, Ltd., 2015. The First Find of Permian Ceratodontids (Dipnoi, Osteichthyes) in Russia O. A. Lebedeva, A. G. Sennikova,b, V. K. Golubeva,b, N. I. Krupinac, G. Niedzwiedzkid, and T. Suleje aBorissiak Paleontological Institute, Russian Academy of Sciences, Profsoyuznaya ul. 123, Moscow, 117997 Russia email: [email protected], [email protected], [email protected] bKazan Federal University, Kremlevskaya ul. 18, Kazan, 420008 Russia cEarth Science Museum, Moscow State University, Moscow, 119991 Russia email: [email protected] dDepartment of Organismal Biology, Evolutionary Biology Center, Uppsala University, Norbyvägen 18A, Uppsala, 75236 Sweden email: [email protected] eInstitute of Paleobiology, Polish Academy of Sciences, Twarda 51/55, Warsaw, 00818 Poland email: [email protected] Received January 30, 2015 Abstract—A huge dipnoan, Permoceratodus gentilis Krupina, gen. et sp. nov. (order Ceratodontiformes), from the terminal Permian beds (Zhukovian Regional Stage, Vyatkian Stage, Upper Permian) of the Sokovka locality (Vladimir Region) is described. It is characterized by the evolutionarily advanced high extent of fusion of skull roof bones and conservative well defined structures of the seismosensory system of the head. This combination distinguishes the new taxon from other Ceratodontiformes. A set of conservative and advanced characters is observed in many vertebrates of the Vyazniki faunal assemblage. Large tetrapods and fishes characterize the terminal developmental stage of the Permian fauna of Eastern Europe, which was fol lowed by impoverishment of the taxonomic composition accompanied by a decrease in body size. The dip noan described here, like some other vertebrates of this assemblage, belong to highrank taxa, which had just appeared in the Paleozoic, but reached flourishing in the Mesozoic. Keywords: Ceratodontiformes, lungfish, paleosynecology, ecological crisis, Upper Permian, Upper Vyatkian, Russia, Vladimir Region DOI: 10.1134/S0031030115110052 INTRODUCTION HISTORICAL ACCOUNT Among Late Permian vertebrate localities of the The Vyazniki vertebrate fauna was discovered dur East European Platform, a group of Vyazniki localities ing B.P. Vjuschkov’s (Paleontological Institute, Acad (Fig. 1) occupies an important place. It is situated on emy of Sciences of the USSR, Moscow: PIN) excava the right bank of the Klyazma River in the town of tions in 1951, 1955, and 1956. Vjuschkov dated it by Vyazniki (Vladimir Region) (Efremov and Vjuschkov, the Severodvinian Age based on relatively small rep 1955; Sennikov and Golubev, 2006b; Newell et al., tiles, which he regarded as insufficiently derived com 2010). In this area, the Vyazniki faunal and floral pared to the forms of the terminal Permian Sokolki assemblages of the terminal Permian, which is only fauna including gorgonopians and pareiasaurs and known in European Russia, is particularly representa also based on a conchostracan assemblage identified tive. This assemblage displays the last antecrisis devel by N.I. Novozhilov. Subsequent detailed studies of opmental stage of the continental biota at the very end vertebrates excavated by Vjuschkov undertaken by of the Permian. Therefore, a thorough examination of PIN researchers (Ivakhnenko, 1990; Shishkin, 1990; this assemblage is of great importance for gaining an Sennikov, 1995, 1996; Ivakhnenko et al., 1997; Gol understanding of the causes and processes of the ubev, 1998, 2000a, 2000b; etc.) has demonstrated that Permo–Triassic ecological crisis and mass extinction the Vyazniki assemblage should be dated as terminal on land. Permian, being the youngest in Eastern Europe, 1112 THE FIRST FIND OF PERMIAN CERATODONTIDS (DIPNOI, OSTEICHTHYES) 1113 30° 40° 50° ° 60 Bykovka 7 6 3 4 5 9 8 2 1 Moscow Vyazniki Klyazma River M7 Vyazniki 12 50° 10 1 km 11 13 Fig. 1. Geographical position of the Vyazniki group of localities: (1) Bykovsky Ovrag, (2) Zelenaya Gorka, (3) Bykovskoe Ozero, (4) Bykovka (Vyazniki2), (5) Sokovka (Vyazniki2), (6) Metallist, (7) Petrino, (8) Vyazniki1, (9) Yartsevo, (10) Balymotikha1, (11) Balymotikha2, (12) Tolmachevo, (13) Fedurniki. younger than the Sokolki (pareiasaurian–gorgono and isolated points in European Russia (Sennikov and pian) assemblage (Fig. 2). Golubev, 2005, 2006a, 2006b, 2007, 2013; Sennikov et al., 2014). Similar boundary assemblages have not In 1999, Sennikov renewed field studies in the yet been revealed anywhere in the world. vicinity of Vyazniki and Gorokhovets. He found the previously known bone beds in the Vyazniki locality Joint studies and excavations of the group of (Bykovka, Yartsevo) and discovered a new site, Gor Vyazniki localities were performed by the Borissiak okhovets, which has yielded the taxonomically richest Paleontological Institute of the Russian Academy of Permian vertebrate fauna of the Sokolki Assemblage Sciences, Moscow (PIN); Institute of Paleobiology of (Sennikov et al., 2003). This initiated a thorough and the Polish Academy of Sciences, Warsaw (IPP); and complex study of PermoTriassic faunal and floral the Faculty of Biology of Warsaw University in 2008, localities and reference sections in this region. Since 2010, and 2013, headed by A.G. Sennikov (PIN) and then, experts not only from PIN, but also from the T. Sulej (IPP). In September, 2013, the Sokovka local Geological Institute of the Russian Academy of Sci ity was intensely excavated. This locality is situated in ences, Moscow (GIN), Saratov and Kazan state uni a short and relatively deep gully cutting the right slope versities, and paleontologists and geologists from of the Klyazma River valley in the northwestern sub Poland, Germany, and Great Britain participated in urbs of the town of Vyazniki, just east of a sand quarry this research. near the village of Bykovka. The main excavation was performed in the right slope of the gully, where the fos Since 2003, new Late Permian bonebearing local sils became exposed over an area of about 20 m2 ities with fish and tetrapod fossils were discovered in (Fig. 3) and rich material of Late Permian vertebrates the vicinity of Vyazniki: Sokovka, Metallist, Balymo was collected. In the left slope, at the same hypsomet tikha, etc. (Fig. 1); in addition, the first Early Triassic ric level, excavation was performed within a small area (Vokhmian Regional Stage) vertebrate locality in and yielded only individual isolated vertebrate bones, Fedurniki (Sennikov and Golubev, 2013) was discov including remains of a huge dipnoan found by ered. Insects, ostracods, and macroflora were found G. Niedzwiedzki. This specimen is described below. for the first time. New material of bivalves and con chostracans was obtained; new fish taxa were identi To date, only individual isolated teeth of the dip fied and described. A palynological assemblage similar noan Gnathorhiza have been recorded in the Upper to that of the type section of the Permian–Triassic Permian of European Russia (Minikh, 1977; boundary beds of China is recognized; it displays the M. Minikh and A. Minikh, 1998; A. Minikh and socalled “algal” or “fungal” episode. In general, the M. Minikh, 2009). In the Induan Stage and Lower data on various biotic groups confirmed the suggestion Olenekian Substage of the Lower Triassic, remains of that the Vyazniki assemblage is a separate terminal Gnathorhiza become rather abundant and are used as Permian faunal and floral assemblage (Fig. 2) corre an index form (Minikh, 1977; M. Minikh and sponding to a stage of crisis destabilization of the Per A. Minikh, 1998). The first ceratodontid teeth of the mian biota, showing somewhat transitional state, and genus Ceratodus appear in Russia in the Upper Ole preceding complete extinction. This stage of destabili nekian Substage of the Lower Triassic and extend fur zation was probably rather short and fixed in Vyazniki ther up to the terminal Ladinian Stage of the Middle PALEONTOLOGICAL JOURNAL Vol. 49 No. 10 2015 1114 LEBEDEV et al. ISC RSC Regional Tetrapod zone Faunal assemblage stage Stage Stage Series Series System System Substage Bukobayian Mastodonsaurus assemblage Ladian Ladian Middle Middle Donguzian Eryosuchus assemblage sian sian Ani Ani Kannemeyeroid superassemblage Trematosaurus subassemblage Gamian Inflectosaurus assemblage Yarengian subassemblage Vyborosaurus– Angusaurus Triassic Triassic lian vkian subassemblage UstmyFedoro Olenekian Olenekian BenthosuchusAngusaurus assemblage kian subassemblage Lower Lower Slud Wetlugasaurus Parotosuchus Proterosuchian superassemblage WetlugasaurusThoosuchus subassemblage Vetlugian Benthosuchus gusevae Rybinskian assemblage subassemblage Benthosuchus Tupilakosaurus wetlugensis Spasskoe assemblage mian Vokh Induan Induan Zhukovian Archosaurus rossicus Vyazniki assemblage Nefyo Chroniosuchus Changh dovian Scutosaurus paradoxus karpinskii Sokolki subassemblage Jarilinus Bykovian mirabilis Chroniosaurus Sokolki Scutosaurus Proelginia levis llyinskoe subassemblage permiana Chroniosaurus dongusensis Putya Upper tinian Deltavjatia vjatkensis Kotelnich subassemblage Upper Lower Upper Lower Upper Suchonica vladimiri Sundyr assemblage LLU Sukhonian Ulemosaurus svijagensis Isheevo assemblage Permian Permian Lower Urzhumian Ocher Estemmenosuchus sub Mezen Guadalupian Lopingian Wordian Capitanian Wuchiapingian uralensis assemblage Urzhumian Severodvinian Vyatkian Dinocephalian Theriodontian assemblage superassemblage superassemblage Titanophoneus Biarmian ian singian nian Golyusherma Ocher LU Parabradysaurus
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