Stratigraphical Occurrence of Vertebrate Remains in the Upper Devonian of Severnaya Zemlya (Russia)

Acta Geologica Polonica, Vol. 49 (1999), No.2, pp. 125-131

Stratigraphical occurrence of vertebrate remains in the Upper Devonian of Severnaya Zemlya (Russia)

ERVINS LUKSEVICS

Latvian Museum of Natural History, K. Barona Str. 4, Riga LV 1050, Latvia. E-mail: [email protected]

ABSTRACT:

LUKSEVICS, E. Stratigraphical occurrence of vertebrate remains in the Upper Devonian of Severnaya Zemlya (Russia). Acta Geologica Polonica, 49 (2), 125-131. Warszawa.

The stratigraphical distribution of vertebrate remains in the standard section of the Upper Devonian of October Revolution Island of the Severnaya Zemlya archipelago has been studied. Fossil fish and agnathans were collected in 1978 and 1979 from the outcrops of the Matusevich, Vavilov and Malyutka formations along the Matusevich, Bol' shaya and other rivers. The fish fauna consists of representatives of the main taxonomic groups of Devonian vertebrates, i.e., psammosteid heterostracans, placoderms, acan thodians, chondrichthyans, porolepiform and osteolepiform "rhipidistians", and dipnoans. S~mmarizing all available data on the distribution of vertebrates, it is possible to suggest a very late Givetian - Frasnian age for the Matusevich Formation, a late Frasnian age of the Vavilov Formation, and a latest Frasnian - early Famennian age for the Malyutka Formation of Severnaya Zemlya. Keywords: Biostratigraphy, Devonian, Arctic region, Vertebrates.

INTRODUCTION thoroughly studied groups. The distribution of fish and agnathan remains in relation to the facies The understanding of Upper Devonian verte changes of the basin, and the main taphonomical brate assemblages from the Northern Hemisphere is features of the Upper Silurian and Lower Devonian very variable. Fish and agnathans from Scotland vertebrates assemblages have been described and the Baltic sections have been studied for almost (KARATAJOTE-TALIMAA & al. 1983). The Middle two centuries and are well known. However, those and Upper Devonian deposits in October from remote regions of the Russian Arctic were dis Revolution Island have not been so completely covered only recently. Abundant remains of fossil studied, particularly the uppermost part of the vertebrates were collected on Octo~r Revolution section. Nevertheless, this part of the section is con Island of the Severnaya Zemlya archipelago during sidered to be palaeontologically very important the study of the key section of the Silurian and because of the abundance and good preservation of Devonian in 1978-1979 (KURIK & al. 1982). Until the fossil fishes (KURIK & al. 1982). now the Upper Silurian and Lower Devonian verte brates, especially acanthodians (VALIUKEVICIUS 1997), osteostracans (MARK-KuRIK & JANVIER MATERIAL 1995; AFANASSIEVA & KARATAJOTE-TALIMAA 1997; AFANASSIEVA 1998) and thelodonts (KARATAJOTE In the northern part of October Revolution TALIMAA & MARss 1997), have been the most Island, large collections of vertebrate remains from 126 ERVINS LUKSEVICS

Matusevich have been analysed from 19 levels within the Matusevich Formation; additional mate rial from other sites, most probably representing the lowermost part of the same formation (KURIK, pers. comm. ), is under study. The outcrops of the Vavilov Formation, which are composed of red coloured sandstone, siltstone, and argillites with rare layers of grey limestone, as well as those of the Malyutka Formation, of similar composition, pro .1 vided much poorer fossil material, which has been collected from only 5 levels. The fauna of fishes and agnathans from the Upper Devonian deposits of October Revolution Island in the collections studied consists mainly of antiarchs, but contains members of all the main groups of Devonian vertebrates, such as psam mosteid heterostracans, placoderms, acanthodians, chondrichthyans, porolepiform and osteolepiform "rhipidistians", and dipnoans. Placoderms belong Fig. 1. Map of the central part of the Severnaya Zemlya archi ing to two genera of antiarchs, Asterolepis and pelago (from the Geological Map of the USSR, reduced for this Bothriolepis, dominate the assemblages numeri publication); numbers on the map indicate the localities: 1, cally. Disarticulated bones of the head and trunk October Cape, locality 139; 2, right tributary of the river armour, plates of the pectoral fin armour and, in Strelka, locality 12; 3, river Bol'shaya, locality 14; 4, river several cases, complete armours of various species Bol'shaya, locality 21; 5, river Matusevich, localities 7,17,18, of Bothriolepis make up the largest part of the ver 18a, 19; Symbol in rectangle: glaciers tebrate remains. The placoderms belonging to this genus are found in all stratigraphic units of the Upper Devonian of Severnaya Zemlya. the the Middle and Upper Devonian sections were collected by the participants of two expeditions: BIOSTRATIGRAPHY mainly by V.N. KARATAJOTE-TALIMAA and J. V ALIUKEVICUS in 1978, and by E. MARK-KuRIK Two assemblages, characterized by faunal ele and A. KHAPILIN in 1979. The 1978 expedition col ments that have been found in association with sev lected fish remains mainly from the outcrops along eral species of Bothriolepis, were used as a basis the river Matusevich, localities 7, 17, 18, 18a, and for further analysis. The first assemblage, which is 19 (5, Text-fig. 1). This collection is kept at the found throughout the Matusevich Formation, and Institute of Geology of Lithuania, Vilnius. Fossil in the lower part of the Vavilov Formation, differs vertebrates excavated in 1979 are stored at the from the second, younger one, which occurs in the Institute of Geology, Tallinn Technical University, Malyutka Formation, in the presence of psam and come from several localities: site number 139 mosteid heterostracans. Everywhere on the East at October Cape; locality number 12, the outcrop European Platform, the simultaneous appearance on the right tributary of the river Strelka, sites num of psammosteids and bothriolepidoid antiarchs ber 14 and 21 along the river Bol'shaya. Specimen indicates a Givetian - Frasnian age (LYARSKAYA & number prefixes in the text denote their respective LUKSEVICS, 1992), whereas psammosteids are not institutions: LGI, Institute of Geology of known in the Famennian deposits. Lithuania, Vilnius, Lithuania; Pi, Institute of The oldest fish remains belonging to the family Geology, Tallinn Technical University, Estonia. Bothriolepididae have been found in Member 2 of Most of the studied material comes from the the Matusevich Formation (Text-fig. 2; locality 7, Matusevich Formation, which comprises mainly bed 72). These are indeterminable fragments of the red coloured and, more rarely, light grey or green pectoral fin armour plates of a bothriolepid antiarch. ish-grey, sandstone, clay and siltstone. The verte The vertebrate assemblage from the upper part brate remains from outcrops along the river of Member 2 and the lower part of Member 3 of the UPPER DEVONIAN VERTEBRATE REMAINS 127

en Q) Q) .c ·c ....E Layer Assemblage Distribution of taxa EZJ] Q) E Rocks §~ 0 Q) Cf) Ll.. :2 -l ~ 2 I'll .:.... ':.::.:::.,.:.: ~ ~...... VI I . ::::I 3 ~ ~ I'll ::E .;. :',.' .: ...... ~ 4

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Fig. 2. Distribution of vertebrates in the section along the river Matusevich. Horizontal dotted lines separate several veltebrate assemblages; stratigraphical column simplified after KURSHS (1982) (Matusevich Formation members are given by V. KURSS, unpublished data); lithological symbols: 1 - sandstone, 2 - siltstone, 3 - clay, 4 -limestone, 5 - marl; abbreviations. B. - Bothriolepis, Form. - Formation, Ps. - Psammosteus 128 ERVINS LUKSEVICS

Matusevich Formation (locality 19, beds 60 - 63) subspecies, Bothriolepis leptocheira curonica, is richer. It comprises, in addition to abundant and which is found in Latvia. well preserved skeletons of Bothriolepis cf. obrutschewi GROSS, the remains of a second repre sentative of the same genus, Bothriolepis sp. 1 AGE AND CORRELATION WITH OTHER (single nuchal plate), the heterostracan REGIONS Psammosteus cf. maeandrinus AGASSIz, acantho dians, chondrichthyans (tooth of 'Stethacanthus' The dating of almost all the Devonian forma type), the porolepiform sarcopterygians including tions of Severnaya Zemlya is based on vertebrate scales of Glyptolepis sp., as well as indeterminable assemblages and on correlation with the sections of fragments of dipnoan bones. the Taimyr Peninsula, Spitsbergen and the East The remaining lower part of Member 3 of the European Platform (MATUKHIN & al. 1997). The Matusevich Formation is characterized by the position of the Givetian - Frasnian boundary is still occurrence of indeterminable remains of uncertain. It was previously suggested that the Bothriolepis sp. 2 (locality 19, bed 50). Middle - Upper Devonian transition in the The upper part of Member 3 of the Matusevich Severnaya Zemlya section lays within the ,1I I Formation (locality 19, beds 33 - 49) yields a rich Gremyaschiy Formation, and that the overlying vertebrate assemblage characterized by the presence deposits of the Matusevich, Vavilov and Malyutka of a bothriolepid which closely resembles formations, on the limited palaeontological data Bothriolepis trautscholdi JAEKEL from the Middle available, belonged to the Upper Devonian (KURIK Frasnian of the Main Devonian Field. In addition, & al. 1982). At the same time, the probable occur the assemblage contains other placoderm fishes, rence of Famennian deposits in Severnaya Zemlya such as two other forms of Bothriolepis, Asterolepis was also suggested (KURIK & al. 1982). New data cf. maxima AGASSIZ, known from the Nairn on the distribution of the Lower and Middle Sandstone of Scotland, Eastmanosteus? sp., as well Devonian placoderms (MARK-KuRIK 1998), as as the sarcopterygians Holoptychius sp., well as the Upper Devonian bothriolepidoid Eusthenopteron sp., and dipnoans. antiarchs, permits correlation of the lowermost part The upper part of the Matusevich Formation, of the Matusevich Formation (including Member 1 members 4 and 5 (locality 19, beds 1 - 33; locality and part of Member 2) with part of the Gauja 17), possibly contains a poor assemblage, yielding Formation, and correlation of the remainder of only a few taxa including Bothriolepis cf. maxima Member 2 and the lowermost part of Member 3 of GROSS (probably a new species), three other the Matusevich Formation, with the Amata species of Bothriolepis, Holoptychius sp. and Formation of the Baltic succession respectively. indeterminable acanthodians. Unfortunately, the data on the distribution of The lower part of the Vavilov Formation (local vertebrate assemblages are insufficient to locate ity 18) contains a slightly different assemblage. the boundary between the Givetian and Frasnian in Fish and agnathan remains recovered from a lime the October Revolution Island succession more stone (bed 5) dissolved in dilute acetic acid include precisely. The oldest placoderm remains belonging tesserae of a heterostracan similar to Psammosteus to the genus Bothriolepis have been reported from falcatus OBRUCHEV from the East European the Eifelian of China (PAN 1981). In Antarctica, a Platform, disarticulated plates of small succession of 23 bothriolepidid taxa was recorded Bothriolepis, and abundant scales of Holoptychius in the Givetian - Lower Frasnian succession sp. The last remains of psammosteids in the section (YOUNG 1988), but in Laurussia bothriolepidids occur in this bed. appeared slightly later. The first certain remains of Only one species of vertebrate, namely bothriolepidids (Bothriolepis prima GROSS, B. Bothriolepis leptocheira TRAQuAIR, represented by obrutschewi) from the Main Devonian Field occur articulated plates of the trunk, head and pectoral fin in the Amata Regional Stage (the Amata Formation armour, has been identified from the sandstones of of Latvia and Estonia, the upper part of the Sven the Malyutka Formation (locality 18a, upper part). toji Formation of Lithuania, and the Staritsa Beds This species is most probably represented in of the Yam-Tesovo Formation in the Leningrad Severnaya Zemlya by a new subspecies, which dif Region of Russia), although questionable remains fers slightly from the nominal subspecies, described of bothriolepidids have been reported from the from Scotland (MILES 1968), and from the other underlying Gauja Formation (KARATAJDTE- UPPER DEVONIAN VERTEBRATE REMAINS 129

TALlMAA 1966). The age of the Gauja Formation is press, IVANOV & LUKSEVICS 1996), of the Gauja still under discussion and hence the Givetian - Formation, or even at some level in the Amata Frasnian boundary has been placed variously at the Formation (see discussion in MARK-KuRIK 1998). base (SOROKIN 1981), or at the top (ESIN & al., in On a basis of the analysis of the distribution of ver-

Se Sta Baltic sequence Placoderm Severnaya Zeml va sequence ne ge Regional Beds zonation Formations Members s Stages U Sl<:ervelis Ketleri B. ciecere P F Zagare interzone a Svete ------P m Tervete e Muri B. ornata E n Muri ------n Akmene Phyllolepis R 1 Kursa a Joniskis interzone ------1------n Eleja B. leptocheira D Purvi.Q.i (=curonica) Malyutka Amula E Stipinai interzone Vavilov Pamusis 1------5 V F Katlesi B. maxima 4 1------r AI'tovo 0 a Buregi s Daugava Il'men' B. trautscholdi N n Svinord Matusevich 1 Porkhov I a Dubniki 3 n Chudovo interzone A Plavi.Q.as Pskov B. cellulosa Snetnaya Gora N Amata Bothriolepis 1------prima-obrutschewi 2 1--- 1-- 1------Gi Gauja Asterolepis ? 1 D2 ve ornata ti Abava* Watsonosteus Gremyaschiy an Burtnieki Pycnosteus tuberculatus * * Vatutin

Fig. 3. Preliminary correlation of the Baltic and Severnaya Zemlya Middle - Upper Devonian sections based on the distribution of verte brates (correlation of the Gremiaschiy Formation after MARK-KuRIK, 1998); *Regional Substage not officially accepted by the Baltic Stratigraphic Association; **Heterostracan Zone 130 ERVINS LUKSEVICS tebrate assemblages and conodonts from South the gigas conodont Zone of the Standard Conodont Timan, the Givetian - Frasnian boundary in the Zonation (IVANOV & LUKsEvIcs 1996). Main Devonian Field could not be traced below the The next vertebrate-based zone in the sequence base ofthe Amata Formation (EsIN & al., in press). of the Main Devonian Field, the Bothriolepis lep Therefore, it seems that bothriolepidids probably tocheira zone [previously known as the B. curoni appeared on the East European Platform at the end ca zone: recent revision of the genus has shown of the Late Givetian. The Early Frasnian age of the that B. curonica GROSS is con specific with B. lep oldest remains of bothriolepidids from North tocheira (see LUKsEvIcs, in press)], corresponds to America is confirmed by their occurrence within the triangularis conodont Zone. Bothriolepis lep the middle asymmetricus conodont Zone tocheira from the upper part of the Malyutka (SANDBERG & al. in YOUNG 1988). Formation indicates a Famennian age at least for The vertebrate assemblage of the Gremyaschiy this part of the succession. Summarizing all of the Formation, which includes p1acoderms such as the available data on the distribution of vertebrates, it antiarchs Asterolepis ex gr. dellei GROSS and A. cf. is possible to suggest a very late Givetian - early ornata EICHWALD sensu AGASSIZ (KURIK & al. Famennian age for the stratigraphical interval com 1982), and Watsonosteus sp.?, Homostius sp., prising the Matusevich, Vavilov and Malyutka for Byssacanthus? sp. (MARK-KuRIK 1998), is similar to mations of Severnaya Zemlya. that from the uppermost Givetian of the Baltic area. Unfortunately, the lowermost part of the Matusevich Formation is insufficiently character Acknowledgements ized by fossils to permit the Givetian - Frasnian boundary to be precisely located. The occurrence of Warm thanks are due to Dr. VALENTINA KARATAJOTE Bothriolepis cf. obrutschewi in members 2 and 3 of TALIMAA (Institute of Geology of Lithuania, Vilnius) and the Matusevich Formation (see MARK-KuRIK 1998) to Dr. ELGA MARK-KuRIK (Institute of Geology, Tallinn makes it possible to correlate this part of the section Technical University), who donated their material for with the Amata Formation of the Baltic succession investigation; to Professor VISVALDIS KURSS (Institute of (Text-fig. 3), the Ust'-Chirka and the base of the Geology, University of Latvia, Riga) and Dr. VLADIMIR Ust'-Yarega formations of Middle Timan, as well MENNER (Institute of Geology and Exploitation of as with the Kumushka Formation of North Timan. Combustible Fuels, Moscow), who provided stratigraph The distribution of the assemblage containing ic information; Dr. PER E. AHLBERG (Natural History Bothriolepis cf. trautscholdi allows the upper part Museum, London), who improved my English text. I am of Member 3 of the Matusevich Formation to be greatly indebted to Dr. ELGA MARK-KuRIK and anony correlated with the Lower - Middle Frasnian inter mous referees for helpful discussions and critical review val, which corresponds to the Dubniki and Daugava of the manuscript. My trip to Great Britain to make a Regional stages of the Main Devonian Field, and comparative study of bothriolepidoids was financed by possibly to the Rassokha Formation of North NATO grant OUTR. LG 961235. The travel expenses to Timan. However, the occurrence of another placo the symposiums in St. Petersburg and Warsaw were derm, Asterolepis cf. maxima AGASSIZ, a taxon financed by UNESCO-IUGS IGCP Project 406. originally described from Scotland, and closely This is a contribution to the IGCP Project 406 resembling A. radiata ROHON from the Amata and "Circum-Arctic Palaeozoic Vertebrates". Plavipas formations of the Baltic, but greatly differ ing fromA. syasiensis LYARSKAYA, which occurs in the Dubniki Formation together with B. trautschol REFERENCES di, contradicts such a correlation. The age of the upper part of the Matusevich AFANASSIEVA, O.B. 1998. New data on osteostracans Formation, consisting of members 4 and 5, could be from Severnaya Zemlya (Russia). In: M. GINTER, & determined as late Frasnian, on the basis of an M.V.H. WILSON, (Eds) , Circum-Arctic Palaeozoic assemblage yielding Bothriolepis cf. maxima; these Faunas and Facies. Special Publication 4 of deposits correlate well with those of the Katlesi and Ichthyolith Issues, pp. 2-3. Warsaw. Pamusis regional stages of the Baltic (the Snezha AFANASSIEVA, O.B. & KARATAJOTE-TALIMAA, V.N. 1997. and Prilovat' formations of northwestern Russia). Osteostracans of Severnaya Zemlya. In: M.V.H. The Bothriolepis maxima Zone of the vertebrate WILSON, (Ed.), Circum-Arctic Palaeozoic zonation of the Main Devonian Field corresponds to Vertebrates: Biological and Geological Significance.

J, UPPER DEVONIAN VERTEBRATE REMAINS 131

Special Publication 2 of Ichthyolith Issues, p. 10. deposits of Latvia. In: V.S. SOROKIN (Ed.), Buckow. Palaeontologiya i stratigrafiya fanerozoya Latvii i ESIN, D., GINTER, M., IVANov, A., LEBEDEv, 0., Baltiyskogo morya, pp. 46-62. Zinatne; Riga. [In LUKSEVICS, E., AVKHIMOVICH, V., GOLUBTSOV, V. & Russian]. PETUKHOVA, L. (in press). Vertebrate correlation of MARK-KuRIK, E. 1998. Placoderms of the Lower and the Upper Devonian on the East European Platform. Middle Devonian of Severnaya Zemlya. In: M. Courier Forschungsinst. Senckenberg. GINTER, & M.V.H. WILSON, (Eds) , Circum-Arctic IVANov, A. & LUKSEVICS, E. 1996. Late Devonian verte Palaeozoic Faunas and Facies. Special Publication 4 brates of the Timan. Daba un muzejs, 6, 22-32. Riga. of Ichthyolith Issues,pp. 33-35. Warsaw. KARATAJOTE-TALIMAA, V.N. 1966. Bothriolepids of MARK-KuRIK, E. & JANVIER, P. 1998. Early Devonian Sventoji Regional Stage of Baltic. In: A.A. GRIGELIS osteostracans from Severnaya Zemlya, Russia. (Ed.), Palaeontologiya i stratigrafiya Pribaltiki i Journal of Vertebrate Paleontology, 15 (3), pp. 449- Byelorussii, I (VI), 191-279. Vilnius. [In Russian]. 462. KARATAJOTE-TALIMAA, VN., MARK-KuRIK, E., KURSHS MATUKHIN, R., MENNER, V., ABUSHIK, A., KARATAJOTE V., MATUKHIN R.G., & MENNER V.V. 1983. Facial TALIMAA, V., LUKSEVICS, E., MANNIK, P., MARK relations and embedding types of vertebrates in the KURIK,E.,MARSS, T.,MoDZALEVSKAYA, T.,NEsToR, Upper Silurian and Lower Devonian of Severnaya H., SOBOLEV, N., STUKALINA, G., & VALIUKEVICIUS, Zemlya. In: D.L. KALJO, & E.R. KLAAMANN, (Eds) , J. 1997. Dating of the Silurian and Devonian Theory and practice of ecostratigraphy, 251-258. Formations on Severnaya Zemlya. In: A. IVANov, Valgus; Tallinn. [In Russian]. M.V.H. WILSON, & A. ZHURAVLEV, (Eds) , KARATAJOTE-TALIMAA, V.N. & MARSS, T. 1997. Palaeozoic Strata and Fossils of the Eurasian Arctic. Thelodonts from the Silurian and Lower Devonian Special Publication 3 of Ichthyolith Issues, pp. 28- of the Severnaya Zemlya archipelago, Russian 29. St. Petersburg. Arctic. In: A. IVANov, M.V.H. WILSON, & A. MILES, R.S. 1968. The Old Red Sandstone Antiarchs of ZHURAVLEV, (Eds), Palaeozoic Strata and Fossils of Scotland: family Bothriolepididae. 130 pp. London. the Eurasian Arctic. Special Publication 3 of PAN JIANG, 1981. Devonian antiarch biostratigraphy of Ichthyolith Issues, pp. 13-14. St. Petersburg. China. Geological Magazine, 118, 69-75. London. KURIK, E., KURSHS, V., MARKOVSKY, V.A., MATUKHIN, SOROKIN, V.S. (Ed.) 1981. Devonian and Carboniferous R.G., MENNER, V.V., MODZALEVSKAYA, T.L., of the Pre-Baltic Region, pp. 1- 502. Ziniitne; Riga. PATRUNOV, D.K., SAMOILOVICH, Y.G., SMIRNOVA, [In Russian]. M.A., TALIMAA, V.N.,KHAPILIN,A.F., CHERKESOVA, VALIUKEVICIUS, J. 1997. An unusually squamated acan S.V. & ABUSHIK, A.F. 1982. Silurian and Devonian thodian from Severnaya Zemlya. In: M.V.H. stratigraphy. of Severnaya Zemlya. Trudy Inst. WILSON, (Ed.), Circum-Arctic Palaeozoic Geologii i Geofiziki SO AN SSSR, 483, pp. 65-73. Vertebrates: Biological and Geological Significance. Moscow. [In Russian]. Special Publication 2 of Ichthyolith Issues, p. 25. KURSHS, V.M. 1982. The provenances of Severnaya Buckow. Zemlya Devonian deposits. Geologiya i geofizika, YOUNG, G.C. 1988. Antiarchs (Placoderm fishes) from AN SSSR, Sibirskoe otdelenie, 2, pp. 19-27. [In the Devonian Aztec siltstone, Southern Victoria Russian with English summary]. Land, Antarctica. Palaeontographica, Abt. A, 202, LYARSKAYA, L.A. & LUKSEVICS, E. 1992. Distribution of 1-125. Stuttgart. agnathans and fishes in the Silurian and Devonian

Manuscript submitted: 15th October 1998 Revised version accepted: 20th December 1998