An overview of fossil mammalian remains from the Upper Palaeolithic site Kostenki 17 (European Russia)

E.A. Petrova and A.A. Bessudnov

Proceedings of the Zoological Institute RAS, 2025, 329(2): 90–133   ·   https://doi.org/10.31610/trudyzin/2025.329.2.90

Full text  

Abstract

For the first time, the results of the study of collections from the 1950s excavations and new palaeontological material from the multilayered Early Palaeolithic site of Kostenki 17 are presented. The fauna includes 15 species of mammals, including widespread species with a Holarctic range, representatives of deciduous and mixed forests, and migrants from arid areas – steppes, semi-deserts.

Key words

biodiversity, multilayered Upper Paleolithic site Kostenki 17, Late Pleistocene mammal fauna, Russian Plain

Submitted February 20, 2025  ·  Accepted April 14, 2025  ·  Published June 3, 2025

References

Alekseeva E.V. and Baryshnikov G.F. 2020. Late Pleistocene and Holocene remains of carnivorous mammals (Carnivora) from the Bliznets Cave in southern part of the Russian Far East. Proceedings of the Zoological Institute of the Russian Academy of Sciences, 324: 388–444. [In Russian]. https://doi.org/10.31610/trudyzin/2020.324.3.388

Anikovich M.V., Popov V.V. and Platonova N.I. 2008. Paleolithic of Kostenki-Borschevo region in the context of Upper Paleolithic of Europe. Proceedings of the Kostenki-Borschevo archaeological expedition IHMC RAS. Vol. 1. Nestor-History, St. Petersburg, 304 p. [In Russian].

Aristov A.A. and Baryshnikov G.F. 2001. Mammals of Russia and Adjacent Territories. Carnivores and Pinnipeds. Zoological Institute RAS, Saint Petersburg, 560 p. [In Russian].

Averianov A.O. 1995. Late Pleistocene hare, Lepus tanaiticus (Lagomorpha: Leporidae) of Siberia. Proceedings of the Zoological Institute of the Russian Academy of Sciences, 263: 121–162. [In Russian].

Averianov A.O. 1998. Late pleistocene hares (Lepus) of the Russian Plain. Illinois State Museum Scientific Papers, 27: 41–68.

Averianov A.O. 2001. Lagomorphs (Mammalia) from the Pleistocene of Eurasia. Paleontological Journal, 35(2): 191–199. [In Russian].

Averianov A.O. and Kuzmina I.E. 1993. Don hare Lepus tanaiticus Gureev, 1964 from the paleolithic sites in Kostenki. Proceedings of the Zoological Institute of the Russian Academy of Sciences, 249: 66–92. [In Russian].

Averianov A.O., Kuznetsova T.V. and Nikol’skii P.A. 2003. Pleistocene hares from the East Siberian Arctic (Lagomorpha: Leporidae). Russian Journal of Theriology, 2(2): 71–76. https://doi.org/10.15298/rusjtheriol.02.2.02

Baryshnikov G. 2006. Late Pleistocene arctic fox (Alopex lagopus) from Crimea, Ukraine. Quaternary International, 142–143: 208–217. https://doi.org/10.1016/j.quaint.2005.03.018

Baryshnikov G.F. 2012. A review of fossil vertebrate remains from Pleistocene layers of Akhstyrskaya cave (North-West Caucasus). Proceedings of the Zoological Institute of the Russian Academy of Sciences, 316(2): 93–138. [In Russian]. https://doi.org/10.31610/tru-dyzin/2012.316.2.93

Baryshnikov G. 2015. Late Pleistocene Canidae remains from Geographical Society Cave in the Russian Far Eas. Russian Journal of Theriology, 14(1): 65–83. https://doi.org/10.15298/rusjtheriol.14.1.03

Baryshnikov G.F. 2020. Large mammals of Kudaro Pleistocene fauna from Caucasus. Proceedings of the Zoological Institute of the Russian Academy of Sciences, 9: 1–398. [In Russian].

Baryshnikov G.F. 2022. Fauna of the Paleolithic site of Ilskaya in the Northern Caucasus, Russia. Prehistoric Archaeology. Journal of Interdisciplinary Studies, 2: 5–52. [In Russian].

Baryshnikov G.F., Dmitrieva E.L., Krakhmalnaya T.V. and Sher A.V. 1998. Origin, evolution, systematics. In: V.E. Sokolov and L.V. Zhirnov (Eds). Saiga: phylogeny, systematics, ecology, conservation and use. Printing house of the Russian Agricultural Academy, Moscow: 9–24. [In Russian].

Baryshnikov G.F., Mol D. and Tikhonov A.N. 2009. Finding of the Late Pleistocene carnivores in Taimyr Peninsula (Russia, Siberia). Russian Journal of Theriology, 8(2): 107–113.

Baryshnikov G.F. and Tikhonov A.N. 2009. The mammals of Russia and adjacent territories. Ungulata. Part 1. Perissodactyles and Artiodactyles (suids, moschids, cervids). Nauka, Saint Petersburg, 164 p. [In Russian].

Baumann C., Starkovitch B.M., Drucker D.G., Münzel S.C., Conard N.J. and Bocherens H. 2020a. Dietary niche partitioning among Magdalenian canids in southwestern Germany and Switzerland. Quaternary Science Reviews, 227: 106032. https://doi.org/10.1016/j.quascirev.2019.106032

Baumann C., Wong G.L., Starkovich B.M., Münzel S.C. and Conard N.J. 2020b. The role of foxes in the Palaeolithic economies of the Swabian Jura (Germany). Archaeological and Anthropological Sciences, 12: 208. https://doi.org/10.1016/j.quascirev.2019.106032

Belan N.G. 1983. Late Pleistocene reindeer of the Dnieper Region. Bulletin of Zoology, 5: 20–26. [In Russian].

Belan N.G. 1985. Late Pleistocene broad-toed horse of the Desna Basin. Proceedings of the Zoological Institute of the USSR Academy of Science, 131: 50–63. [In Russian].

Bessudnov A.A. 2019. Zamyatninskaya archaeological culture of the Late Upper Palaeolithic: a myth or reality? Camera Praehistorica, 1(2): 36–69. [In Russian]. https://doi.org/10.31250/2658-3828-2019-1-36-69

Bessudnov A.A., Dinnis R., Sinitsyn A.A., Dudin A.E., Bessudnov A.N., Petrova E.A., Khlopachev G.A. and Sablin M.V. 2024. New data on the radiocarbon chronology of the sites of the Late Upper Paleolithic period of the center of the Russian Plain. Stratum plus, 1: 161–201. [In Russian]. https://doi.org/10.55086/sp241161201

Bessudnov A.A., Sinitsyn A.A., Dinnis R., Artyushenko A.A., Lada A.R., Stepanova K.N., Malyutina A.A., Bessudnova M.A., Petrova E.A. and Douka K. 2021. Kostenki 17 (Spitsynskaya): New Data on the Stratigraphy, Chronology and Conditions of the Occurrence of the Cultural Layers. Stratum plus, 1: 163–198. [In Russian]. https://doi.org/10.55086/sp211163198

Bessudnov A.A., Stepanova K.N., Artuyshenko A.A., Lada A.R., Kupriyanova M.D., Petrova E.A., Malyutina A.A., Smolkina V.S. and Sinitsyn A.A. 2023. Main directions of the new stage of work at the Paleolithic site of Kostenki 17 (Spitsynskaya). In: I.V. Shershen’ (Ed.). Rogachev readings: proceedings of the Kostenki Museum-Reserve. Press-Burger, Voronezh: 83–95. [In Russian].

Binder W.J., Thompson E.N. and Van Valkenburgh B. 2002. Temporal variation in tooth fracture among Rancho La Brea dire wolves. Journal of Vertebrate Paleontology, 22(2): 423–428. https://doi.org/10.1671/0272-4634(2002)022[0423:TVITFA]2.0.CO;2

Binder W.J. and Van Valkenburgh B. 2010. A comparison of tooth wear and breakage in Rancho La Brea sabertooth cats and dire wolves across time. Journal of Vertebrate Paleontology, 30(1): 255–261. https://doi.org/10.1080/02724630903413016

Bocherens H. 2015. Isotopic tracking of large carnivore palaeoecology in the mammoth steppe. Quaternary Science Reviews, 117: 42–71. https://doi.org/10.1016/j.quascirev.2015.03.018

Boeskorov G.G. and Baryshnikov G.F. 2013. Late Quaternary carnivores of Yakutia. Nauka, Saint Petersburg, 199 p. [In Russian].

Boriskovsky P.I. 1963. Essays on the Palaeolithic of the Don Basin: Understudied Old Stone Age Sites of Kostenki. Materials and Studies in the Archaeology of the USSR, 121: 232 p. [In Russian].

Boriskovsky P.I., Praslov N.D. and Anikovich M.V. 1982. Kostenki 17 (Spitsyn site). In N.D. Praslov and A.N. Rogachev (Eds). The Palaeolithic of the Kostenki-Borshchevo Region on the Don 1879–1979: results of field investigations. Nauka, Leningrad, 181–186. [In Russian].

Boulbes N. and Van Asperen E.N. 2019. Biostratigraphy and Palaeoecology of European Equus. Frontiers in Ecology and Evolution, 7: 301. https://doi.org/10.3389/fevo.2019.00301

Bronk Ramsey C. 2009. Bayesian analysis of radiocarbon dates. Radiocarbon, 51: 337–360. https://doi.org/10.1017/S0033822200033865

Burova N.D., Bessudnov A.A. and Bessudnov A.N. 2019. Zooarchaeological analysis of large mammals bone remains from the Late Upper Palaeolithic site of Divnogor’ye 1 (Voronezh Region). Camera praehistorica, 2(3): 40–65. https://doi.org/10.31250/2658-3828-2019-2-40-65

Cirilli O., Machado H., Arroyo-Cabrales J., Barrón-Ortiz C.I., Davis E., Jass C.N., Jukar A.M., Landry Z., Marín-Leyva A.H., Pandolfi L. et al. 2022. Evolution of the Family Equidae, Subfamily Equinae, in North, Central and South America, Eurasia and Africa during the Plio-Pleistocene. Biology, 11: 1258. https://doi.org/10.3390/biology11091258

Croitor R.V. 2010. The History of Reindeer in the Palaeolithic of Moldova. Stratum plus, 1: 137–165. [In Russian].

Croitor R. 2018. Plio-Pleistocene deer of Western Palearctic: taxonomy, systematics, phylogeny. Institute of Zoology of the Academy of Sciences of Moldova, Chişinău, 140 p.

Dalén L., Nystrom V., Valdiosera C., Germonpre M., Sablin M., Turner E., Angerbjorn A., Arsuaga J.L. and Gotherstrom A. 2007. Ancient DNA Reveals Lack of Postglacial Habitat Tracking in the Arctic Fox. Proceedings of the National Academy of Sciences, 104: 6726–6729. https://doi.org/10.1073/pnas.0701341104

David A.I. 1979. Animal remains from excavations in the Brynzeny Grotto 1. In I.M. Gania (Ed.). Micro- and macrofauna of the Late Phanerozoic of the southwest of the USSR. Shtiintsa, Kishinev: 42–57. [In Russian].

David A.I. 1980. Theriofauna of the Pleistocene of Moldova. Shtiintsa, Kishinev, 185 p. [In Russian].

Ding L., Chen C., Wang H. and Zhang B. 2016. Complete mitochondrial DNA sequence of Lepus tolai (Leporidae: Lepus). Mitochondrial DNA Part A, 27(3): 2085–2086. https://doi.org/10.3109/19401736.2014.982568

Doan K., Niedziałkowska M., Stefaniak K., Sykut M., Jędrzejewska B., Ratajczak-Skrzatek U., Piotrowska N., Ridush B. et al. 2022. Phylogenetics and phylogeography of red deer mtDNA lineages during the last 50 000 years in Eurasia. Zoological Journal of the Linnean Society, 194: 431–456. https://doi.org/10.1093/zoolinnean/zlab025

Dudin A.E. 2018. Planigraphy of the third complex of the cultural layer Ia on the site of Kostenki 11. Transactions of the Institute for Material Culture, 17: 45–54. [In Russian].

Dudin A.E. and Fedyunin I.V. 2019. The third bone-earth dwelling complex of the Kostenki 11/Ia site. In: S.N. Lisitsyn and N.I. Platonova (Eds). Man and mammoth in the Paleolithic of Europe. Part II. The Dnieper-Don historical and Cultural region. Ars longa, St. Petersburg: 221–235. [In Russian].

Englund J. 2006. Cranial and skeletal size in red foxes, Vulpes vulpes (Carnivora, Canidae) in areas with large variation in food abundance. Russian Journal of Theriology, 5(1): 25–34.

Ermolova N.M. 1978. Theriofauna of the Angara Valley in the Late Anthropogene. Nauka, Novosibirsk, 220 p. [In Russian].

Fedorova R.V. 1963. Natural conditions during the period of habitation of Upper Paleolithic man in the area of Kostenki village, Voronezh region, according to the data of spore-pollen analysis of sediments from the Spitsyn site (Kostenki XVII). Essays on the Palaeolithic of the Don Basin: Understudied Old Stone Age Sites of Kostenki. Materials and Studies in the Archaeology of the USSR, 121: 220–229. [In Russian].

Flerov K.K. 1979. Systematics and evolution. V.E. Sokolov (Ed.). Bison, Nauka, Moscow: 49–90. [In Russian].

Flower L.O.H. 2014. Canid Evolution and Palaeoecology in the Pleistocene of Western Europe, with Particular Reference to the Wolf Canis lupus L. 1758. Ph.D. thesis. Royal Holloway University of London, London, 558 p.

Flower L.O.H. and Schreve D.C. 2014. An investigation of palaeodietary variability in European Pleistocene canids. Quaternary Science Reviews, 96: 188–203. https://doi.org/10.1016/j.quascirev.2014.04.015

Flower L.O.H., Schreve D.C. and Lamb A.L. 2021. Nature of the beast? Complex drivers of prey choice, competition and resilience in Pleistocene wolves (Canis lupus L., 1754). Quaternary Science Reviews, 272: 107212. https://doi.org/10.1016/j.quascirev.2021.107212

Fordham D.A., Browna S.C., Canteria E., Austina J.J., Lomolinoe M.V., Haythornea S., Armstrongg E., Bocherensh H., Manicaj A., Rey-Iglesiad A., Rahbekb C., Nogués-Bravob D. and Lorenzend E.D. 2024. 52,000 years of woolly rhinoceros population dynamics reveal extinction mechanisms. PNAS, 121(24): e2316419121. https://doi.org/10.1073/pnas.2316419121

Foronova I.V. 1990. Fossil horses of the Kuznetsk Basin. Publishing house of the Institute of Geology and Geophysics of the Siberian Branch of the USSR Academy of Sciences, Novosibirsk, 131 p. [In Russian].

Foronova I.V. 2001. Quaternary mammals of the southeast of Western Siberia (Kuznetsk Basin): phylogeny, biostratigraphy, paleoecology. Publishing House of the Siberian Branch of the Russian Academy of Sciences, Branch “Geo”, Novosibirsk, 243 p. [In Russian].

Fox-Dobbs K., Leonard J. and Koch P. 2008. Pleistocene megafauna from eastern Beringia: paleoecological and paleoenvironmental interpretations of stable carbon and nitrogen isotope and radiocarbon records. Palaeogeography, Palaeoclimatology, Palaeoecology, 261: 30–46. https://doi.org/10.1016/j.palaeo.2007.12.011

Fritts S.H. and Mech L.D. 1981. Dynamics, movements, and feeding ecology of a newly protected wolf population Landon in northwestern Minnesota. Wildlife Monographs, 80: 3–79.

Gavrilov K.N. 2015. “Dwellings” of the Anosovka-Mezin type: origin and interpretation. Stratum plus, 1: 187–203. [In Russian].

Geffen E., Waidyaratne S., Dalén L., Angerbjörn A., Vila C., Hersteinsson P., Fuglei E., White P.A., Goltsman M., Kapel C.M.O. et al. 2007. Sea Ice Occurrence Predicts Genetic Isolation in the Arctic Fox. Molecular Ecology, 16: 4241–4255. https://doi.org/10.1111/j.1365-294x.2007.03507.x

Geist V. 1998. Deer of the World: their Evolution, Behavior and Ecology. Stackpole Books, Mechanicsburg: 1–416.

Germonpre M. 1993. Osteometric data on Late Pleistocene mammals from the Flemish Valley, Belgium. Department of Paleontology Royal Belgian Institute of Natural Sciences, Brussels, 135 p.

Germonpre M. and Sablin M.V. 2004. Systematics and osteometry of Late Glacial foxes from Belgium. Bulletin de L’ Insitut royal des Sciences naturelles de Belgique, Sciences de la Terre, 74: 175–188.

Giannoulis T., Stamatis C., Tsipourlianos A. and Mamuris Z. 2018. Mitogenomic analysis in European brown hare (Lepus europaeus) proposes genetic and functional differentiation between the distinct lineages. Mitochondrial DNA Part A, 29(3): 353–360. https://doi.org/10.1080/24701394.2016.1278540

Gimranov D.O., Kosintsev P.A. and Gasilin V.V. 2015. Species diagnostics of corsac (Vulpes corsac), fox (Vulpes vulpes), and arctic fox (Vulpes lagopus) according to ontological characteristics of mandible teeth. Zoologicheskiy Zhurnal, 94(11): 1338–1350. [In Russian]. https://doi.org/10.7868/S0044513415090093

Gittleman J.L. and Van Valkenburgh B. 1997. Sexual dimorphism in the canines and skulls of carnivores: effects of size, phylogeny, and behavioural ecology. Journal of Zoology, 242: 97–117. https://doi.org/10.1111/j.1469-7998.1997.tb02932.x

Gromova V.I. 1949. History of horses (genus Equus) in the Old World. Proceedings of the Paleontological Institute of the USSR Academy of Science, 17(1): 1–374. [In Russian].

Gureev A.A. 1964. Fauna of the USSR. Mammals, 3(10). Lagomorphs (Lagomorpha). Nauka, Moskva–Leningrad, 276 p. [In Russian].

Haynes G. 2017. Finding meaning in mammoth age profiles. Quaternary International, 443: 65–78. https://doi.org/10.1016/j.quaint.2016.04.012

Hoffecker J.F., Holliday V.T., Stepanchuk V.N. and Lisitsyn S.N. 2018. The hunting of horse and the problem of the Aurignacian on the central plain of Eastern Europe. Quaternary International, 492: 53–63. https://doi.org/10.1016/j.quaint.2018.05.044

Hoffecker J.F., Kuzmina I.E., Syromyatnikova E.V., Anikovich M.V., Sinitsyn A.A., Popov V.V. and Holliday V.T. 2010. Evidence for kill-butchery events of early Upper Paleolithic age at Kostenki, Russia. Journal of Archaeological Science, 37: 1073–1089. https://doi.org/10.1016/j.jas.2009.12.008

Kosintsev P.A. 2007. Late Pleistocene large mammal faunas from the Urals. Quaternary International, 160: 112–120. https://doi.org/10.1016/j.quaint.2006.09.012

Kosintsev P.A. and Borodin A.V. 1990. Theriofauna of the eastern slope of the Northern Urals in the Late Pleistocene and Holocene. Proceedings of the Zoological Institute of the USSR Academy of Sciences, 212: 120–134. [In Russian].

Kurtén B. 1965. The carnivora of the Palestine caves. Acta zoologica Fennica, 107: 1–74.

Kuzmina I.E. 1975. Some data on mammals of the Middle Urals in the Late Pleistocene. Bulletin of the Commission for the Study of the Quaternary, 43: 63–77. [In Russian].

Kuzmina I.E. 1997. Horses of North Eurasia from the Pliocene till the present time. Proceedings of the Zoological Institute of the Russian Academy of Sciences, 273: 1–221. [In Russian].

Kuzmina I.E. and Sablin M.V. 1993. Arctic foxes of the Late Pleistocene from the Upper Desna River. Proceedings of the Zoological Institute of the Russian Academy of Sciences, 249: 93–104. [In Russian].

Kuzmina I.E. and Sablin M.V. 1994. Wolf, Canis lupus L. from the Late Paleolithic sites Kostenki on the Don River. Proceedings of the Zoological Institute of the Russian Academy of Sciences, 256: 44–58. [In Russian].

Landon D.B., Waite C.A., Peterson R.O. and Mech L.D. 1998. Evaluation of age determination techniques for Gray wolves. Journal of Wildlife Management, 62(2): 674–682. https://doi.org/10.2307/3802343

Larsson P., von Seth J., Hagen I.J., Götherström A., Androsov S., Germonpré M., Bergfeldt N., Fedorov S., Eide N.E., Sokolova N. et al. 2019. Consequences of Past Climate Change and Recent Human Persecution on Mitogenomic Diversity in the Arctic Fox. Philosophical Transactions of the Royal Society B, 374: 20190212. https://doi.org/10.1098/rstb.2019.0212

Lazarev P.A. 1980. Anthropogene horses of Yakutia. Science, Moscow, 189 p. [In Russian].

Leonard J.A., Vilà C., Fox-Dobbs K., Koch P.L., Wayne R.K. and Van Valkenburgh B. 2007. Megafaunal extinctions and the disappearance of a specialised wolf ecomorph. Current Biology, 17: 1146–1150. https://doi.org/10.1016/j.cub.2007.05.072

Levkovskaya G.M., Shumilovskikh L.S., Anikovich M.V., Platonova N.I., Hoffecker J.F., Lisitsyn S.N., Pospelova G.A., Kuzmina I.E. and Sanko A.F. 2015. Supra-regional correlations of the most ancient paleosols and Paleolithic layers of Kostenki-Borschevo region (Russian Plain). Quaternary International, 365: 114–134. https://doi.org/10.1016/j.quaint.2014.11.043

Lipecki G. and Wojtal P. 2015. Carnivores from the openair Gravettian site Kraków Spadzista. In: J. Wilczyński, G. Haynes and P. Wojtal (Eds). A Gravettian site in Southern Poland, Kraków Spadzista. ISEA PAS, Krakow: 117–157.

Marques J.P., Seixas F.A., Farelo L., Callahan C.M., Good J.M., Montgomery W.I., Reid N., Alves P.C., Boursot P. and Melo-Ferreira J. 2020. An annotated draft genome of the mountain hare (Lepus timidus). Genome Biology and Evolution, 12(1): 3656–3662. https://doi.org/10.1093/gbe/evz273

Meiri M., Kosintsev P., Conroy K., Meiri S., Barnes I. and Lister A. 2018. Subspecies dynamics in space and time: A study of the red deer complex using ancient and modern DNA and morphology. Journal of Biogeography, 45: 367–380. https://doi.org/10.1111/jbi.13124

Michell C.T., Pohjoismäki J.L., Spong G. and Thulin C.G. 2022. Mountain- and brown hare genetic polymorphisms to survey local adaptations and conservation status of the heath hare (Lepus timidus sylvaticus Nilsson, 1831). Scientific Data, 9(1): 667. https://doi.org/10.1038/s41597-022-01794-5

Moska M., Jakubiec J., Wierzbicki H., Strzała T. and Kozyra K. 2016. Low genetic variability of the edible ormouse (Glis glis) in Stolowe Mountains National Park (Poland) – preliminary results. Mammal Research, 61(4): 409–415. https://doi.org/10.1007/s13364-016-0282-0

Niedziałkowska M., Doan K., Górny M., Sykut M., Stefaniak K., Piotrowska N., Jędrzejewska B., Ridush B. et al. 2020. Winter temperature and forest cover have shaped red deer distribution in Europe and the Ural Mountains since the Late Pleistocene. Journal of Biogeography, 48: 147–159. https://doi.org/10.1111/jbi.13989

Niedziałkowska M., Górny M., Gornia J., Popovíc D., Baca M., Ratajczak-Skrzatek U., Kovalchuk O., Sykut M. et al. 2024. Impact of global environmental changes on the range contraction of Eurasian moose since the Late Pleistocene. Science of the Total Environment, 957: 177235. https://doi.org/10.1016/j.scito-tenv.2024.177235

Niedziałkowska M., Tarnowska E., Ligmanowska J., Jędrzejewska B., Podgórski T., Radziszewska A., Ratajczyk I., Kusza S., Bunevich A.N., Danila G., Shkvyria M., Grzybowski T. and Wózniak M. 2021. Clear phylogeographic pattern and genetic structure of wild boar Sus scrofa population in Central and Eastern Europe. Scientific Reports, 11: 9680. https://doi.org/10.1038/s41598-021-88991-1

Nuzhnyi N.D., Praslov N.D. and Sablin M.V. 2014. The first case of confirmation of successful mammoth hunting in Europe (Kostenki 1, Russia). In: G.A. Khlopachev (Ed.). The collection of archaeological sources of the Kunstkamera, 4. MAE RAS, St. Petersburg: 108–117. [In Russian].

Orlando L., Metcalf J.L., Alberdi M.T., Telles-Antunes M., Bonjean D., Otte M. et al. 2009. Revising the recent evolutionary history of equids using ancient DNA. The Proceedings of the National Academy of Sciences, 106: 21754–21759. https://doi.org/10.1073/pnas.0903672106

Panitsina V.A., Bodrov S.Y., Boulygina E.S., Slobodova N.V., Kosintsev P.A. and Abramson N.I. 2023. In Search of the Elusive North: Evolutionary History of the Arctic Fox (Vulpes lagopus) in the Palearctic from the Late Pleistocene to the Recent Inferred from Mitogenomic Data. Biology, 12: 1517. https://doi.org/10.3390/biology12121517

Patton H., Hubbard A., Andreassen K., Auriac A., Whitehouse P.L., Stroeven A.P., Shackleton C., Winsborrow M., Heyman J. and Hall A.M. 2017. Deglaciation of the Eurasian ice sheet complex. Quaternary Science Reviews, 169: 148–172. https://doi.org/10.1016/j.quascirev.2017.05.019

Patton H., Hubbard A., Andreassen K., Winsborrow M. and Stroeven A.P. 2016. The build-up, configuration, and dynamical sensitivity of the Eurasian ice-sheet complex to Late Weichselian climatic and oceanic forcing. Quaternary Science Reviews, 153: 97–121. https://doi.org/10.1016/j.quascirev.2016.10.009

Petrova E.A. 2009. History of magafauna from the Middle – Late Neoplestocene in the Volga-Kama Region. PhD thesis. Zoological Institute of the Russian Academy of Sciences, Saint Petersburg, 23 p. [In Russian].

Petrova E.A., Burova N.D. and Bessudnov A.A. 2025. Mammalian fauna of the Upper Palaeolithic site Kostenki 17 (Spitsynskaya) in the context of human hunting and gathering. Proceedings of the Zoological Institute of the Russian Academy of Sciences, 329(3) (in press).

Petrova E.A., Voyta L.L., Bessudnov A.A. and Sinitsyn A.A. 2023. An integrative paleobiological study of woolly mammoths from the Upper Paleolithic site Kostenki 14 (European Russia). Quaternary Science Reviews, 302: 107948. https://doi.org/10.1016/j.quas-cirev.2022.107948

Plasteeva N.A. and Klementyev A.M. 2017. Late Pleistocene horse Equus (Equus) ferus (Perissodactyla, Equidae) from the Cis-Baikal and Transbaikalia. Proceedings of the Zoological Institute of the Russian Academy of Sciences, 321(2): 180–198. [In Russian]. https://doi.org/10.31610/trudyzin/2017.321.2.180

Plasteeva N.A., Burova N.D. and Kosintsev P.A. 2012. Caballoid horses (Equus ferus Boddaert, 1785) from Eastern Europe in the end of Late Pleistocene. Bulletin of Moscow Society of Naturalists. Biological series, 117(5): 3–11. [In Russian].

Plis K., Niedziałkowska M., Borowik T., Lang J., Heddergott M., Tiainen J., Bunevich A., Šprem N. et al. 2022. Pan-European phylogeography of the European roe deer (Capreolus capreolus). Ecology and Evolution, 12: e8931. https://doi.org/10.1002/ece3.8931

Ponomarev D. and Puzachenko A. 2017. Changes in the morphology and morphological diversity of the first lower molar of narrow-headed voles (Microtus gregalis, Arvicolinae, Rodentia) from northeastern European Russia since the Late Pleistocene. Quaternary International, 436: 239–252. https://doi.org/10.1016/j.quaint.2015.05.047

Praslov N.D. and Rogachev A.N. (Eds). 1982. The Palaeolithic of the Kostenki-Borshchevo Region on the Don 1879–1979: results of field investigations. Nauka, Leningrad, 294 p. [In Russian].

Prilepskaya N.E., Belyaev R.I., Burova N.D., Bachura O.P. and Sinitsyn A.A. 2020. Determination of season-of-death and age-at-death by cementum increment analysis of horses Equus ferus (Boddaert, 1785) from cultural layer IVa at Upper Paleolithic site Kostenki 14 (Markina Gora) (Voronezh region, Russia). Quaternary International, 557: 110–120. https://doi.org/10.1016/j.quaint.2020.05.055

Prost S., Knapp M., Fleming J., Hufthammer A.K., Kosintsev P., Stiller M. and Hofreiter M. 2010. Short communication: a phantom extinction? New insights into extinction dynamics of the Don hare Lepus tanaiticus. Journal of Evolutionary Biology, 23: 2022–2029. https://doi.org/10.1111/j.1420-9101.2010.02062.x

Pryor A.J.E., Beresford-Jones D.G., Dudin A.E., Ikonnikova E.M., Hoffecker J.F. and Gamble C. 2020. The chronology and function of a new circular mammoth-bone structure at Kostenki 11. Antiquity, 94(374): 323–341. https://doi.org/10.15184/aqy.2020.7

Rabiniak E., Rekovets L., Kovalchuk O., Baca M., Popović D., Strzała T. and Barkaszi Z. 2023. Hares from the Late Pleistocene of Ukraine: a phylogenetic analysis and the status of Lepus tanaiticus (Mammalia, Lagomorpha). Biologia. https://doi.org/10.1007/s11756-023-01499-z

Reimer P., Austin W., Bard E., Bayliss A., Blackwell P., Bronk Ramsey C., Butzin M., Cheng H. et al. 2020. The IntCal20 Northern Hemisphere radiocarbon age calibration curve (0–55 cal kBP). Radiocarbon, 62: 725–757. https://doi.org/10.1017/RDC.2020.41

Rekovets L.I. 1985. Small mammals of Desna-Dnieper late palaeolithic. Naukova Dumka, Kiev, 166 p. [In Russian].

Rekovets L.I. and Topachevsky I.V. 1988. Lagomorphs (Lagomorpha, Mammalia) from the late palaeolithic fauna of Mezhirich. Paleontological digest, 25: 56–60. [In Russian].

Rey-Iglesia A., Pryor A.J.E., Wilson T., Teeter M.A., Margaryan A., Khaskhanov R., Le Meillour L., De Jager D., Troché G.B., Welker F., Szpak P., Dudin A.E. and Lorenzen E.D. 2025. Ancient biomolecular analysis of 39 mammoth individuals from Kostenki 11-Ia elucidates Upper Palaeolithic human resource use. Quaternary Environments and Humans, 3(1): 100049. https://doi.org/10.1016/j.qeh.2024.100049

Rogachev A.N. 1957. Multilayered sites of the Kostenki-Borshchevo region on the Don and the question of cultural development during the Upper Palaeolithic on the Russian Plain. Materials and Studies in the Archaeology of the USSR, 59: 10–134. [In Russian].

Rogachev A.N. 1962. On the Anosovka-Mezin type of Paleolithic dwellings on the Russian Plain. Brief communications of the Institute of Archeology, 92: 12–17. [In Russian].

Rogachev A.N. and Anikovich M.V. 1984. Late Paleolithic of the Russian Plain and Crimea. In P.I. Boriskovsky (Ed.). The Palaeolithic of the USSR. Archaeology of the USSR, 1. Nauka, Moscow: 162–271. [In Russian].

Sablin M.V. 1994. Late Pleistocene arctic fox (Alopes lagopus rossicus) from Kostenki in Voronezh Region. Proceedings of the Zoological Institute of the Russian Academy of Sciences, 256: 59–68. [In Russian].

Sablin M.V. 2013. The arctic foxes from Gbnnersdorf. The faunal remains from Gönnersdorf. Verlag des Römisch-Germanischen Zentralmuseums, Mainz: 277–280.

Sablin M.V. 2017. The fauna of large mammals from Eliseevichi 1 site. In G.A. Khlopachev (Ed.). Kamennyy vek Rossiysko-Belorusskogo porubezh’ya. MAE RAN, Saint Petersburg: 21–45. [In Russian].

Sablin M.V. and Kuzmina I.E. 1992. Late Pleistocene reindeer from the Upper Don. Proceedings of the Zoological Institute of the Russian Academy of Sciences, 246: 72–80. [In Russian].

Sablin M.V., Burova N.D. and Petrova E.A. 2018. Horses and ancient people: zooarchaeological investigation of Muhkai 2a. Proceedings of the Zoological Institute of the Russian Academy of Sciences, 322(3): 333–356. [In Russian]. https://doi.org/10.31610/tru-dyzin/2018.322.3.333

Sedov S.N., Sinitsyn A.A., Lev S.Yu., Bessudnov A.A., Bessudnov A.N., Sycheva S.A., Romanis T.V., Sheinkmann V.S. and Korkka M.A. 2022. Paleosols of the Upper Palaeolithic sites in the East European Plain reflect the environmental fluctuations of centennial to millennial scale during MIS 3 and MIS 2. Geomorphology, 53(5): 69–77. https://doi.org/10.31857/S0435428122050157

Sergin V.Ya. 2019. Dwellings or sanctuaries? Stratum plus, 1: 99–110. [In Russian].

Sharko F., Slobodova N., Boulygina E., Cheprasov M., Gladysheva-Azgari M., Tsygankova S., Rastorguev S., Novgorodov G., Boeskorov G., Grigorieva L., Hwang W.S., Tikhonov A. and Nedoluzhko A. 2023. Ancient DNA of the Don-hares assumes the existence of two distinct mitochondrial clades in Northeast Asia. Genes, 14: 700. https://doi.org/10.3390/genes14030700

Sinitsyn A.A. 2015. Perspectives on the Palaeolithic of Eurasia: Kostenki and related sites. In: N. Sanz (Ed.). Human Origin Sites and the World Heritage Convention in Eurasia. UNESCO, Paris and Mexico City: 163–189.

Sinitsyn A.A. 2017. Kostenki 14 (Markina gora). In: N.E. Chalykh (Ed.). Archaeological research in the Central Chernozem region 2016. Novyy vzglyad, Lipetsk: 66–69. [In Russian].

Sinitsyn A.A., Hoffecker J.F., Sinitsyna G.V., Spiridonova E.A., Guskova E.G., Forman S., Ocherednoi A.K., Bessudnov A.A., Mironov D.C. and Raynolds B. 2004. Kostenki 14 (Markina gora). In: M.V. Anikovich and N.I. Platonova (Eds). Kostenki and the Early Upper Paleolithic of Eurasia: general trends, local developments. Istoki, Voronezh: 39–59. [In Russian].

Sinitsyn A.A., Stepanova K.N. and Petrova E.A. 2019. New direct evidence of mammoth hunting from Kostenki. Prehistoric Archaeology. Journal of Interdisciplinary Studies, 1: 147–156. [In Russian]. https://doi.org/10.31600/2658-3925-2019-1-149-158

Smith S., Sandoval-Castellanos E., Lagerholm V.K., Napierala H., Sablin M., Von Seth J., Fladerer F.A., Germonpré M., Wojtal P., Miller R., Stewart J.R. and Dalén L. 2017. Nonreceding hare lines: genetic continuity since the Late Pleistocene in European mountain hares (Lepus timidus). Biological Journal of the Linnean Society, 120(4): 891–908. https://doi.org/10.1093/biolinnean/blw009

Sokolov I.I. and Rashek V.L. 1961. The development of teeth and skull as indeces of age in Saiga tatarica. Bulletin of Moscow Society of Naturalists. Biological series, 66(6): 77–98. [In Russian].

Spiridonova E.A. 2002. Palynological examination of the age of sediments in the stratigraphic column of the site of Kostenki 14 (Markina Gora). In: A.A. Sinitsyn, V.Ya. Sergin and J.F. Hoffeker (Eds). Features of the development of the Upper Paleolithic of Eastern Europe. Kostenki in the context of the Paleolithic of Eurasia 1. IHMC RAS, St. Petersburg: 237–246.

Street M. and Turner E. 2013. The faunal remains from Gönnersdorf. Verlag des Römisch-Germanischen Zentralmuseums, Mainz, 357 p.

Svistun V.I. 1968. Fauna of Late Anthropogene mammals of the Romankovo alluvial locality. Natural setting and past fauna, 4: 3–56. [In Russian].

Taberlet P., Fumagalli L., Wust-Saucy A.G. and Cosson J.F. 1998. Comparative phylogeography and postglacial colonization routes in Europe. Molecular Ecology, 7: 453–464. https://doi.org/10.1046/j.1365-294x.1998.00289.x

Tarroux A., Berteaux D. and Bêty J. 2010. Northern Nomads: Ability for Extensive Movements in Adult Arctic Foxes. Polar Biology, 33: 1021–1026. https://doi.org/10.1007/s00300-010-0780-5

Tatarinov K.A. 1966. Pleistocene Vertebrates of Podolia and Sub-Carpathians. Bulletin of the Commission for Quaternary Research Academy of Sciences USSR, 32: 51–62. [In Russian].

Urantowka A.D., Kroczak A. and Mackiewicz P. 2017. The influence of molecular markers and methods on inferring the phylogenetic relationships between the representatives of the Arini (parrots, Psittaciformes), determined on the basis of their complete mitochondrial genomes. BMC Ecology and Evolution, 17: 166. https://doi.org/10.1186/s12862-017-1012-1

Van Asperen E.N. 2010. Ecomorphological adaptations to climate and substrate in late Middle Pleistocene caballoid horses. Palaeogeography, Palaeoclimatology, Palaeoecology, 297: 584–596. https://doi.org/10.1016/j.palaeo.2010.09.007

Van Asperen E.N., Stefaniak K., Proskurnyak I. and Ridush B. 2012. Equids from Emine-Bair-Khosar Cave (Crimea, Ukraine): co-occurrence of the stenonid Equus hydruntinus and the caballoid E. ferus latipes based on skull and postcranial remains. Palaeontologia Electronica, 15(1): 5A. https://doi.org/10.26879/280

Van Ballenberghe V., Erickson A.W. and Byman D. 1975. Ecology of the timber wolf in Northeastern Minnesota. Wildlife Monographs, 43: 3–43.

Van Valkenburgh B. 1988a. Trophic diversity in past and present guilds of large predatory mammals. Paleobiology, 14(2): 155–173. https://doi.org/10.1017/S0094837300011891

Van Valkenburgh B. 1988b. Incidence of tooth breakage among large, predatory mammals. The American Naturalist, 131(2): 291–302. https://doi.org/10.1086/284790

Van Valkenburgh B. 1991. Iterative evolution of hypercarnivory in canids (Mammalia: Carnivora): evolutionary interactions among sympatric predators. Paleobiology, 17(4): 340–362. https://doi.org/10.1017/S0094837300010691

Van Valkenburgh B. 2007. Déjàvu: theevolutionoffeeding morphologies in the Carnivora. Annual Meeting of the Society for Integrative and Comparative Biology (3–7 January 2007, Phoenix, Arizona, USA). Oxford University Press, Oxford: 1–17. https://doi.org/10.1093/icb/icm016

Van Valkenburgh B. and Koepfli K.P. 1993. Cranial and dental adaptations to predation in canids. In: N. Dunstone and M.L. Gorman (Eds). Mammals as Predators. Symposia of the Zoological Society of London, 65. Oxford University Press, London: 15–37. https://doi.org/10.1093/oso/9780198540670.003.0002

Vasil’ev S.K. 2005. Deer (Genera Megaloceros, Cervus, Alces) of the Late Pleistocene of the Novosibirsk Priobye. Fauna of the Urals and Siberia in the Pleistocene and Holocene, 4: 89–112. [In Russian].

Velichko A.A., Pisareva V.V., Sedov S.N., Sinitsyn A.A. and Timireva S.N. 2009. Paleogeography of Kostenki 14 (Markina gora). Archaeology, Ethnology & Anthropology of Eurasia, 37: 25–50. https://doi.org/10.1016/j.aeae.2010.02.002

Vereshchagin N.K. and Kuzmina I.E. 1977. Remains of mammals from Paleolithic site on the Don and Desna Rivers. Proceedings of the Zoological Institute of the USSR Academy of Science, 72: 77–110. [In Russian].

Vershinina A.O., Heintzman P.D., Froese D.G., Zazula G., Cassatt-Johnstone M., Dalen L., Der Sarkissian C., Dunn S.G., Ermini L., Gamba C. et al. 2021. Ancient horse genomes reveal the timing and extent of dispersals across the Bering Land Bridge. Molecular Ecology, 23: 6144–6161. https://doi.org/10.1111/mec.15977

Vinogradov B.S. 1937. Jerboas. Publishing house of the USSR Academy of Sciences, Moscow–Leningrad, 197 p. [In Russian].

Von den Driesch A. 1976. A guide to the measurement of animal bones from archaeological sites. Peabody Museum Bulletin, 1: 1–136.

Weinstock J. 2002. Environment, body size and sexual dimorphism in late glacial reindeer. In: D. Ruscillo (Ed.). Recent advances in ageing and sexing animal bones. Oxbow Books, Oxford: 246–252.

Weinstock J., Willerslev E., Sher A., Tong W., Ho S.Y.W., Rubenstein D. et al. 2005. Evolution, systematics, and phylogeography of Pleistocene horses in the new world: a molecular perspective. PLOS Biology, 3: 241. https://doi.org/10.1371/journal.pbio.0030241

Westerman M., Krajewski C., Kear B.P., Meehan L., Meredith R.W., Emerling C.A. and Springer M.S. 2016. Phylogenetic relationships of dasyuromorphian marsupials revisited. Zoological Journal of the Linnean Society, 176(3): 686–701. https://doi.org/10.1111/zoj.12323

Wilczyński J., Wojtal P., Robličková M., Oliva M. 2015. Dolní Vȇstonice I (Pavlovian, the Czech Republic) – results of zooarchaeological studies of the animal remains discovered on the campsite (excavation 1924–52). Quaternary International, 379: 58–70. https://doi.org/10.1016/j.quaint.2015.05.059

Wojtal P. and Wilczyński J. 2015. Zooarchaeological studies of large mammal remains from Kraków Spadzista site-trench C2 and trech E1 (2011–2012 excatations). In: J. Wilczyński, G. Haynes and P. Wojtal (Eds). A Gravettian site in Southern Poland, Kraków Spadzista. ISEA PAS, Krakow: 93–111.

Zheltova M.N., Burova N.D., Zaretskaya N.E., Zaitseva G.I. and Sementsov A.A. 2021. “Hare Tracks” in the Upper Palaeolithic in the centre of the East-European Plain (an overview). Quaternary International, 587–588: 310–325. https://doi.org/10.1016/j.quaint.2020.10.017

 

© Zoological Institute of the Russian Academy of Sciences
Last modified: June 3, 2025