Skip to main content
Log in

On Time and Environment of Stephanorhinus kirchbergensis Jäger 1839 (Mammalia, Rhinoceratidae) in Altai and Northeastern Russia

  • Published:
Biology Bulletin Aims and scope Submit manuscript


The remains of the extinct Merck’s rhinoceros (Stephanorhinus kirchbergensis (Jäger 1839)), well studied in Western Europe, are rare in Russia. However, thanks to the work of a number of researchers, the geography of the finds and the reconstructed range of the species have been significantly expanded. The time of the optimal existence of Merck’s rhinoceros in Yakutia is now recognized as the Middle Pleistocene; the latest finds, dating from the beginning of the late Pleistocene, are known from the southeast of Western Siberia. We provide new radiocarbon dates for the root of a tooth and bone tissue from a previously unstudied lower jaw of the Merck’s rhinoceros from Altai (AltR), whose taxonomic identity we confirm using genomic analysis. Both dates provide an age estimate of around 40 thousand years, which corresponds to the Karginsky time (MIS 3), and are the youngest for the species on the territory of Russia. The pollen spectrum from the soil filling the bone canal characterizes plant communities of open landscapes with forest areas on the upland or in the floodplain, and reflects either local features of the environment or communities of the cold stage within the Karginsky interstadial. A second Merck’s rhinoceros from the Chondon River (ChR), in extreme northeast Yakutia, was determined by previous researchers to have lived either 45–70 thousand years ago or during the beginning of the Middle Pleistocene. Considering what habitats were available in the region, we propose that the ChR could have lived during the last—Kazantsevo—interglacial (MIS 5e) or later. Both finds, AltR and ChR, extend the temporal range of the species existence.

This is a preview of subscription content, log in via an institution to check access.

Access this article

Subscribe and save

Springer+ Basic
EUR 32.99 /Month
  • Get 10 units per month
  • Download Article/Chapter or Ebook
  • 1 Unit = 1 Article or 1 Chapter
  • Cancel anytime
Subscribe now

Buy Now

Price excludes VAT (USA)
Tax calculation will be finalised during checkout.

Instant access to the full article PDF.

Fig. 1.
Fig. 2.
Fig. 3.
Fig. 4.
Fig. 5.

Similar content being viewed by others


  1. Alekseeva, L.I., Teriofauna rannego antropogena Vostochnoi Evropy (Theriofauna of the Early Anthropogen of Eastern Europe), Moscow: Nauka, 1977.

  2. Ancient DNA: Methods and Protocols, Shapiro, B., Barlow, A., Heintzman, P.D., Hofreiter, M., Paijmans, J.L.A., and Soares, A.E.R., Eds., New York: Springer–Verlag, 2012.

    Google Scholar 

  3. Andreev, A.A., Schirrmeister, L., Tarasov, P.E., Ganopolski, A., Brovkin, V., Siegert, C., Wetterich, S., and Hubberten, H.-W., Vegetation and climate history in the Laptev Sea region (Arctic Siberia) during Late Quaternary inferred from pollen records, Quat. Sci. Rev., 2011, vol. 30, pp. 2182–2199.

    Article  Google Scholar 

  4. Arkhipov, S.A. and Votakh, M.R., History of vegetation in the Middle–Late Wurm and Holocene in the valley of the Upper Ob River, in Pleistotsen Sibiri i smezhnykh oblastei (Pleistocene of Siberia and Adjacent Regions), Moscow: Nauka, 1973, pp. 130–143.

  5. Asperen, E.N. and Kahlke, R.-D., Dietary variation and overlap in central and northwest European Stephanorhinus kirchbergensis and S. hemitoechus (Rhinocerotidae, Mammalia) influenced by habitat diversity, Quat. Sci. Rev., 2015, vol. 107, pp. 47–61.

    Article  Google Scholar 

  6. Belyaeva E.I., The remains of a fossil rhinoceros from the vicinity of Rybinsk city, Byull. Kom. Izuch. Chetvertichn. Perioda, Akad. Nauk SSSR, 1939, vol. 5, pp. 69–92.

    Google Scholar 

  7. Beug, H.-J., Leitfaden der Pollen Bestimmung für Mitteleuropa and Angrenzende Gebiete, Munich: Verlag Dr. Friedrich Pfeil, 2004.

    Google Scholar 

  8. Billia, E.M.E., First records of Stephanorhinus kirchbergensis (Jäger, 1839) (Mammalia, Rhinocerotidae) from the Kuznetsk Basin (Kemerovo, Kuzbass area, southeast of western Siberia), Boll. Paleontol. Ital., 2007, vol. 46, pp. 95–100.

    Google Scholar 

  9. Billia, E.M.E., The skull of Stephanorhinus kirchbergensis (Jäger 1839) (Mammalia, Rhinocerotidae) from the Irkutsk region (southwest eastern Siberia), Quat. Int., 2008a, vol. 179, pp. 20–24.

    Article  Google Scholar 

  10. Billia, E.M.E., Revision of the fossil material attributed to Stephanorhinus kirchbergensis (Jäger 1839) (Mammalia, Rhinocerotidae) preserved in the museum collections of the Russian Federation, Quat. Int., 2008b, vol. 179, pp. 25–37.

    Article  Google Scholar 

  11. Billia, E.M.E., The famous Stephanorhinus kirchbergensis (Jäger 1839) “Irkutsk skull” (Mammalia, Rhinocerotidae) from eastern Siberia briefly compared with those from Krapina and Warsaw (Eastern Europe), Muz. Olteniei Craiova, 2010, vol. 26, pp. 296–302.

    Google Scholar 

  12. Billia, E.M.E., Stephanorhinus kirchbergensis (Jäger, 1839) (Mammalia, Rhinocerotidae) from European Russia: a new, detailed inventory of sires and referred material, Centr. Eur. Geol., 2014, vol. 57, no. 2, pp. 165–195.

    Article  Google Scholar 

  13. Billia, E.M.E. and Zervanovȧ, J., New Stephanorhinus kirchbergensis (Jäger, 1839) (Mammalia, Rhinocerotidae) records in Eurasia. Addenda to a previous work, Geol., Paleontol., Paletnol., 2015, vol. 36, pp. 55–68.

    Google Scholar 

  14. Bocherens, H., Isotopic tracking of large carnivore palaeoecology in the mammoth steppe, Quat. Sci. Rev., 2015, vol. 117, pp. 42–71.

    Article  Google Scholar 

  15. Bronk Ramsey, C., Higham, T., Bowles, A., and Hedges, R., Improvement to the pretreatment of bone at Oxford, Radiocarbon, 2004, vol. 46, no. 1, pp. 155–163.

    Article  Google Scholar 

  16. Brown, T.A., Nelson, D.E., Vogel, J.S., and Southon, J.R., Improved collagen extraction by modified Longin method, Radiocarbon, 1988, vol. 30, pp. 171–177.

    Article  CAS  Google Scholar 

  17. Burkanova, E.M., Billia, E.M.E., and Persico, D., Stephanorhinus kirchbergensis (Jäger, 1839) (Mammalia, Rhinocerotidae) from the Po valley (Lombardia, Northern Italy): possible diet/nutrition and living conditions, Quat. Int., 2020, vol. 554, pp. 164–169.

    Article  Google Scholar 

  18. Cherskii, I.D., Description of the skull of rhinoceros different from Rh. tichorinus (Rh. Merkii Jaeg.), Zap. Imper. Akad. Nauk, 1874, vol. 25, pp. 65–74.

    Google Scholar 

  19. Chytrý, M., Horsȧk, M., Danihelka, J., Ermakov, N., German, D.A., Hȧjkev, M., Hȧjkovȧ, P., Koċí, M., Kubešovȧ, S., Lustyk, P., Nekola, J.C., Pvelkovȧ Riċȧnkovȧ, V., Preislerovȧ, Z., Resl, P., and Valachoviċ, M., A modern analogue of the Pleistocene steppe-tundra ecosystem in southern Siberia, Boreas, 2019, vol. 48, pp. 36–56. 0300-9483

  20. Dabney, J., Knapp, M., Glocke, I., Gansauge, M.-T., Weihmann, A., Nickel, B., Valdiosera, C., Garcia, N., Paabo, S., Arsuaga, J.-L., and Meyer, M., Complete mitochondrial genome sequence of a Middle Pleistocene cave bear reconstructed from ultrashort DNA fragments, Proc. Natl. Acad. Sci. U.S.A., 2013, vol. 110, pp. 15758–15763.

    Article  CAS  Google Scholar 

  21. Edgar, R.C., MUSCLE: multiple sequence alignment with high accuracy and high throughput, Nucleic Acids Res., 2004, vol. 32, pp. 1792–1797.

    Article  CAS  Google Scholar 

  22. Giterman, R.E., Istoriya rastitel’nosti Severo-Vostoka SSSR v pliotsene i pleistotsene (Vegetation History of the Northeastern USSR in Pliocene and Pleistocene), Moscow: Nauka, 1985.

  23. Grichuk, V.P., Technique for treatment of sedimentary rocks poor in organic residues for pollen analysis, Probl. Fiz. Geogr., 1940, vol. 8, pp. 53–58.

    Google Scholar 

  24. Gromova, V., New materials on the Quaternary fauna of the Volga region and the history of mammals of Eastern Europe and Northern Asia in general, Byull. Kom. Izuch. Chetvertichn. Perioda, Akad. Nauk SSSR, 1932, vol. 2, pp. 69–184.

    Google Scholar 

  25. Gromova, V.I., The remains of the Merck’s rhinoceros (Rhinoceros mercki Jaeg.) from the Lower Volga River, Tr. Paleontol. Inst., Akad. Nauk SSSR, 1935, vol. 4, pp. 91–136.

    Google Scholar 

  26. Gubin, S.V. and Zanina, O.G., Changes of soil cover during the development of ice complex deposits in the Kolyma lowland, Part 1, Kriosfera Zemli, 2013, vol. 17, no. 4, pp. 48–56.

    Google Scholar 

  27. Gubin, S.V. and Zanina, O.G., Changes of soil cover during the development of ice complex deposits in the Kolyma lowland, Part 2, Kriosfera Zemli, 2014, vol. 18, no. 1, pp. 77–82.

    Google Scholar 

  28. Kienast, F., Wetterich, S., Kuzmina, S., Schirrmeister, L., Andreev, A.A., Tarasov, P., Nazarova, L., Kossler, A., Frolova, L., and Kunitsky, V.V., Paleontological records indicate the occurrence of open woodlands in a dry inland climate at the present-day Arctic coast in western Beringia during the Last Interglacial, Quat. Sci. Rev., 2011, vol. 30, pp. 2134–2159.

    Article  Google Scholar 

  29. Kirillova, I.V., Chernova, O.F., Kukarskikh, V.V., Shidlovskiy, F.K., and Zanina, O.G., The first finding of a rhinoceros of the genus Stephanorhinus in Arctic Asia, Dokl. Biol. Sci., 2016, vol. 471, pp. 300–303.

    Article  CAS  Google Scholar 

  30. Kirillova, I.V., Chernova, O.F., van der Made, J., Kukarskih, V.V., Shapiro, B., van der Plicht, J., Shidlovskiy, F.K., Heintzman, P.D., Kolfschoten, T., and Zanina, O.G., Discovery of the skull of Stephanorhinus kirchbergensis (Jäger 1839) above the Arctic Circle, Quat. Res., 2017, vol. 3, pp. 537–550.

    Article  Google Scholar 

  31. Kirillova, I.V., Borisova, O.K., Chernova, O.F., van Kolfschoten, T., van der Lubbe, J.H.J.L., Panin, A.V., Pečnerová, P., van der Plicht, J., Shidlovskiy, F.K., Titov, V.V., and Zanina, O.G., “Semi-dwarf” woolly mammoths from the East Siberian Sea coast (continental Russia), Boreas, 2020 vol. 49, no. 2, pp. 269–285.

    Article  Google Scholar 

  32. Kosintsev, P., Mitchell, K.J., Devièse, T., van der Plicht, J., Kuitems, M., Petrova, E., Tikhonov, A., Higham, T., Comeskey, D., Turney, C., Cooper, A., van Kolfschoten, T., Stuart, A.J., and Lister, A.M., Evolution and extinction of the giant rhinoceros Elasmotherium sibiricum sheds light on late Quaternary megafaunal extinctions, Nat. Ecol. Evol., 2019, vol. 3, no. 1, pp. 31–38.

    Article  Google Scholar 

  33. Kosintsev, P.A., Zykov, S.V., Tiunov, M.P., Shpansky, A.V., Gasilin, V.V., Gimranov, D.O., and Devjashin, M.M., The first find of Merck’s rhinoceros (Mammalia, Perissodactyla, Rhinocerotidae, Stephanorhinus kirchbergensis Jager, 1839) remains in the Russian Far East, Dokl. Biol. Sci., 2020, vol. 491, pp. 47–49.

    Article  CAS  Google Scholar 

  34. Kotowski, A., Badura, J., Borówka, R.K., Stachowicz-Rybka, R., Hrynowiecka, A., Tomkowiak, J., Bieniek, B., Przybylski, B., Ciszek, D., Ratajczak, U., Urbański, K., Shpansky, A.V., and Stefaniak, K., Stephanorhinus kirchbergensis from Gorzów Wielkopolski (Poland)—preliminary data and perspectives, Proc. INQUA–SEQS 2017 Int. Conf. “Quaternary Stratigraphy and Hominids around Europe: Tautavel (Eastern Pyrenees),” Tautavel-Ufa, 2017, p. 32.

  35. Laukhin, S.A., Shilova, G.N., and Velichkevich, F.Yu., Paleobotanical characteristics and paleoclimates of the Karginsky period in the West Siberian Plain, Vestn. Arkheol., Antropol. Etnogr., 2006, vol. 7, pp. 203–225.

    Google Scholar 

  36. Laukhin, S.A., Pushkar’, V.S., and Cherepanova, M.V., The current state of reconstructions of environment in the north of Siberia in the Karginskoe period (Late Pleistocene), Byull. Mosk. O-va. Ispyt. Prir., Otd. Geol., 2012, vol. 87, no. 6, pp. 37–48.

    Google Scholar 

  37. Laukhin, S.A., Pushkar’, V.S., and Cherepanova, M.V., Correlation of natural events of the Karginsky period of Pleistocene (analogs of MIS-3) from the Ob River region to the Sea of Okhotsk, Byull. Mosk. O-va. Ispyt. Prir., Otd. Geol., 2015, vol. 90, no. 2, pp. 23–34.

    Google Scholar 

  38. Lomachenkov, V.S., Geologicheskoe stroenie i rel’ef mezhdurechii nizov’ev r. Omoloi, r. Yany i r. Chondona (Geological Structure and Relief of Interfluves of the Lower Omoloi, Yana, and Chondon Rivers), Moscow: Rosgeolfond, 1956.

  39. Lozhkin, A.V. and Anderson, P.M., Forest or no forest: implications of the vegetation record for climatic stability in Western Beringia during Oxygen Isotope Stage 3, Quart. Sci. Rev., 2011, vol. 30, pp. 2160–2181.

    Article  Google Scholar 

  40. Němec, M., Wacker, L., Hajdas, I., and Gäggeler, H., Alternative methods for cellulose preparation for AMS measurement, Radiocarbon, 2010, vol. 55, nos. 2–3, pp. 1358–1370.

    Article  Google Scholar 

  41. Panychev, V.A., Radiouglerodnaya khronologiya allyuvial’nykh otlozhenii Predaltaiskoi ravniny (Radiocarbon Chronology of Alluvial Deposits in Cis-Altai Plain), Novosibirsk: Nauka, 1979.

  42. Pozdnyakov, L.K., Daurskaya listvennitsa (Dahurian (Gmelin) Larch), Moscow: Nauka, 1975.

  43. Reimer, P.J., Bard, E., Bayliss, A., Beck, J.W., Blackwell, P.G., Bronk Ramsey, C., Buck, C.E., Cheng, H., Edwards, R.L., Friedrich, M., Grootes, P.M., Guilderson, T.P., Haflidason, H., Hajdas, I., Hatt, C., et al., IntCal13 and MARINE13 radiocarbon age calibration curves 0–50000 years calBP, Radiocarbon, 2013, vol. 55, no. 4, pp. 1869–1887.

    Article  CAS  Google Scholar 

  44. Schirrmeister, L., Siegert, C., Kuznetsova, T., Kuzmina, S., Andreev, A., Kienast, F., Meyer, H., and Bobrov, A., Paleoenvironmental and paleoclimatic records from permafrost deposits in the Arctic region of Northern Siberia, Quat. Int., 2002, vol. 89, pp. 97–118.

    Article  Google Scholar 

  45. Shpanskii, A.V., New records of Merck’s rhinoceros (Stephanorhinus kirchbergensis Jäger 1839) (Rhinocerotidae, Mammalia) in the Ob River region near Tomsk, Geosfernye Issled., 2016, no. 1, pp. 24–39.

  46. Shpanskii, A.V., Paleozoogeography of Merck’s rhinoceros (Stephanorhinus kirchbergensis Jäger 1839) (Rhinocerotidae, Mammalia), Geosfernye Issled., 2017, no. 3, pp. 73–89.

  47. Shpansky, A.V. and Billia, E.M.E., Records of Stephanorhinus kirchbergensis (Jäger, 1839) (Mammalia, Rhinocerotidae) from the Ob’ River at Krasny Yar (Tomsk region, southeast of Western Siberia), Russ. J. Theriol., 2012, vol. 1, pp. 47–55.

    Article  Google Scholar 

  48. Shpansky, A.V. and Boeskorov, G.G., Northernmost record of the Merck’s rhinoceros Stephanorhinus kirchbergensis (Jäger) and taxonomic status of Coelodonta jacuticus Russanov (Mammalia, Rhinocerotidae), Paleontol. J., 2018, vol. 52, no. 4, pp. 445–462.

    Article  Google Scholar 

  49. Stamatakis, A., RAxML version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies, Bioinformatics, 2014, vol. 30, pp. 1312–1313.

    Article  CAS  Google Scholar 

  50. Titov, V.V. and Tesakov, A.S., Late Miocene (Turolian) vertebrate faunas of the southern European Russia, in Fossil Mammals of Asia: Neogene Biostratigraphy and Chronology, Wang, X., Flynn, L.J., and Fortelius, M., Eds., New York: Columbia Univ. Press, 2013, pp. 536–543.

    Google Scholar 

  51. Troll, C.J., Kapp, J., Rao, V., Harkins, K.M., Cole, C., Naughton, C., Morgan, J.M., Shapiro, B., and Green, R.E., A ligation-based single-stranded library preparation method to analyze cell-free DNA and synthetic oligos, BMC Genomics, 2019, vol. 20, no. 1, art. ID 1023.

    Article  CAS  Google Scholar 

  52. Unifitsirovannaya regional’naya stratigracheskaya schema chetvertichnykh otlozhenii Zapadno-Sibirskoi ravniny (Unified Regional Stratigraphic Scheme of Quaternary Deposits of the West Siberian Plain), Novosibirsk: Sib. Nauchno-Issled. Inst. Geol., Geofiz. Miner. Syr’ya, 2000.

  53. van der Made, J., The rhinos from the Middle Pleistocene of Neumark–Nord (Saxony–Anhalt), Veroffentlichungen Landesamtes Denkmalpflege Archaol., 2010, vol. 62, pp. 433–500.

    Google Scholar 

  54. Vasil’ev, S.K., Lobachev, Yu.V., and Lobachev, A.Yu., New data on the locations of the Late Pleistocene megafauna on the Chumysh and Chik rivers (Altai krai and Novosibirsk oblast), in Problemy arkheologii, etnografii, antropologii Sibiri i sopredel’nykh territorii (Problems of Archeology, Ethnography, Anthropology of Siberia and Adjacent Territories), Novosibirsk: Inst. Arkheol. Etnogr., Sib. Otd., Ross. Akad. Nauk, 2014, vol. 20, pp. 15‒18.

  55. Vasil’ev, S.K., Serednev, M.A., Milyutin, K.I., Slyusarenko, I.Yu., Kozlikin, M.B., and Chekha, A.M., Collection of paleotheriological material on the Chumysh River (Altai krai) and on the Ob River near Bibikha village (Novosibirsk oblast) in 2015, in Problemy arkheologii, etnografii, antropologii Sibiri i sopredel’nykh territorii (Problems of Archeology, Ethnography, Anthropology of Siberia and Adjacent Territories), Novosibirsk: Inst. Arkheol. Etnogr., Sib. Otd., Ross. Akad. Nauk, 2015, vol. 21, pp. 36–40.

  56. Vershinina, A.O., Kapp, J.D., Baryshnikov, G.F., and Shapiro, B., The case of an arctic wild ass highlights the utility of ancient DNA for validating problematic identifications in museum collections, Mol. Ecol. Resour., 2020, vol. 20, no. 5, pp. 1182–1190.

    Article  CAS  PubMed  Google Scholar 

  57. Volkova, V.S., Paleogeography of the Karginsky Interglacial (interstage) period in Western Siberia 50 (55)–23 thousand years ago, Byull. Kom. Izuch. Chetvertichn. Perioda, Ross. Akad. Nauk, 2001, no. 64, pp. 89–93.

  58. von den Driesch A., A Guide to the Measurement of Animal Bones from Archaeological Sites, Peabody Mus. Bull., no. 1, Cambridge, Ma: Peabody Museum Press, 1978.

  59. Wacker, L., Němec, M., and Bourquin, J., A revolutionary graphitization system: fully automated, compact and simple, Nucl. Instrum. Methods Phys. Res., 2010, vol. 268, nos. 7–8, pp. 931–934.

    Article  CAS  Google Scholar 

  60. Zykin, V.S., Zykina, V.S., and Orlova, L.A., Reconstruction of environmetal and climate in Late Pleistocene in the south of Western Siberia from the sediments of the Lake Aksor basin, Arkheol., Etnogr. Antropol. Evrazii, 2003, no. 4, pp. 2–16.

Download references


We are grateful to F.K. Shidlovskiy for providing the mandible of Merck’s rhinoceros (AltR) for our study.

We are grateful to the editors of the Zoological Journal and reviewers who undoubtedly improved our manuscript.


The pollen analysis was performed within the frameworks of state contract with the Institute of Plant and Animal Ecology, Ural Branch, Russian Academy of Sciences (no. AAAA-A19-119031890086-0). The analysis of the results of 14C dating and the isotope composition was performed under State Assignment of the Institute of Geography, Russian Academy of Sciences no. 0148-2019-0006. This study was supported by the Russian Foundation for Basic Research, project no. 18-04-00982.

Author information

Authors and Affiliations


Corresponding authors

Correspondence to I. V. Kirillova, A. O. Vershinina, E. P. Zazovskaya, O. G. Zanina, S. Cutler, P. A. Kosintsev, E. G. Lapteva, O. F. Chernova or B. Shapiro.

Ethics declarations

The authors declare that they have no conflicts of interest. This article does not contain any studies involving animals or human participants performed by any of the authors.

Additional information

Translated by N. Smolina

Rights and permissions

Reprints and permissions

About this article

Check for updates. Verify currency and authenticity via CrossMark

Cite this article

Kirillova, I.V., Vershinina, A.O., Zazovskaya, E.P. et al. On Time and Environment of Stephanorhinus kirchbergensis Jäger 1839 (Mammalia, Rhinoceratidae) in Altai and Northeastern Russia. Biol Bull Russ Acad Sci 48, 1674–1687 (2021).

Download citation

  • Received:

  • Revised:

  • Accepted:

  • Published:

  • Issue Date:

  • DOI: