Fossil bear material from the oldest deposits in the Jasna Strzegowska cave (Silesia, southern Poland)

• Autorzy: Marciszak Adrian Przemysław , Lipecki Grzegorz

Identyfikatory

DOI

10.7306/gq.1556

• Języki publikacji: EN

Abstrakty

EN

Fossil bear material from the lowermost deposits of the Jasna Strzegowska cave is described and its taxonomic status assessed. Comparison with bear remains from other Early and early Middle Pleistocene localities of Eurasia shows the presence of two bear species: Ursus etruscus and U. deningeri, based on morphological and size characters of the teeth and postcranial bones. The teeth of U. deningeri from the Jasna Strzegowska cave are larger, wider and have much more complicated occlusal surfaces when compared with the teeth of U. etruscus. Both bear species are characterized by relatively short and robust metapodials, although those of U. etruscus are on average less massive. Some differences in muscle attachments and articular surfaces have also been found. Metapodials of primitive arctoid bears, that co-occurred with U. etruscus and U. deningeri, are much longer and slimmer. U. etruscus and U. deningeri probably did not live in the Jasna Strzegowska cave at the same time. Based on its time range in Eurasia and morphological characteristics, the age of U. etruscus is estimated as Early Pleistocene, between 1.8-1.3 Ma. This is its first description from Poland, and the remains described are in size and morphology indistinguishable from material from other European localities. For U. deningeri, an early Middle Pleistocene age is suggested.

Słowa kluczowe

EN

teeth; metapodials; morphotypes; evolution; lineage

• Wydawca: Państwowy Instytut Geologiczny - Państwowy Instytut Badawczy
• Czasopismo: Geological Quarterly
• Rocznik: 2020
• Tom: Vol. 64, No. 4
• Strony: 861--875
• Opis fizyczny: Bibliogr. 109 poz., rys., tab., wykr.

Twórcy

autor

Marciszak Adrian Przemysław

• University of Wrocław, Faculty of Biological Sciences, Institute of Environmental Biology, Department of Palaeozoology, Sienkiewicza 21, 50-335 Wrocław, Poland

autor

Lipecki Grzegorz

• Polish Academy of Sciences, Institute of Systematics and Evolution of Animals, Sławkowska 17, 31-016 Kraków, Poland

Bibliografia

• 1. Abbazzi, L., 2010. La fauna de Cérvidos de Barranco León y Fuente Nueva 3. In: Ocupaciones Humanas en el Pleistoceno Inferior y Medio de la Cuenca de Guadix-Baza, Memoria Científica (eds. I. Toro, B. Martínez-Navarro and J. Agustí): 273-290. Junta de Andalucía, Consejería de Cultura. E.P.G. Arqueología Monográfico.
• 2. Argant, A., 1991. Carnivores quaternaries de Bourgogne. Documents des Laboratoires de Geologie de la Faculte des Sciences de Lyon, 115: 1-301.
• 3. Argant, A., 2004. Les Carnivores du gisement Pliocène final de Saint-Vallier (Drôme, France). Geobios, 37: 133-182.
• 4. Argant, A., 2010. Carnivores (Canidae, Felidae et Ursidae) de Romain-la-Roche (Doubs, France). Revue de Paléobiologie de Genève, 29: 495-601.
• 5. Athen, K., 2007. Biometrische Untersuchungen des Stylopodiums, Zygopodiums und Metapodiums pleistozäner Ursiden im Hinblick auf die Evolution des Höhlenbären und die Klassifizierung des Fundmaterials Einhornhöhle/Harz. Ph.D. thesis, Eberhard-Karls-Universität, Tübingen.
• 6. Ballesio, R., 1986. Les carnivores du gisement pléistocène d'Oubeidiyeh (Israel). Mémoires et travaux du Centre de recherche français de Jérusalem, 5: 63-91.
• 7. Baryshnikov, G., 2006. Morphometrical variability of cheek teeth in cave bears. Scientific Annals, School of Geology, Aristotle University of Thessaloniki, 98: 81-102.
• 8. Baryshnikov, G.F., 2007. Semya medvedey (Carnivora, Ursidae) (in Russian). Nauka, Sankt-Petersburg.
• 9. Baryshnikov, G., Foronova I., 2001. Pleistocene small cave bear (Ursus rossicus) from the South Siberia, Russia. Cadernos Laboratorio Xeolóxico de Laxe, 26: 373-398.
• 10. Baryshnikov, G., Kalmykov, N., 2005. Middle Pleistocene Cave Bear (Ursus deningeri von Reichenau) from Transbaikalia (Russia). Mitteilungen der Kommission für Quartärforschung der Österreichischen Akademie der Wissenschaften, 14: 13-16.
• 11. Basilici, G., Faraone, A.G., Gentili, S., 1991. Un nuovo reperto di Macaca nelle brecce ossifere pleistoceniche di Monte Peglia (Terni, Italia centrale). Bolletino della Società Paleontologica Italiana, 30: 251-254.
• 12. Bernsen, J.J.A., 1932. Eine Revision der Fossilen Säugetierfauna aus den Tonen van Tegelen. (VI & VII) Ursus etruscus & Mustela sp. Natuurhistorisch Maandblad, 21: 20-26.
• 13. Berzi, A., 1966. L'orso di Gaville nel Valdarno Superiore. Palaeontographia Italica, 60: 19-32.
• 14. Bishop, M.J., 1982. The mammal fauna of the early Middle Pleistocene cavern infill site of Westbury-sub-Mendip, Somerset. Special Papers in Paleontology, 28: 1-107.
• 15. Bocheński, Z., Bocheński, Z.M., Tomek, T., 2012. A history of Polish birds. Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Kraków.
• 16. Boeuf, O., 1997. À propos de Chilhac, Senèze, Blassac-la-Girondie (Haute-Loire, France), gisements du Pliocène terminal, leur intéręt biochronologique. In: Actes du Congrès BiochroM'97 (eds. J.-P. Aguilar, S. Legendre and J. Michaux): 661-668. Mémoire et Travaux EPHE, Institut de Montpellier-2.
• 17. Boivin, P., Barbet, P., Boeuf, O., Devouard, B., Besson, J.-C., Hénot, J.-M., Devidal, J.-L., Constantin, C., Charles, L., 2010. Geological setting of the lower Pleistocene fossil deposits of Chilhac (Haute-Loire, France). Quaternary International, 223-224: 107-115.
• 18. Clot, A., Duranthon, F., 1990. Les mammiferes fossiles du Quaternaire dans les Pyrénées. Accord, Toulouse.
• 19. Clot, A., Chaline, J., Heintz, E., Jammot, D., Mourer-Chanviré, C., Rage, J.C., 1976. Montoussé 5 (Hautes-Pyrénées), un nouveau remplissage de fissure à faune de Vertébrés du Pléistocène inférieur. Geobios, 9: 511-514.
• 20. Dehm, R., 1962a. Altpleistozäne Säuger von Schernfeld bei Eichstätt in Bayern. Mitteilungen der Bayerischen Staatssammlung für Paläontologie und Historische Geologie, 2: 17-61.
• 21. Dehm, R., 1962b. Eine weitere altpleistozäne Spaltenfüllung bei Eichstätt in Bayern. Mitteilungen der Bayerischen Staatssammlung für Paläontologie und Historische Geologie, 2: 63-64.
• 22. Delson, E., Faure, M., Guérin, C., Aprile, L., Argant, J., Blackwell, B.A.B., Debard, E., Hancourt-Smith, W., Martin-Suarez, E., Monguillon, A., Parenti, F., Pastre, J.F., Sev, S., Skinner, A.R., Swisher, C.C., Valli, A.M.F., 2006. Franco-American renewed research at the Late Villafranchian locality of Senèze (Haute-Loire, France). Courier Forschunginstitut Senckenberg, 256: 275-290.
• 23. Dimitrijević, V., 1990. Prvi rezultati istrazivanja sisarske faune iz Trlice kod Pljevlja (in Croatian). In: Zbornik radova 12. kongresa geologa Jugoslavije, knj. 1: Stratigrafija, sedimentologija, paleontologija, 328-336. Ohrid.
• 24. Dimitrijević, V., Zic, J., Mijović, D., 2003. Pleisocene fauna on Trlica near Pljevlja (Montenegro). ProGEO News, 1: 6-7.
• 25. Erdbrink, D.P., 1953. A review of fossil and recent bears of the Old World; with remarks on their phylogeny, based upon their dentition. Jan de Lange, Deventer.
• 26. Forsyth Major, C.J., 1887. Considerazioni sulle faune dei mammiferi pliocenici e post-pliocenici della Toscana. Palaeontographica, 1: 1-82.
• 27. Forsyth Major, C.J., 1890. Note on a Pliocene mammalian fauna at Olivola in the Upper Val di Magra (Prov. Massa-Carrara), Italy. Geological Magazine, 3: 305-308.
• 28. Freudenberg, W., 1914. Die Säugetiere des älteren Quartärs von Mitteleuropa mit besonderer Berücksichtigung der Fauna von Hundsheim und Deutschaltenburg in Niederösterreich etc. Geologische und Paläontologische Abhandlungen, Neue Folge, 12: 1-219.
• 29. García, N.G., 2003. Osos y otros carnívoros de la Sierra de Atapuerca. Fundación Oso de Asturias, Oviedo.
• 30. García, N., Arsuaga, J.L., 2001. Ursus dolinensis: a new species of Early Pleistocene ursid from Trinchera Dolina, Atapuerca (Spain). Comptes Rendus de l'Académie des Sciences. Série II, Sciences de la Terre et des Planètes, 332: 717-725.
• 31. Gautier, F., Heintz, E., 1974. Le gisement villafranchien de La Puebla de Valverde (Province de Teruel, Espagne). Bulletin Muséum nationale d'Histoire naturelle 3a série, 228: 113-133.
• 32. Gibbard, P., Cohen, K.M., 2008. Global chronostratigraphical correlation table for the last 2.7 million years. Episodes, 31: 243-247.
• 33. Gibbard, P., Head, M.J., 2009a. The definition of the Quaternary system/period and the Pleistocene series/epoch. Quaternaire, 20: 125-133.
• 34. Gibbard, P., Head, M.J., 2009b. IUGS ratification of the Quaternary System/Period and the Pleistocene Series/Epoch with a base at 2.58 MA. Quaternaire, 20: 411-412.
• 35. Hilpert, B., 2006. Die Ursiden aus Hunas - Revision und Neubearbeitung der Bärenfunde aus der Steinberg-Höhlenruine bei Hunas (Gde. Pommelsbrunn, Mittelfranken, Bayern). Friedrich-Alexander-Universität Erlangen-Nürnberg, Erlangen.
• 36. ICZN, 1999. International code of zoological nomenclature. International Trust for Zoological Nomenclature, London.
• 37. Jánossy, D., 1986. Pleistocene vertebrae faunas of Hungary. Akadémiai Kiadó, Budapest.
• 38. Kahlke, R.-D., Garcia, N., Kostopoulos, D.S., Lacombat, F., Lister, A.M., Mazza, P.P.A., Spassov, N., Titov, V.V., 2011. Western Palaearctic palaeoenvironment conditions during communities, and implications for hominin dispersal in Europe. Quaternary Science Reviews, 30: 1368-1395.
• 39. Koufos, G.D., 2018. New material and revision of the Carnivora, Mammalia from the Lower Pleistocene locality Apollonia 1, Greece. Quaternary, 1: 1-38.
• 40. Koufos, G.D., Kostopoulos, D.S., 1997. New carnivore material from the Plio-Pleistocene of Macedonia (Greece) with the description of a new canid. Münchner Geowissenschaften, 34: 33-63.
• 41. Koufos, G.D., Kostopoulos, D.S., 2016. The Plio-Pleistocene large mammal record of Greece: implications for early human dispersals into European. In: Human Evolution in the Southern Balkans (eds. K. Harvati and M. Roksandic): 269-280. Springer, Dordrecht.
• 42. Koufos, G.D., Konidaris, G.E., Harvati, K., 2018. Revisiting Ursus etruscus (Carnivora, Mammalia) from the Early Pleistocene of Greece with description of new material. 2018. Quaternary International, 497: 222-239.
• 43. Kowalski, K., 1951. Jaskinie Polski, 1 (in Polish). Państwowe Muzeum Archeologiczne, Warszawa.
• 44. Kowalski, K., 1959. Katalog ssaków plejstocenu Polski (in Polish). Państwowe Wydawnictwo Naukowe, Warszawa.
• 45. Krause, J., Unger, T., Nocon, A., Malaspinas, A.-S., Kolokotronis, S.-O., Stiller, M., Soibelzon, L., Springgs, H., Dear, P.H., Briggs, A.W., Bray, S.C.E., O'Brien, S.J., Rabeder, G., Matheus, P., Cooper, A., Slatkin, M., Pääbo, S., Hofreiter, M., 2008. Mitochondrial genomes reveal an explosive radiation of extinct and extant bears near the Miocene-Pliocene boundary. BMC Evolutionary Biology, 8: 220.
• 46. Kurtén, B., 1956. The bears and hyenas of the interglacials. Quaternaria, 4: 1-13.
• 47. Kurtén, B., 1959. On the bears of the Holsteinian Interglacial. Stockholm Contributions in Geology, 2: 73-102.
• 48. Kurtén, B., 1969. Die Carnivoren-Reste aus dem Kiesen von Süßenborn bei Weimar. Paläontologische Abhandlungen, Abteilung A - Paläozoologie, 3: 735-756.
• 49. Kurtén, B., Crusafon-Pairó, M., 1977. Villafranchian Carnivores (Mammalia) from La Puebla de Valverde (Teruel,Spain). Commentationes Biologicae, 85: 1-39.
• 50. Kurtén, B., Poulianos, A.N., 1977. New stratigraphic and faunal material from Petralona Cave with special reference to the Carnivora. Anthropos, 4: 47-130.
• 51. Lewis, M., Pacher, M., Turner, A., 2010. The larger Carnivora of the West Runton Freshwater Bed. Quaternary International, 228: 116-135.
• 52. Madurell-Malapeira, J., Alba, D. M., Moyà-Solà, S., 2009. Carnivora from the late Early Pleistocene of Cal Guardiola (Terrassa, Vallès-Penedès Basin, Catalonia, Spain). Journal of Paleontology, 83: 969-974.
• 53. Madurell-Malapeira, J., Morales, J., Vinuesa, V., Boscaini, A., 2015. Úrsidos, hiénidos y félidos del Pleistoceno interior de Cueva Victoria (Cartagena, Murcia). Mastia, 11-13: 401-432.
• 54. Malez, M., 1959. A new old-Pleistocene fauna in Dalmatia. Bulletin Scientifique, 5: 13-14.
• 55. Malez, M., 1961. Staropleistocenska fauna koštane breče poluotoka Marjana kod Splita (in Croatian). Palaeontologia Jugoslavica, 4: 1-43.
• 56. Martínez-Navarro, B., 1991. Revisión sistemática y estudio cuantitativo de la fauna de macromamíferos del yacimiento de Venta Micena (Orce, Granada). Ph.D. dissertation, Universidad Autónoma de Barcelona, Barcelona.
• 57. Martinez-Navarro, B., Belmaker, M., Yosef-Bar, O., 2009. The large carnivores from Ubeidiya (early Pleistocene, Israel): biochronological and biogeographical implications. Journal of Human Evolution, 56: 514-524.
• 58. Martínez-Navarro, B., Palmqvist, P., Madurell, J., Ros-Montoya, S., Espigares, M.P., Torregrosa, V., Pérez-Claros, J.A., 2010. La fauna de grandes mamíferos de Fuente Nueva 3 y Barranco León 5. Estado de la cuestión. In: Ocupaciones Humanas en el Pleistoceno Inferiory Medio de la Cuenc de Guadix-Baza, Memoria Científica (eds. I. Toro, B. Martínez-Navarro and J. Agustí): 197-236. Arqueología Monográfico, Andalucía.
• 59. Mazza, P., Rustioni, M., 1992. Morphometric revision of the Eurasian species Ursus etruscus Cuvier. Palaeontographia Italica, 79: 101-146.
• 60. Mazza, P., Rustioni, M., 1994. On the phylogeny of Eurasian bears. Palaeontographica A, 230: 1-38.
• 61. Medin, T., Martínez-Navarro, B., Rivals, F., Madurell-Malapeira, J., Ros-Montoya, S., Espigares, M.P., Figueirido, B., Rook, L., Palmqvist, P., 2017. Late Villafranchian Ursus etruscus and other large carnivorans from the Orce sites (Guadix-Baza basin, Andalusia, southern Spain): taxonomy, biochronology, paleobiology, and ecogeographical context. Quaternary International, 431: 20-41.
• 62. Medin, T., Martínez-Navarro, B., Madurell-Malapeira, J., Figueirido, B., Kopaliani, G., Rivals, F., Kiladze, G., Palmqvist, P., Lordkipanidze, D., 2019. The bears from Dmanisi and the first dispersal of early Homo out of Africa. Scientific Reports, 9: 17752.
• 63. Mirosław-Grabowska, J., Cyrek, K., 2009. Archaeology and stratigraphy of the Jasna Strzegowska Cave. Studies of the Faculty of Earth Sciences, University of Silesia, 56: 273-282.
• 64. Moullé, P.-E., 1992. Les grands mammifères du Pleistocène inferieur de la grotte du Vallonnet (Roquebrune-Cap-Martin, Alpes-Maritimes). Etude paléontologique des Carnivores, Equidé, Suidé et Bovidés. Ph.D. dissertation, Museum National d'Histoire Naturelle à l'Institut de Paléontologie Humaine, Paris.
• 65. Musil, R., 1972. Die Bären der Stranska Skala. Anthropos, 20: 107-112.
• 66. Musil, R., 1974. Lažánky bei Tišnov - eine neue Fundstätte der Biharfauna. Acta Musei Moraviae, Scientiae naturales, 59: 87-93.
• 67. Musil, R., 1995. Large fauna of talus cones at the Stránská Skála Hill. Anthropos Neue Serie, 26: 65-83.
• 68. Musil, R., 2001. Die Ursiden-Reste aus der Unterpleistozän von Untermassfeld. Monographien des Römisch-Germanischen Zentralsmuseums, 40: 633-658.
• 69. Nadachowski, A., Stefaniak, K., Szynkiewicz, A., Marciszak, A., Socha, P., Schick, P., August, P., 2011. Biostratigraphic importance of the Early Pleistocene fauna from Żabia Cave (Poland) in Central Europe. Quaternary International, 248: 204-218.
• 70. Nadachowski, A., Lipecki, G., Ratajczak, U., Stefaniak, K., Wojtal, P., 2016. Dispersal events of the saiga antelope (Saiga tatarica) in Central Europe in response to the climatic fluctuations in MIS 2 and the early part of MIS 1. Quaternary International, 420: 357-362.
• 71. Newton, E.T., 1913. On the remains of Ursus etruscus (= U. arvernensis) from the Pliocene deposits of Tegelen sur Meuse. Verhandelingen van het Geologisch Mijnbouwkundig Genootschap voor Nederland en Koloniën, Geologische Serie, 1: 249-254.
• 72. Petrucci, M., Sardella, R., 2009. Ursus etruscus Cuvier, 1823 from the Early Pleistocene of Monte Argentario (Southern Tuscany, Central Italy). Bollettino della Societŕ Paleontologica Italiana, 48: 89-94.
• 73. Petrucci, M., Cipullo, A., Martínez-Navarro, B., Rook, L., Sardella, R., 2013. The late Villafranchian (Early Pleistocene) carnivores (Carnivora, Mammalia) from Pirro Nord (Italy). Palaeontographica Abteilung A Paläozoologie, Stratigraphie, 298: 113-145.
• 74. Prószyński, M., 1952. Geological observations of the Bug River Basin (in Polish with English summary). Biuletyn Państwowego Instytutu Geologicznego, 65: 313-340.
• 75. Przesmycki, P., 1912. Jaskinie na Wyżynie Małopolskiej (in Polish). Ziemia, 3: 443-444.
• 76. Rabeder, G., 1999. Die Evolution des Höhlenbären-Gebisses. Mitteilung der Kommission für Quartärforschung der Österreichischen Akademie der Wissenschaften, 11: 1-102.
• 77. Rabeder, G., Pacher, M., Withalm, G., 2010. Early Pleistocene bear remains from Deutsch-Altenburg (Lower Austria). Mitteilungen der Kommission für Quartärforschung der Österreichischen Akademie der Wissenschaften, 17: 1-135.
• 78. Radulescu, C., Samson, P.-M., Petculescu, A., Stiucă, E., 2003. Pliocene large mammals of Romania. Coloquios de Paleontología, 1: 549-558.
• 79. Ratajczak, U., Shpansky, A.V., Malikov, D.G., Stefaniak, K., Nadachowski, A., Wojtal, P., Ridush, B., Krakhmalnaya, T.V., Stepanchuk, V., Mackiewicz, P., 2016. Quaternary skulls of the saiga antelope from Eastern Europe and Siberia: Saiga borealis versus Saiga tatarica - One species or two? Quaternary International, 420: 329-347.
• 80. Rode, K., 1931. Über die Bären von Taubach und Ehringsdorf. Paläontologische Zeitschrift, 13: 61-72.
• 81. Rode, K., 1935. Untersuchungen uber das Gebiss der Bären. Monographien zur Geologie und Paläontologie, 2: 1-162.
• 82. Rook, L., Croitor, R., Delfino, M., Ferretti, M.P., Gallai, G., Pavia, M., 2013. The Upper Valdarno Plio-Pleistocene vertebrate record: an historical overview, with notes on palaeobiology and stratigraphic significance of some important taxa. Bollettino della Societŕ Paleontologica Italiana, 132: 140-125.
• 83. Rustioni, M., Mazza, P., 1993a. The genus Ursus in Eurasia: dispersal events and stratigraphical significance. Rivista Italiana di Paleontologia e Stratigrafia, 98: 487-494.
• 84. Rustioni, M., Mazza, P., 1993b. The Late Villafranchian bear from Pietrafitta (Perugia, Central Italy). Palaeontographia Italica, 80: 51-62.
• 85. Rybicka, M., Cyrek, K., 1997. Wyniki weryfikacyjnych badań wykopaliskowych w Jaskini Jasnej w Strzegowej, woj. Katowickie, w 1991 roku (in Polish). Łódzkie Sprawozdania Archeologiczne, 3: 5-13.
• 86. Sawicki, L., 1949. Rozwój badań w zakresie prehistorii czwartorzędu w Polsce (in Polish). Sprawozdania Państwowego Muzeum Archeologicznego, 2: 13-30.
• 87. Sawicki, L., 1953. Stan badań nad wiekiem człowieka kopalnego w Polsce (in Polish). Acta Geologica Polonica, 3: 171-186.
• 88. Schaub, S., 1943. Die ober Pliocäne Säugetierfauna von Seneze (Haute Loire) und ihre verbreitungsgeschichtliche Stellung. Eclogae Geologicae Helvetiae, 36: 270-289.
• 89. Sher, A. V., Weinstock, B.J., Baryshnikov, G.F., Davydov, S.P., Boeskorov, G.G., Zazhigin, V.S., Nikolskiy, P.A., 2011. The first record of “spelaeoid” bears in Arctic Siberia. Quaternary Science Reviews, 30: 2238-2249.
• 90. Sotnikova, M.V., 1989. Khishchnye mlekopitayushchiye pliotsenarannego pleistotsena (stratigraficheskoye znacheniye) (in Russian). Nauka, Moskva.
• 91. Stefaniak, K., 2015. Neogene and Quaternary Cervidae from Poland. Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Kraków.
• 92. Stefaniak, K., Socha, P., Tyc, A., Cyrek, K., Nadachowski, A., 2009. Caves, rock shelters and palaeontological sites in quarries of the Czestochowa Upland - catalogue of important speleological features. Studies of the Faculty of Earth Sciences, University of Silesia, 56: 307-354.
• 93. Symeonidis, N., 1992. Lower Pleistocene (Villafranchian) mammalian fossils from the basin of Sesklon (Volos). Annales Geologiques des Pays Helleniques, 35: 1-41.
• 94. Szelerewicz, M., Górny, A., 1986. Jaskinie Wyżyny Krakowsko-Wieluńskiej (in Polish). Polskie Towarzystwo Turystyczno-Krajoznawcze „Kraj”, Warszawa-Kraków.
• 95. Tchernov, E., Tsoukala, E., 1997. Middle Pleistocene (Early Toringian) carnivore remains from northern Israel. Quaternary Research, 48: 122-136.
• 96. Terzea, E., 1996. Biochronology of the Pleistocene deposits at Betfia (Bihor, Komnnii). Acta Zoologica Cracoviensia, 39: 531-540.
• 97. Torre, D., Albianelli, A., Azzaroli, A., Ficcarelli, G., Magi, M., Napoleone, G., Sagri, M., 1993. Paleomagnetic calibration of the Late Villafranchian mammalian faunas from the Upper Valdarno, Central Italy. Memorias Societa Geologica Italiana, 49: 335-344.
• 98. Torres, T.D., 1988. Osos (Mammalia, Carnivora, Ursidae) del Pleistoceno Ibérico: I. Filogenia; Distribución estratigráfica y geográfica. Estudio anatómico y métrico del cráneo. Boletín Geológico y Minero, 99: 3-46.
• 99. Torres, T.D., 1992. Los restos de oso del yacimiento de Venta Micena (Orce, Granada) y el material de Ursus etruscus C. Cuvier del Villafranquiense Europeo. In: Proyecto Orce-Cueva Victoria (1988-1992). Presencia humana en el Pleistoceno inferior de Granada y Murcia (ed. J. Gibert): 87-106. Museo de Prehistoria, Orce.
• 100. Vekua, A., 1996. Die Wirbeltierfauna des Villafranchium von Dmanisi und Ihre biostratigraphische Bedeutung. Jahrbuch des Römisch-Germanischen Zentralmuseums, 42: 77-180.
• 101. Viret, J., 1954. Le loess a bancs durcis de Saint-Vallier (Drome) etsa faune de mammiféres villafranchiens. Nouvelles Archives du Muséum d'Histoire Naturelle de Lyon, 4: 1-200.
• 102. Vislobokova, I.A., Agadjanian, A.K., 2015. New data on large mammals of the Pleistocene Trlica fauna, Montenegro, the Central Balkans. Paleontological Journal, 6: 86-102.
• 103. von Reichenau, W., 1904. Über eine neue fossile Bären-Art Ursus deningeri aus den fluviatilen Sanden von Mosbach. Jahrbücher des Nassauischen Vereins für Naturkunde, 57: 1-11.
• 104. von Reichenau, W., 1906. Beiträge zur näheren Kenntnis der Carnivoren aus den Sanden von Mauer und Mosbach. Abhandlunge der Grossherzoglich Hessischen Geologischen Landesanstalt zu Darmstadt, 4: 189-313.
• 105. Wagner, J., 2010. Pliocene to early Middle Pleistocene ursine bears in Europe: a taxonomic overview. Journal of the National Museum of Prague, Natural History Series, 179: 197-215.
• 106. Wagner, J., Čermák, S., 2012. Revision of the early Middle Pleistocene bears (Ursidae, Mammalia) of Central Europe, with special respect to possible co-occurrence of spelaeoid and arctoid lineages. Bulletin of Geosciences, 87: 461-496.
• 107. Wagner, J., Jiangzuo, Q., Lenardić, J.M., Liu, J., 2017. Taxonomic revision of bears from the locality Šandalja I (Croatia) and its biostratigraphic consequences. Fossil Imprint, 73: 533-544.
• 108. Wiszniowska, T., 1989. Middle Pleistocene Carnivora (Mammalia) from Kozi Grzbiet in the Świętokrzyskie Mts., Poland. Acta Zoologica Cracoviensia, 32: 589-630.
• 109. Wolsan, M., 1989. Drapieżne - Carnivora (in Polish). Folia Quaternaria, 59-60: 177-197.

Uwagi

Opracowanie rekordu ze środków MNiSW, umowa Nr 461252 w ramach programu "Społeczna odpowiedzialność nauki" - moduł: Popularyzacja nauki i promocja sportu (2021).