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The latest Ursus spelaeus in Italy, a new contribution to the extinction chronology of the cave bear

  • Ivan Martini (a1), Mauro Coltorti (a1), Paul P.A. Mazza (a2), Marco Rustioni (a3) and Fabio Sandrelli (a1)...

The skeleton of a young prime adult cave bear, Ursus spelaeus, was found in Chiostraccio Cave (Siena, Tuscany, central Italy), only slightly buried under rock falls. The specimen was dated yielding a conventional age of 24,030 ± 100 14C yr BP (29,200–28,550 cal yr BP), which makes it the latest known representative of the species in Italy. The skeleton was accompanied by the remains of wolf (Canis lupus), wild boar (Sus scrofa), aurochs (Bos primigenius), red deer (Cervus elaphus), roe deer (Capreolus capreolus), bat (Vespertinus murinus), and crow (Corvus monedula). The site seems confirming that the latest Italian U. spelaeus populations shared the risk of intrusion. The association of the cave bear with other animals suggests that the assemblage is an attritional palimpsest of remains of different species not originally associated in life. Cave bears were probably more vegetarian than brown bears and possibly became extinct when plant productivity dropped at the onset of MIS 2. Central and southern Italy may have offered isolated and sheltered refugia for cave bears.

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Andreucci, S., Clemmersen, L.B., Murray, A.S., and Pascucci, V. Middle to late Pleistocene coastal deposits of Alghero, northwest Sardinia (Italy): chronology and evolution. Quaternary International 222, (2010). 316.
Andrews, P., and Turner, A. Life and death of the Westbury bears. Annales Zoolgici Fennici 28, (1992). 139149.
Argenti, P., and Mazza, P.P.A. Mortality analysis of the Late Pleistocene bears from Grotta Lattaia, central Italy. Journal of Archaeological Science 33, (2006). 15521558.
Bennett, K.D., Tzedakis, P.C., and Willis, K.J. Quaternary refugia of north European trees. Journal of Biogeography 18, (1991). 103115.
Bianchi-Demicheli, F., and Oppizzi, N. Ricerche speleologiche e paleontologiche nella regione del Monte Generosa, la Caverna Generosa. Bolletino Società Ticinese di Scienze Naturali 89, (2001). 6166.
Bocherens, H., Bridault, A., Drucker, D.G., Hofreiter, M., Münzel, S.C., Stiller, M., and van der Plicht, J. The last of its kind? Radiocarbon, ancient DNA and stable isotope evidence from a late cave bear from Rochedane (France). Quaternary International (2013). (in press)
Bon, M., and Boscato, P. Associazioni faunistiche dell'Olocene Antico nell'Italia del Sud, confronto tra un sito adriatico (Grotta delle Mura, Monopoli, Bari) ed uno tirrenico (Grotta della Serratura, Marina di Camerota, Salerno). II Quaternario 9, 2 (1996). 567572.
Boscato, P. I macromammiferi dell'Aurignaziano e del Gravettiano antico di Grotta Paglicci. Palma di Cesnola, A. Paglicci. L'Aurignaziano e il Gravettiano antico: Claudio Grenzi Editore, Foggia. (2005). 4961.
Chiesa, S., Coltorti, M., Cremaschi, M., Ferraris, M., and Prosperi, L. Loess sedimentation and quaternary deposits in the Marche province. Proceedings of the International Meeting INQUA Commission on Loess and Paleogeography, the Loess in Northern and Central Italy. A Loess Basin Between the Alps and the Mediterranean Region. Quaderni di Geodinamica Alpina e Quaternaria (1990). 103130.
Clark, P.U., Dyke, A.S., Shakun, J.D., Carlson, A.E., Clark, J., Wohlfarth, B., Hostetler, S.W., and McCabe, A.M. The Last Glacial Maximum. Science 325, (2009). 710714.
Coltorti, M., and Dramis, F. Sedimentological characteristics of stratified slope waste deposits in the Umbria–Marche Apennines (central Italy) and their genetic implication. Processus et mesure de l'erosion, Ed. CNRS. (1987). 145152. (Paris)
Coltorti, M., and Dramis, F. The significance of stratified slope waste deposits in the quaternary Umbria Marche Apennines (central Italy). Zeitschrift für Geomorphologie. Neue Folge Supplementbande 71, (1988). 5970.
Coltorti, M., and Dramis, F. The chronology of Upper Pleistocene stratified slope-waste deposits in central Italy. Permafrost and Periglacial Processes 6, (1995). 235242.
Coltorti, M., Melis, E., and Patta, D. Geomorphology, stratigraphy and facies analysis of some Late Pleistocene and Holocene key deposits along the coast of Sardinia (Italy). Quaternary International (2010). 117.
Coltorti, M., Della Fazia, J., Paredes Rios, F., and Tito, G. Nuagapua (Chaco, Bolivia): evidence for the latest occurrence of megafauna in association with human remains in South America. Journal of South American Earth Sciences 33, (2012). 5667.
Coltorti, M., Firuzabadi, D., and Pieruccini, P. Geomorphological map and land units at 1:200,000 scale of the Siena Province (Southern Tuscany, Italy). Journal of Maps 2011, (2012). 115.
Dansgaard, W., Johnsen, S.J., Clausen, H.B., Dahl-Jensen, D., Gundestrup, N.S., Hammer, C.U., Hvidberg, C.S., Steffensen, J.P., Sveinbjörnsdottir, A.E., Jouzel, J., and Bond, G. Evidence for general instability of past climate from a 250-kyr ice-core record. Nature 364, (1993). 218220.
Erdbrink, D.P. A review of fossil and recent bears of the Old World with remarks on their phylogeny based upon their dentition, 2 parts. (1953). Drukkerij Jan De Lange, Deventer, The Netherlands.
Ficcarelli, G., Bertini, A., Coltorti, M., Mazza, P., Mezzabotta, C., Rook, L., and Torre, D. Hypotheses on the causes of extinction of the South American mastodons during the last maximum glacial. Journal of South American Earth Sciences 10, 1 (1997). 2938.
Ficcarelli, G., Coltorti, M., Moreno Espinosa, M., Pieruccini, P.L., Rook, L., and Torre, D. A model for the Holocene extinction of the mammal megafauna in Ecuador (South America). Journal of South American Earth Sciences 15, (2003). 835845.
Follieri, M., Giardini, M., Magri, D., and Sadori, L. Palynostratigraphy of the last glacial period in the volcanic region of central Italy. Quaternary International 47, 48 (1998). 320.
Fornaca Rinaldi, G., and Radmilli, A.M. Datazione con il metodo Th 230/238 di stalagmiti contenute in depositi musteriani. Atti Società Toscana di Scienze Naturali 75, (1968). 639646.
Forsten, A., and Dimitrijević, V. Pleistocene horses (genus Equus) in the central Balkans. Geoloski Anali Balkaanskoga Poluostrva 65, (2004). 5575.
Gandin, A., Giamello, M., Guasparri, G., Mugnaini, S., and Sabatini, G. The calcare cavernoso of the Montagnola Senese (Siena, Italy): mineralogical-petrographic and petrogenetic features. Mineralogica et Petrographica Acta 43, (2000). 271289.
Giraudi, C., and Frezzotti, M. Late Pleistocene glacial events in the Central Apennines, Italy. Quaternary Research 48, (1997). 280290.
Grandal-d'Anglade, A., and López González, F. Sexual dimorphism and ontogenetic variation in the skull of the cave bear (Ursus spelaeus Rosenmüller) of the European Upper Pleistocene. Geobios 38, 3 (2005). 325337.
Grove, A.T., and Rackham, O. The Nature of Mediterranean Europe, an Ecological History. (2001). Yale University Press, New Haven and London.
Heaton, T.J., Blackwell, P.G., and Buck, C.E. A Bayesian approach to the estimation of radiocarbon calibration curves: the IntCal09 methodology. Radiocarbon 51, (2009). 11511164.
Hewitt, G.M. Some genetic consequences of ice ages, and their role in divergence and speciation. Biological Journal of the Linnean Society 58, (1996). 247276.
Hewitt, G.M. Post-glacial re-colonization of European biota. Biological Journal of the Linnean Society 68, (1999). 87112.
Hewitt, G.M. The genetic legacy of the Quaternary ice ages. Nature 405, (2000). 907913.
Hofreiter, M., Capelli, C., Krings, M., Waits, L., Conard, N., Münzel, S., Rabeder, G., Nagel, D., Paunovic, M., Jambresic, G., Meyer, S., Weiss, G., and Pääbo, S. Ancient DNA analyses reveal high mitochondrial DNA sequence diversity and parallel morphological evolution of Late Pleistocene cave bears. Molecular and Biological Evolution 19, (2002). 12441250.
Hofreiter, M., Serre, D., Rohland, N., Rabeder, G., Nagel, D., Conard, N., Munzel, S., and Paabo, S. Lack of phylogeography in European mammals before the last glaciation. Proceedings of the National Academy of Sciences of the United States of America 101, (2004). 12,96312,968.
Huntley, B. Europe. Huntley, B., and Webb, T. Vegetation History. (1988). Kluwer Academic Publishers, London. 341384.
Huntley, B. European post-glacial forests, compositional changes in response to climatic change. Journal of Vegetation Science 1, (1990). 507518.
Huntley, B. Pollen-climate response surfaces and the study of climate change. Gray, J.M. Applications of Quaternary Research. Quaternary Proceedings 2 (1992). Quaternary Research Association, Cambridge. 9199.
Iacoviello, F., and Martini, I. Provenance and geological significance of red mud and other clastic sediments of the Mugnano cave (Montagnola Senese, Italy). International Journal of Speleology 41, 2 (2012). 317328.
Iacoviello, F., and Martini, I. Clay minerals in cave sediments and terra rossa soils in the Montagnola Senese karst massif (Italy). Geological Quarterly 53, 3 (2013). 527536.
Jaurand, E. Il glacialismo negli Appennini. Testimonianze geomorfologiche e riferimenti cronologici e paleoclimatici. Bollettino della Società Geografica Italiana 4, 3 (1999). 399432.
Johnsen, S.J., Dahl-Jensen, D., Gundestrup, N., Steffensen, J.P., Clausen, H.B., Miller, H., Masson-Delmotte, V., Sveinbjörnsdottir, A.E., and White, J. Oxygen isotope and palaeotemperature records from six Greenland ice-core stations: Camp Century, Dye-3, GRIP, GISP2, Renland and NorthGRIP. Journal of Quaternary Science 16, (2001). 299307.
Kahlke, R.-D., García, N., Kostopoulos, D.S., Lacombat, F., Lister, A.M., Mazza, P.P.A., Spassov, N., and Titov, V.V. Western Palaearctic palaeoenvironmental conditions during the Early and early Middle Pleistocene inferred from large mammal communities, and implications for hominin dispersal in Europe. Quaternary Science Reviews Special Issue, Ecological scenarios for human evolution during the Early and early Middle Pleistocene in the Western Palaearctic. Quaternary Science Reviews 30, (2011). 13681395.
Knapp, M., Rohland, N., Weinstock, J., Baryshnikov, G., Sher, A., Nagel, D., Rabeder, G., Pinhasi, R., Schmidt, H.A., and Hofreiter, M. First DNA sequences from Asian cave bear fossils reveal deep divergences and complex phylogeographic patterns. Molecular Ecology 18, (2009). 12251238.
Koby, F. Les usures séniles des canines d'Ursus spelaeus et la préhistoire. Verhandlungen. Naturforschende Gesellschaft (Basel) 51, (1939). 7695.
Kurtén, B. Life and death of the Pleistocene cave bear: a study in paleoecology. Acta Zoologica Fennica 95, (1958). 159.
Kurtén, B. The Cave Bear Story. (1976). Columbia University Press, New York.
Lowe, J.J., Rasmussen, S.O., Björck, S., Hoek, W.Z., Steffensen, J.P., Walker, M.J.C., Yu, Z., the INTIMATE group, Precise dating and correlation of events in the North Atlantic region during the Last Termination, a revised protocol recommended by the INTIMATE group. Quaternary Science Reviews 27, (2008). 617.
Lugli, S. Timing of post-depositional events in the Burano Formation of the Secchia Valley (Upper Triassic, northern Apennines) clues from gypsum–anhydrite transition and carbonate metasomatism. Sedimentary Geology 140, (2001). 107122.
Malez, M. Kvartarni sisavci (Mammalia) iz velike pećine na Ravnoj gori (SR Hrvatska, Jugoslavija). Rad Zavoda za znanstveni rad Jugoslavenske akademije znanosti i umjetnosti. Varaždin 1, (1986). 33139.
Martini, I. Cave clastic sediments and implications for speleogenesis: new insights from the Mugnano Cave (Montagnola Senese, Northern Apennines, Italy). Geomorphology 134, (2011). 452460.
Mattson, D.J. Diet and morphology of extant and recently extinct northern bears. Ursus 10, (1998). 479496.
Mazza, P., Rustioni, M., and Boscagli, G. Evolution of ursid dentition, with inferences on the functional morphology of the masticatory apparatus in the genus Ursus. Moggi-Cecchi, J. Aspects of Dental Biology: Paleontology, Anthropology and Evolution. (1995). Publisher International Institute for the study of Man, Florence. 147157.
Monegato, G., Pini, R., Ravazzi, C., Reimer, P., and Wick, L. Correlation of Alpine glaciation and global glacioeustatic changes through integrated lake and alluvial stratigraphy in N-Italy. Journal of Quaternary Science 26, (2011). 791804.
Münzel, S.C., Hofreiter, M., Stiller, M., Mittnik, A., Conard, N.J., and Bocherens, H. Pleistocene bears in the Swabian Jura (Germany): genetic replacement, ecological displacement, extinctions and survival. Quaternary International 245, (2011). 225237.
Musil, R. Ursus spelaeus — Der Höhlenbär. Museum für Ur-und Frühgeschichte Thüringens. (1980). Weimar, Germany.
Oeschger, H., Siegenthaler, V., Schotterer, U., and Gugelmann, A. A box diffusion model to study the carbon dioxide exchange in nature. Tellus 27, (1975). 168192.
Pacher, M. Upper Pleistocene cave assemblages at Alpine sites in Austria and adjacent regions. Preistoria Alpina 39, (2003). 115128.
Pacher, M., and Stuart, A.J. Extinction chronology and palaeobiology of the cave bear (Ursus spelaeus). Boreas 38, (2009). 189206.
Pascucci, V. Karst and paleoenvironments of the Siena area (central Italy). Guerrieri, L., Rischia, I., and Serva, L. 32nd International Geological Conference. (2004). Florence, Italy. 2028.
Pascucci, V., and Bianciardi, G. Geologia e morfologia, una serie di modesti rilievi caratterizzati da una struttura complessa e da un notevole sviluppo di fenomeni carsici. Manganelli, G., and Favilli, L. La Montagnola Senese, una guida naturalistica. (2001). WWF Italia, Rome. 1123.
Rabeder, G. Neues vom Höhlenbären, Zur Morphogenetik der Backenzähne. Die Höhle 34, (1983). 6785.
Rabeder, G., Nagel, D., and Pacher, M. Der Höhlenbär. (2000). Thorbecke, Stuttgart.
Ramakrishnan, U., and Hadly, E.A. Using phylochronology to reveal cryptic population histories: review and synthesis of 29 ancient DNA studies. Molecular Ecology 18, (2009). 13101330.
Randi, E. Conservation genetics of carnivores in Italy. Comptes Rendus Biologies 326, Suppl. 1 (2003). S54S60.
Ravazzi, C., Deaddis, M., De Amicis, M., Marchetti, M., Vezzoli, G., and Zanchi, A. The last 40 ka evolution of the Central Po Plain between the Adda and Serio rivers. Géomorphologie, Relief, Processus, Environnement 2, (2012). 131154.
Reimer, P.J., Baillie, M.G.L., Bard, E., Bayliss, A., Beck, J.W., Blackwell, P.G., Bronk Ramsey, C., Buck, C.E., Burr, G.S., Edwards, R.L., Friedrich, M., Grootes, P.M., Guilderson, T.P., Hajdas, I., Heaton, T.J., Hogg, A.G., Hughen, K.A., Kaiser, K.F., Kromer, B., McCormac, F.G., Manning, S.W., Reimer, R.W., Richards, D.A., Southon, J.R., Talamo, S., Turney, C.S.M., van der Plicht, J., and Weyhenmeyer, C.E. IntCal09 and Marine09 radiocarbon age calibration curves, 0–50,000 years cal BP. Radiocarbon 51, (2009). 11111150.
Reumer, J.W.F. Habitat fragmentation and the extinction of mammoths (Mammuthus primigenius, Proboscidea, Mammalia), arguments for a causal relationship. Kahlke, R.-D., Maul, L.C., and Mazza, P.P. Late Neogene and Quaternary biodiversity and evolution, regional developments and interregional correlations. Vol. I. Proceedings of the 18th International Senckenberg Conference (VI International Palaeontological Colloquium in Weimar). Courier Forschungsinstitut Senckenberg 256, (2007). 279286. (Stuttgart)
Rowe, K., Heske, E., Brown, P., and Paige, K. Surviving the ice, northern refugia and postglacial colonization. Proceedings of the National Academy of Sciences of the United States of America 101, (2004). 10,35510,359.
Starkel, L. Environmental changes at the Younger Dryas–Preboreal transition and during the early Holocene: some distinctive aspects in central Europe. The Holocene 1, (1991). 234242.
Stewart, J.R., and Lister, A.M. Cryptic northern refugia and the origins of modern biota. Trends in Ecology & Evolution 16, (2001). 608613.
Stiller, M., Baryshnikov, G., Bocherens, H., d'Anglade, G., Hilpert, B., Munzel, S.C., Pinhasi, R., Rabeder, G., Rosendahl, W., Trinkaus, E., Hofreiter, M., and Knapp, M. Withering away — 25, 000 years of genetic decline preceded cave bear extinction. Molecular and Biological Evolution 27, (2010). 975978.
Stiner, M.C. Mortality analysis of Pleistocene bears and its paleoanthropological relevance. Journal of Human Evolution 34, (1998). 303326.
Stuart, A.J. Mammalian extinctions in the late Pleistocene of northern Eurasia and North America. Biological Reviews 66, (1991). 453562.
Stuart, A.J. Late Pleistocene megafaunal extinctions: a European perspective. MacPhee, R.D.E. Extinctions in Near Time, Causes, Contexts and Consequences. (1999). Kluwer Academic/Plenum, New York. 257269.
Stuart, A.J., and Lister, A.M. Patterns of Late Quaternary megafaunal extinctions in Europe and northern Asia. Courier Forschungsinstitut Senckenberg 259, (2007). 287297.
Stuart, A.J., Kosintsev, P.A., Higham, T.F.G., and Lister, A.M. Pleistocene to Holocene extinction dynamics in giant deer and woolly mammoth. Nature 431, (2004). 684689.
Stuiver, M., Reimer, P.J., Bard, E., Beck, J.W., Burr, G.S., Hughen, K.A., Kromer, B., McCormac, G., van der Plicht, J., and Spurk, M. IntCal98 radiocarbon age calibration, 24,000–0 cal BP. Radiocarbon 40, (1998). 10411083.
Taberlet, P., and Bouvet, J. Mitochondrial DNA polymorphism, phylogeography, and conservation genetics of the brown bear (Ursus arctos) in Europe. Proceedings of the Royal Society of London. Series B 255, (1994). 195200.
Taberlet, P., Fumagalli, L., Wust-Saucy, A.G., and Cosson, J.F. Comparative phylogeography and postglacial colonization routes in Europe. Molecular Ecology 7, (1998). 453464.
Trincardi, F., and Correggiari, A. Quaternary forced regression deposits in the Adriatic basin and the record of composite sea-level cycles. Hunt, D., and Gawthorpe, R. Depositional Response to Forced Regression. Special Publication Geological Society 172, (2000). 245269. (London)
Trincardi, F., Asioli, A., Cattaneo, A., Correggiari, A., and Langone, L. Stratigraphy of the late-Quaternary deposits in the central Adriatic basin and the record of short-term climatic events. Guilizzoni, P., and Old¢eld, F.L. Palaeoenvironmental Analysis of Italian Crater Lake and Adriatic Sediments (PALICLAS). Memorie dell'Istituto Italiano di Idrobiologia 55, Pallanza, Italy (1996). 3964.
Tsoukala, E. Contribution to the study of the Pleistocene fauna of large mammals (Carnivora, Perissodactyla, Artiodactyla) from Petralona (Chalkidiki, N. Greece). Comptes Rendus. Académie des Sciences 312, II (1991). 331336.
Valdiosera, C.E., García, N., Anderung, C., Dalén, L., Crégut-Bonnoure, E., Kahlke, R.-D., Stiller, M., Brandström, M., Thomas, M.G., Arsuaga, J.L., Götherström, A., and Barnes, I. Staying out in the cold, glacial refugia and mitochondrial DNA phylogeography in ancient European brown bears. Molecular Ecology 16, (2007). 51405148.
van Andel, T. Glacial environments I: the Weichselian climate in Europe between the end of the OIS-5 and the LGM. van Andel, T., and Davies, W. Neanderthals and Modern Humans in the European Landscape During the Last Glaciation. (2003). Mc Donald Institute Monographs, Cambridge. 919.
Villa, P., and Mahieu, E. Breakage patterns of human long bones. Journal of Human Evolution 21, (1991). 2748.
Weinstock, J. Epiphyseal fusion in brown bears: a population study of grizzlies (Ursus arctos horribilis) from Montana and Wyoming. International Journal of Osteoarchaeology 19, (2009). 416423.
Wolverton, S. Natural-trap ursid mortality and the Kurtén Response. Journal of Human Evolution 50, (2006). 540551.
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