Hostname: page-component-8448b6f56d-wq2xx Total loading time: 0 Render date: 2024-04-19T14:50:51.872Z Has data issue: false hasContentIssue false

Reconstructing cave bear paleoecology from skeletons: a cross-disciplinary study of middle Pleistocene bears from Yarimburgaz Cave, Turkey

Published online by Cambridge University Press:  08 February 2016

Mary C. Stiner
Affiliation:
Department of Anthropology, University of Arizona, Tucson, Arizona 85721. E-mail: mstiner@u.arizona.edu
Hema Achyuthan
Affiliation:
Department of Geosciences, University of Arizona, Tucson, Arizona 85721
Güven Arsebük
Affiliation:
Istanbul Üniversitesi Edebiyat Fakültesi, Prehistorya Anabilim Dali, Istanbul, Turkey
F. Clark Howell
Affiliation:
Department of Anthropology, University of California, Berkeley, California 94720
Steven C. Josephson
Affiliation:
Department of Anthropology, University of Utah, Salt Lake City, Utah 84112
Kenneth E. Juell
Affiliation:
Department of Anthropology, University of Utah, Salt Lake City, Utah 84112
Jeffrey Pigati
Affiliation:
Department of Geosciences, University of Arizona, Tucson, Arizona 85721
Jay Quade
Affiliation:
Department of Geosciences, University of Arizona, Tucson, Arizona 85721

Abstract

Cave bears, an extinct subgenus (Spelearctos) of Ursus, were versatile enough to inhabit large areas of the northern hemisphere during the middle and late Pleistocene, yet they had evolved a specialized dentition that emphasized grinding functions, implying a heavy dietary reliance on tough, fibrous foods (i.e., plants). Isotope studies have yielded conflicting results on cave bear diet, however, often without consideration of the provenance of the samples or the possible contradictions that taphonomic and morphologic evidence might pose to dietary interpretations. It is likely that cave bear habits varied somewhat in response to environmental circumstance, and the limits on their abilities to do so remain unknown. If the larger goal of paleontological inquiry is to reconstruct the adaptations of cave bear species, then variation and commonalities among populations must be tracked closely, and the disparate lines of evidence currently available examined together on a case by case basis. Clearly, no single analytical technique can achieve this. By way of example we present the results of a cross-disciplinary collaboration that combines osteometric, isotopic, and taphonomic approaches to studying the paleoecology of a bear assemblage from Yarimburgaz Cave in northwest Turkey. Reference information on the linkages between diet, hibernation, and population structure in modern bears provides test implications for the investigation. Osteometric techniques demonstrate the presence of two coextant middle Pleistocene bear species in the sample–Ursus (Spelearctos) deningeri, a form of cave bear, and U. arctos or brown bear–the former abundant in the sample, the latter rare. An attritional mortality pattern for the bears and the condition of their bones show that most or all of the animals died in the cave from nonviolent causes in the context of hibernation. The study also elucidates several characteristics of the cave bear population in this region. Osteometric techniques show that the adult sex ratio of the cave bears is only slightly skewed toward females. This pattern lies near one extreme of the full range of possible outcomes in modern bear species and can only reflect a strong dietary dependence on seasonally available plants and invertebrates, showing that hibernation was a crucial overwintering strategy for both sexes; the results specifically contradict the possibility of regular, heavy emphasis on large game (hunted or scavenged) as a winter food source. The nature of wear and breakage to the adult cave bear teeth indicates that food frequently was obtained from cryptic sources, requiring digging and prying, and that extensive mastication was necessary, leading to complete obliteration of some cheek tooth crowns in old individuals. The patterns of tooth damage during life corroborate the dietary implications of the adult sex ratio and also argue for a diet rich in tough, abrasive materials such as nuts, tubers, and associated grit. The carbon and oxygen isotopic compositions of cave and brown bear tooth enamel from the site are virtually identical, and there is no evidence of a strong marine signal in either species, despite the cave's proximity to a modern estuary of the Sea of Marmara; nitrogen isotope ratios could not be examined because of poor protein preservation. The isotope results suggest that both bear species were highly omnivorous in the region during the middle Pleistocene and obtained nearly all of their food from terrestrial and fresh-water habitats. Bone pathologies, usually originating from trauma, occur in some of the adult bears, testifying to long lifespans of some individuals in this fossil population. The Yarimburgaz cave bears also exhibit great size dimorphism between the sexes, based on weight-bearing carpal bone dimensions, with adult males attaining roughly twice the body mass of adult females.

Type
Articles
Copyright
Copyright © The Paleontological Society 

Access options

Get access to the full version of this content by using one of the access options below. (Log in options will check for institutional or personal access. Content may require purchase if you do not have access.)

References

Literature Cited

Andrews, P., and Turner, A. 1992. Life and death of the Westbury bears. Annalidi Zoologica Fennici 28:139149.Google Scholar
Argant, A. 1980. Une brèche fossilifère du Pléistocène moyen de Saône-et-Loire: étude paléontologique du gisement de Château. D.E.S. Sciences naturelles, Sciences de la Terre, Université Cl Bernard, LyonI.Google Scholar
Arsebük, G., Howell, F. C., and Özbaşaran, M. 1990. Yarimburgaz 1988. XI. Kazi sonuçlari toplantisi, I. Ankara Üniversitesi Basimevi, pp. 938.Google Scholar
Arsebük, G., Howell, F. C., and Özbaşaran, M. 1991. Yarimburgaz 1989. XII. Kazi sonuçlari toplantisi, I. Ankara Üniversitesi Basimevi, pp. 1741.Google Scholar
Arsebük, G., and Özbaşaran, M. 1994. Yarimburgaz magâralari: Pleistosen'den bir kesit. XI Türk tarih kongresi. Türk Tarih Kurumu Basimevi, Ankara, 1994, pp. 1727.Google Scholar
Baryshnikov, G. 1989. Les mammifères du Paléolithique Inférieur du Caucase. L'Anthropologie (Paris) 93:813830.Google Scholar
Baryshnikov, G. 1996. Distribution of cave bears in the Pleistocene of Asia. Journal of Vertebrate Paleontology 15(Suppl. 3):17A.Google Scholar
Baryshnikov, G. 1997. Cave bears from the Paleolithic of the Greater Caucasus. In Saunders, J. J., Styles, B. W., and Baryshnikov, G., eds. Quaternary paleozoology in the Northern Hemisphere. Scientific Papers of the Illinois State Museum, Springfield(in press).Google Scholar
Behrensmeyer, A. K. 1978. Taphonomic and ecologic information from bone weathering. Paleobiology 4:150162.CrossRefGoogle Scholar
Betts, F., Blumenthal, N. C., and Posner, A. S. 1981. Bone mineralization. Journal of Crystal Growth 53:6373.CrossRefGoogle Scholar
Blackwell, B., Schwarcz, H. P., Porat, N., Howell, F. C., and Arsebük, F. 1990. Electron spin resonance (ESR) dating of Ursus teeth from Yarimburgaz Cave, Turkey. Geological Society of America Abstracts with Programs 22:A120121.Google Scholar
Bocherens, H., Fizet, M., and Mariotti, A. 1994. Diet, physiology, and ecology of fossil mammals as inferred from stable carbon and nitrogen isotope biochemistry: implications for Pleistocene bears. Palaeogeography, Palaeoclimatology, and Palaeoecology 107:213225.CrossRefGoogle Scholar
Brudevold, F., and Soremark, R. 1967. Chemistry of the mineral phase of enamel. Crystalline organization of dental mineral. Pp. 247277in Miles, A. E. D., ed. Structural and chemical organization of teeth, Vol. 2. Academic Press, London.Google Scholar
Bunnell, F. L., and Tait, D. E. N. 1981. Population dynamics of bears—implications. Pp. 7598in Fowler, C. W. and Smith, T. D., eds. Dynamics of large mammal populations. Wiley, New York.Google Scholar
Chisholm, B. S., Nelson, D. E., and Schwarcz, H. P. 1982. Stablecarbon isotope ratios as a measure of marine versus terrestrial protein in ancient diets. Science 216:11311132.CrossRefGoogle ScholarPubMed
Craighead, J. J., Varney, J. R., and Craighead, F. C. 1974. A population analysis of the Yellowstone grizzly bears. Bulletin of the Montana Forest and Conservation Experiment Station 40:120.Google Scholar
Craighead, J. J., Craighead, F. C., and Sumner, J. 1976. Reproductive cycles and rates in the grizzly bear, Ursus arctos horribilis, of the Yellowstone ecosystem. Pp. 337356in Pelton, et al. 1976.Google Scholar
Currey, J. 1984. The mechanical adaptations of bones. Princeton University Press, Princeton, N.J.CrossRefGoogle Scholar
DeNiro, M. J., and Epstein, S. 1978. Influence of diet on the distribution of carbon isotopes in animals. Geochimica et Cosmochimica Acta 42:495506.CrossRefGoogle Scholar
DeNiro, M. J., and Epstein, S. 1981. Influence of diet on the distribution of nitrogen isotopes in animals. Geochimica et Cosmochimica Acta 45:341351.CrossRefGoogle Scholar
Dong, Z. 1997. Mixture analysis and its preliminary application in archaeology. Journal of Archaeological Science 24:141161.CrossRefGoogle Scholar
Driesch, A. von den. 1976. A guide to the measurement of animal bones from archaeological sites. Peabody Museum Bulletin 1. Peabody Museum of Archaeology and Ethnology, Harvard University, Cambridge.Google Scholar
Elliot, J. C., Holcomb, D. W., and Young, R. A. 1985. Infrared determination of the degree of substitution of hydroxyl by carbonate ions in human dental enamel. Calcified Tissue International 37:372375.CrossRefGoogle Scholar
Farrand, W. R. 1992. Geoarchaeology of Yarimburgaz Cave, Turkey. Pp. 1936in Bermudez, J., Arsuaga, J., and Carbonell, E., eds. Evolución humana en Europa y los yacimientos de la Sierra de Atapuerca, Actas, Vol. 1. Junta de Castilla y León.Google Scholar
Gamble, C. 1986. The Palaeolithic settlement of Europe. Cambridge University Press, Cambridge.Google Scholar
Gargett, R. H. 1996. Cave bears and modern human origins: the spatial taphonomy of Pod Hradem Cave, Czech Republic. University Press of America, Lanham, Md.Google Scholar
Glenn, L. P., Lentfer, J. W., Faro, J. B., Miller, L. H. 1976. Reproductive biology of female brown bears (Ursus arctos), McNeil River, Alaska. Pp. 381390in Pelton, et al. 1976.Google Scholar
Gordon, K. R., and Morejohn, G. V. 1975. Sexing black bear skulls using lower canine and lower molar measurements. Journal of Wildlife Management 39:4044.CrossRefGoogle Scholar
Hilderbrand, G. V., Farley, S. D., Robbins, C. T., Hanley, T. A., Titus, K., and Servheen, C. 1996. Use of stable isotopes to determine diets of living and extinct bears. Canadian Journal of Zoology 74:20802088.CrossRefGoogle Scholar
Howell, F. C., and Arsebük, G. 1989. Report on the 1988 investigations in the Cave of Yarimburgaz (Marmara, Turkey). Unpublished manuscript submitted to theNational Geographic Society, Washington, D.C.Google Scholar
Howell, F. C., and Arsebük, G. 1990. Report on the current status of research on the Cave of Yarimburgaz (Marmara, Turkey). Unpublished manuscript submitted to theNational Geographic Society, Washington, D.C.Google Scholar
Johnson, K. G., and Pelton, M. R. 1980. Environmental relationships and the denning period of black bears in Tennessee. Journal of Mammalogy 61:653660.CrossRefGoogle Scholar
Josephson, S. C., Juell, K. E., and Rogers, A. R. 1996. Estimating sexual dimorphism by method-of-moments. American Journal of Physical Anthropology 100:191206.3.0.CO;2-0>CrossRefGoogle ScholarPubMed
Judd, S. L., Knight, R. R., and Blanchard, B. M. 1986. Denning grizzly bears in the Yellowstone National Park area. Pp. 111117in Zager, P., Garshelis, D., Graber, D., LeCount, A., and Willey, C., eds. Bears—their biology and management. Sixth International Conference on Bear Research and Management, Grand Canyon, Ariz. 1983. Port City Press., Washington D. C.Google Scholar
Koby, F. E. 1940. Les usures séniles des canines d'Ursus spelaeus et al préhistoire. Pp. 7695in Verhandlungen der Naturforschenden Gesellschaft in Basel, Vol. LI.Google Scholar
Koby, F. E. 1949. Le dimorphisme sexuel des canines d'Ursus arctos et d'Ursus spelaeus. Revue suisse de Zoologie 56:675687.Google Scholar
Koby, F. E. 1953. Modifications que les ours des cavernes ont fait subir à leur habitat. Pp. 1527in Premier Congrès International de Spéléologie, Paris, Tome IV(4).Google Scholar
Koch, P. L., Fogel, M. L., and Tuross, N. 1994. Tracing the diets of fossil animals using stable isotopes. Pp. 6392in Lathja, K. and Michiner, R. H., eds. Stable isotopes in ecology and environmental science. Blackwell Scientific, Boston.Google Scholar
Kuhn, S. L., Arsebük, G., and Howell, F. C. 1996. A Middle Pleistocene lithic assemblage from Yarimburgaz Cave, Turkey. Paléorient 22(1):3149.CrossRefGoogle Scholar
Kurtén, B. 1958. Life and death of the Pleistocene cave bear: a study in paleoecology. Acta Zoologica Fennica 95:459.Google Scholar
Kurtén, B. 1973. Transberingian relationships of Ursus arctos Linné (brown and grizzly bears). Commentationes biologicae, Societas Scientiarum Fennica, Helsinki.Google Scholar
Kurtén, B. 1976. The cave bear story. Columbia University Press, New York.Google Scholar
Kurtén, B., and Poulainos, A.-N. 1981. Fossil carnivora of Petralona Cave: status as of 1980. Anthropos 8:156.Google Scholar
Laville, H., Prat, F., and Thibault, C. 1972. Un gisement à faune du Pléistocène moyen: la Grotte de l'Eglise à Cénac-et-Saint-Julien (Dordogne). Quaternaria 16:71119.Google Scholar
Lee-Thorp, J. A., and van der Merwe, N. J. 1987. Carbon isotope analysis of fossil bone apatite. South African Journal of Science 83:712715.Google Scholar
Lee-Thorp, J. A., van der Merwe, N. J., and Brain, C. K. 1989. Isotopic evidence for dietary differences between two extinct baboon species from Swartkrans. Journal of Human Evolution 18:183190.CrossRefGoogle Scholar
Matheus, P. 1995. Diet and co-ecology of Pleistocene shortfaced bears and brown bears from eastern Beringia. Quaternary Research 44:447453.CrossRefGoogle Scholar
McNamee, T. 1984. The grizzly bear. Knopf, New York.Google Scholar
Murie, A. 1985. The grizzlies of Mount McKinley. University of Washington Press, Seattle.Google Scholar
Nelson, B. K., DeNiro, M. J., Schoeninger, M. J., and DePaolo, D. J. 1986. Effects of diagenesis on strontium, carbon, nitrogen, and oxygen concentration and isotopic composition of bone. Geochimica et Cosmochimica Acta 50:19411949.CrossRefGoogle Scholar
Özdogân, M., and Koyunlu, A. 1986. Yarimburgaz magarasi: 1986 yili Çalişmalarinin ilk sonuçlari ve bazi gözlemler. Arkeoloji ve Sanat (Istanbul) 32/33:414.Google Scholar
Pelton, M. R., Lentfer, J. W., and Edgar Folk, G. 1976 Bears—their biology and management. Third (1974) International Conference on Bear Research and Management, Binghamton, N.Y. and Moscow. ICUN new series No. 40. International Union for Conservation of Nature, Morges, Switzerland.Google Scholar
Picton, H. D., and Knight, R. R. 1986. Using climate data to predict grizzly bear litter size. Pp. 4144in Zager, P., Garshelis, D., Graber, D., LeCount, A., and Willey, C., eds. Bears—their biology and management. Sixth International Conference on Bear Research and Management, Grand Canyon, Ariz. 1983. Port City Press, Washington, D.C.Google Scholar
Pigati, J. 1996. Stable and radioisotope evidence on preservation of paleodietary and paleoenvironmental information in enamel bio-apatite. , University of Arizona, Tucson.Google Scholar
Prat, F. 1976. Les carnivores: Ursidés. Pp. 376383in de-Lumley, H., ed. La Préhistoire Française, Tome I. Les Civilisations Paléolithiques et Mésolithiques de la France. C.N.R.S., Paris.Google Scholar
Prat, F. 1988. Les Ursidés de la grotte Vaufrey. Pp. 291318in Rigaud, J.-P., ed. La Grotte Vaufrey: paléoenvironnement, chronologie, activités humaines. Mémoires de la Société Préhistorique Française, Tome XIX. C.N.R.S., Paris.Google Scholar
Prat, F., and Thibault, C. 1976. Le gisement de Nauterie à La Romieu (Gers). Pp. 182in Fouilles de 1967 à 1973. Mauterie I. Mémoires du Museum national d'Histoire naturelle, new series. Séries C, Sciences de la Terre, Tome XXXV. Paris.Google Scholar
Quade, J., Cerling, T. E., Barry, J. C., Morgan, M. E., Pilbeam, D. R., Chivas, A. R., Lee-Thorp, J. A., and van der Merwe, N. J. 1992. A 16-Ma record of paleodiet using carbon and oxygen isotopes in fossil teeth from Pakistan. Chemical Geology (Isotope Geoscience Section) 94:183192.CrossRefGoogle Scholar
Rey, C., Renugopalakrishnan, V., Shimizu, M., Collins, B., and Glimcher, M. J. 1991. A resolution-enhanced Fourier Transform infrared spectroscopic study of the environment of the CO3 ion in the mineral phase of enamel during its formation and maturation. Calcified Tissue International 49:259268.CrossRefGoogle ScholarPubMed
Reynolds, H. V., Curatolo, J. A., and Quimby, R. 1976. Denning ecology of grizzly bears in northeastern Alaska. Pp. 403409in Pelton, et al. 1976.Google Scholar
Rogers, L. L. 1987. Effects of food supply and kinship on social behavior, movements, and population growth of black bears in northeastern Minnesota. Wildlife Monographs No. 97.Google Scholar
Schoeninger, M. J., and DeNiro, M. J. 1982. Carbon isotope ratios of apatite from fossil bone cannot be used to reconstruct the diets of animals. Nature 297:577578.CrossRefGoogle ScholarPubMed
Schütt, G. 1968. Die cromerzeitlichen Bären aus der Einhornhöhle bei Scharzfeld. Mitteilungen der Geologischen Institut T. H. Hannover 7:1121.Google Scholar
Stiner, M. C. 1993. Modern human origins—faunal perspectives. Annual Review of Anthropology 22:5582.CrossRefGoogle Scholar
Stiner, M. C. 1994. Honor among thieves: a zooarchaeological study of Neandertal ecology. Princeton University Press, Princeton, N.J.Google Scholar
Stiner, M. C. 1998. Mortality analysis of Pleistocene bears and its paleoanthropological relevance. Journal of Human Evolution (in press).CrossRefGoogle Scholar
Stiner, M. C., Arsebük, G., and Howell, F. C. 1996. Cave bears and Paleolithic artifacts in Yarimburgaz Cave, Turkey: dissecting a palimpsest. Geoarchaeology 11:279327.3.0.CO;2-Z>CrossRefGoogle Scholar
Sullivan, C. H., and Krueger, H. W. 1981. Carbon isotope analysis of separate chemical phases of fossil bone. Nature 292:333335.CrossRefGoogle ScholarPubMed
Tchernov, E., and Tsoukala, E. 1997. Middle Pleistocene (early Toringian) carnivore remains from northern Israel. Quaternary Research 48:122136.CrossRefGoogle Scholar
Thackeray, J. F., van der Merwe, N. J., Lee-Thorp, J. A., Sillen, A., Lanham, J. L., Smith, R., Keyser, A., and Monteiro, P. M. S. 1990. Changes in carbon isotope ratios in the Late Permian recorded in therapsid tooth apatite. Nature 347:751753.CrossRefGoogle Scholar
Tochon-Danguy, H. J., Geoffrey, M., and Baud, C. A. 1980. Electron-spin-resonance study of the effects of carbonate substitution in the synthetic apatites and apatites from human teeth. Archives of Oral Biology 25:357361.CrossRefGoogle ScholarPubMed
Van Valkenburgh, B. 1989. Carnivore dental adaptations and diet: a study of trophic diversity with guilds. Pp. 410436in Gittleman, J. L., ed. Carnivore behavior, ecology, and evolution. Cornell University Press, Ithaca, N.Y.CrossRefGoogle Scholar
Wainwright, S. A., Briggs, W. D., Currey, J. D., and Gosline, J. M. 1976. Mechanical design in organisms. Princeton University Press, Princeton, N.J.Google Scholar