CAVEBEAR ECOLOGYAND INTERACTIONSWITH PLEISTOCENE HUMANS MARYC. STINER, Department of Anthropology,Building 30, Universityof Arizona,Tucson, AZ 85721, USA,email: [email protected] Abstract:Human ancestors (Homo spp.), cave bears(Ursus deningeri, U. spelaeus), andbrown bears (U. arctos) have coexisted in Eurasiafor at least one million years, andbear remains and Paleolithic artifacts frequently are found in the same caves. The prevalenceof cave bearbones in some sites is especiallystriking, as thesebears were exceptionallylarge relative to archaichumans. Do artifact-bearassociations in cave depositsindicate predation on cave bearsby earlyhuman hunters, or do they testify simply to earlyhumans' and cave bears'common interest in naturalshelters, occupied on different schedules?Answering these and other questions aboutthe circumstancesof human-cave bear associationsis made possible in partby expectations developedfrom research on modem bearecology, time-scaledfor paleontologicand archaeologic applications. Here I review availableknowledge on Paleolithichuman-bear relations with a special focus on cave bears(Middle Pleistocene U. deningeri)from YarimburgazCave, Turkey.Multiple lines of evidence show thatcave bearand human use of caves were temporallyindependent events; the apparentspatial associations between human artifacts andcave bearbones areexplained principally by slow sedimentationrates relative to the pace of biogenicaccumulation and bears' bed preparationhabits. Hibernation-linkedbehaviors and population characteristics of cave bears,based on osteometric,isotopic, and age andsex structureanalyses, indicate that they dependedheavily on seasonalfood supplies,which were rich in resistantplant materials and cryptic, gritty foods. Thereis little evidence of direct ecological interactionamong Pleistocene humans and cave bears. Ursus 11:41-58 Key words: brownbears, cave bears,Mediterranean, mortality, paleodiet, Pleistocene human-bear interactions, sex ratio,Ursus arctos, Ursusdeningeri, Ursus spelaeus Pleistocene sediments in Eurasiancaves often contain By way of anthropologicalbackground, the chronol- complex records of habitationby predatoryspecies. Of ogy of Paleolithic culturesbegins aroundthe Plio-Pleis- these, early humans (Homo spp.), wolves (Canis lupus), tocene boundary and lasts until the Holocene. The foxes (Vulpes spp.), spotted hyenas (Crocuta crocuta), Paleolithic traditionallyis divided into 3 major cultural and bears (Ursus spp.) were especially prolific sources phases: the Lower,the Middle, andthe UpperPaleolithic. of bone refuse. Making sense of cave faunas is an inter- The hominid forms associatedwith these culturesvaried esting, but also challenging, enterprise,beginning with considerably,and there is no simple correspondencebe- the questions of how the assemblagesformed, what spe- tween cultural (behavioral) change and hominid mor- cies were active collectors and modifiers of bones, and phological (skeletal) change. Lower Paleolithicartifacts the extent to which sediment chemistryfavored skeletal are attributedto some of the late Australopithecinespe- preservation. These problemsfall in the methodological cies, which were confined to the African continent, as domainof vertebratetaphonomy, an areaof researchcon- well as to early variantsof the genus Homo, which by cerned with how bone assemblages become part of the 1.4-1.8 million years ago (MYA) spreadfrom Africa into paleontological and archaeologicalrecords. much of Eurasia. First appearingsome 250,000 years While clear answers about assemblage formationhis- ago, Middle Paleolithic artifactsare characterizedby in- tory seldom are easy to obtain, thereis a wealth of infor- novations in stone tool productiontechniques and arti- mation locked in cave sites about the ecology of early fact forms. They are attributedto archaichumans such humans and their relations with potential competitors, as Neandertalsin general (H. sapiens neanderthalensis) mainly of the order Carnivora. The story of human- as well as to earliest anatomicallymoder humans (H. bear interactionsis both importantand enigmatic. Con- sapiens sapiens) in the west Asian cave sites of Qafzeh verging patternsof omnivorywould seem reasonenough and Skhul (Bar-Yosef et al. 1986, Valladaset al. 1988, for Pleistocene humans and bears to have stayed out of Bar-Yosef 1989, Vandermeersch1989). The Upper Pa- each other'sway. Yet the remainsof cave bearsor brown leolithic, which began between 42,000 and 35,000 years bears are found in nearlyevery EurasianMiddle and Up- ago (dependingon region), is markedby spectacularra- per Paleolithic site with preserved bone-sometimes diations in materialculture over a relatively short time many bear bones, but more usually a few. In fact, bear span. Artifacts of all 3 periods commonly are found in remains turn up in Paleolithic cave sites more consis- open sites. Cave sites, usually in limestone solutioncavi- tently than the remains of almost any other large carni- ties, were periodically inhabitedduring the Middle and vore, at least prior to 20,000 years ago, when greater Upper Paleolithic culture periods only, and most of the interspecificexclusion among cave residentsis apparent archaeofaunalrecords of these periods come from caves (Gamble 1986, Stiner 1994). for the simple reasonthat cave sedimentsfavor bone pres- ervation. Paleolithic peoples probably were no more 42 Ursus 11:1999 bound to caves than bears, but both were at times at- Middle Pleistocene (U. etruscus) was relatively small- tractedby the prospectof easy shelter. bodied but evolved into largertypes by the Middle Pleis- Here I review availableknowledge on Pleistocene hu- tocene. This presentationconcerns 2 bears in particular, man-bear relations,as gleaned by archaeologicalinves- cave bears (U. deningeri, U. spelaeus, U. rossicus) and tigations of cave sites, with a special focus on cave bears brown bears (U. arctos and its most immediate ances- in southernEurope and western Asia. The rich litera- tors) (Kurten1976, Baryshnikov1998). Once prevalent ture on moder brown and black bears (U. americanus) throughoutEurasia, all cave bearswere extinctby roughly plays a key role in this kind of research. Specifically, 10,000 years ago (Baryshnikov1999), and most popula- wildlife data are used to build testable predictionsabout tions disappearedconsiderably earlier. Brownbears con- how bears may contributeto faunal assemblage forma- tinue to exist and even doubled their geographic range tion. The faunal patternsinside caves that can be ex- duringthe Late Pleistocene by colonizing the Americas, plained by modern bear behavior are played against as did humans not long thereafter. anthropologicalhypotheses about ancienthuman behav- Because recent humans and bears display strong at- ior. A variety of analytic techniques are marshaledfor tractions to meat as well as to energy-richplant foods, this kind of research,including osteometry, skeletal dam- theirevolutionary histories following biogeographiccon- age analysis, taxonomic and body part profiling, and tact must have affected one anotherto some extent. The mortalityand isotope analyses, to obtain rigorous,if in- ecological links between humansand bears may always direct,evidence aboutPleistocene cave bear(and human) have been relativelyweak, because both tend to be versa- ecology. For relatedpublications see Stiner(1994, 1998) tile, generalistforagers (sensu Foley 1984). These links and Stiner et al. (1996, 1998), and for information on were perennial, however, due to overlappingneeds for Pleistocene carnivore guilds in the MediterraneanBa- foraging territoryand, periodically,for shelter. sin, see Stiner (1990, 1991, 1992, 1993, 1994) and cita- tions therein. PLEISTOCENEHUMAN-BEAR INTERACTIONS:FACT AND FICTION HISTORYOF HOMINID-URSID There is no shortageof bear stories in anthropological COEXISTENCE andpopular literature. Most of them concern"cave men" Between 1.8 and 1.4 million years ago, populationsof (Neandertals)and cave bears. The original site for the a hominid known as Homo ergaster expandedfrom Af- Neandertalcave bear cult is DrachenlochCave in Swit- rica into Eurasia. Hominids' spreadacross southernAsia zerland, made famous by Bachler's radical interpreta- appearsto have been rapid (Klein 1989). Colonization tion of a faunaconsisting almost exclusively of bearbones of continental Europe and the colder regions of north- (Kurten 1976). The sediments of Drachenloch lacked central and eastern Asia by hominids took considerably artifactualmaterial, but allegedly preserveda stone crypt longer, but they certainly reached these regions before with as many as 7 cave bear skulls arrangedneatly in- 500,000 years ago. The first hominids to enter Eurasia side and, nearby,a smaller stone chest packed with bear were omnivores with a notable capacity for huntingver- long bones. Bachler's claim was refutedby Koby (1940) tebrateprey, a propertywhich may have been essential and in Kurten's (1976) highly entertainingbook, The to winter survival in northernhabitats (Foley In Press, Cave Bear Story (also Kurten 1958, 1971, 1973). Stiner In Press). Colonization of new continents put The cave bear cult is a engaging story, if only for the hominidsin contactwith diverse environmentsand novel early spiritualismit implies. The idea that archaic hu- floras and faunas. mans manipulatedbear remains in symbolic ways per- Hominids were late arrivalsto the predatoryguilds of sists, at least partlybecause bear bones frequentlyoccur Africa, where they first evolved, and hominid-camivore