The World of - International Congress, Rome 2001

The Late of woolly (Mammuthus primigenius), straight-tusked ( antiquus) and other in

A.J. Stuart, A.M. Lister Department of Biology, University College, London, UK [email protected]

We are engaged in a research project (funded at present, it is apparent that these range changes by the Natural Environment Research Council - were not the same for each ; for example NERC) on megafaunal throughout the “last stands” of Mammuthus primigenius, Europe within the period ca. 50,000 to 9000 14C Megaloceros giganteus and Palaeoloxodon BP. The work involves a survey of strati- antiquus appear to have been made in very dif- graphic information and available 14C dates, and ferent regions of Europe. Tracking these changes also sampling crucial material for a major involves firstly gathering data from the literature programme of AMS 14C dating. Both of the and from colleagues in each region. By these elephant species present in the European Late means we are building up an approximate pic- : Mammuthus primigenius and ture and specifying the likely latest material of Palaeoloxodon antiquus are included in the our target species for each region. In to project. obtain a much more accurate database, we are Our target species include most of those that sampling the putatively latest material and sub- became extinct, or regionally extinct, after mitting it for 14C dating. ca. 15,000 BP: Mammuthus Late Quaternary extinctions have been vari- primigenius, woolly Coelodonta ously attributed to overkill by hunters antiquitatis; giant Megaloceros giganteus; (Martin 1984; Martin & Steadman 1999), to lion leo; and spotted hyaena Crocuta environmental changes (Graham & Lundelius crocuta. We are also targeting species which 1984; Graham 1990; Guthrie 1984, 1990; retreated to the south of Europe (Iberia, , Lister & Sher 1995) and more recently to ) prior to extinction, probably before “hyperdisease” (MacPhee & Marx 1997). 20,000-30,000 BP: straight-tusked elephant Stuart (1991, 1999) has proposed that the prox- Palaeoloxodon antiquus; narrow-nosed rhinoc- imate cause was by , but only eros hemitoechus (perhaps also when megafaunal populations were already Merck’s rhinoceros Stephanorhinus kirchber- stressed and reduced in numbers and geograph- gensis); and Hippopotamus ical range by major climatic changes. Our cur- amphibius. rent project aims to test these rival hypotheses of samples of elephants in Europe by seeking correlations between 14C and other European megafauna is currently in dated extinction patterns and proxy data for cli- progress, and we are actively seeking more matic/environmental changes on the one hand, material to be dated. and the appearance of modern humans and The strategy is to follow in some detail the upper palaeolithic artefacts on the other. patterns of shrinking geographical distributions Woolly mammoth Mammuthus primigenius, that preceded extinction. From the data available widespread over most of Europe in the Last

722 The Late of woolly mammoth (Mammuthus primigenius)...

Cold Stage, became extinct during the Late L.W. Nelson (eds.), Megafauna and Man. Glacial (Stuart 1991). There are a number of Flagstaff: North Arizona Press. records from north-western and central Europe Lister, A.M. & Sher, A.V. 1995. Ice cores and as late as 13,000 to 12,000 BP, but none so far mammoth extinction. 378: 23-24. after this time. Woolly mammoth may have MacPhee, R.D.E. & Marx, P.A. 1997. The retreated first from , and it is 40,000 plague; humans, hyperdisease anticipated that forthcoming dates will clarify and first contact extinctions. In S.M. this picture. Our research on mammoth extinc- Goodman & B.D. Patterson (eds.), Natural tion in Europe links with ongoing work in Change and Human Impact in Mada- and the Russian Far East (Orlova et al. gascar: 169-217. Washington D.C.: Smi- 2000; Sher 1997). thsonian Institution Press. In marked contrast, Palaeoloxodon antiquus Martin, P.S. 1984. Prehistoric overkill: a global was widespread in north-western, central and model. In P.S. Martin & R. G. Klein (eds.), southern Europe ca. 120,000 BP (“Last Quaternary Extinctions, a Prehistoric ”, Oxygen Isotope Stage 5e) (Stuart Revolution. Tucson: 1991). Prior to extinction, we hypothesise that Press. it retreated to southern Europe (Iberia, Italy, Martin, P.S. & Steadman, D.W. 1999. and probably the Balkans) in the early part of Prehistoric extinctions on islands and conti- the Last Cold Stage (OIS 4 and possibly OIS nents. In R.D.E. MacPhee (ed.), Extinctions 3), presumably in response to climatic cooling in Near Time; Causes, Contexts and Con- and a similar contraction in range of temperate sequences. New York: Kluwer Acade- vegetation. It may prove difficult to determine mic/Plenum Publishers. the timing of its extinction with any accuracy, Orlova, L.A., Kuzmin, Y.V., & Zolnikov, I.D. as Palaeoloxodon material of this age is very 2000. Time-space systematics for mam- rare, and much or all of it close to or beyond the moth (Mammuthus primigenius Blum.) and range of 14C dating. prehistoric humans in Siberia (on the basis of radiocarbon dating). , REFERENCES Ethnology, and Anthropology of 1 (3): 31-41. Graham, R.W. 1990. Evolution of new ecosys- Sher, A.V. 1997. Late-Quaternary extinction of tems at the end of the Pleistocene. In L.E. large in northern Eurasia: A new Agenbroad, J.I. Mead & L.W. Nelson look at the Siberian contribution. In B. (eds.), Megafauna and Man. Flagstaff: Huntley, W. Cramer., A.V. Morgan, H.C. North Arizona Press. Prentice & J.R. M. Allen. (eds.), Past and Graham, R.W. & Lundelius, E.L. 1984. Co- future rapid environmental changes: The evolutionary disequilibrium and Plei- spatial and evolutionary responses of ter- stocene extinctions. In P.S Martin & R. G. restrial biota: 319-339. Berlin-Heidelberg- Klein (eds.), Quaternary Extinctions, a New York: Springer-Verlag. Prehistoric Revolution. Tucson: University Stuart, A.J. 1991. Mammalian extinctions in of Arizona Press. the of northern Eurasia and Guthrie, D.E. 1984. Mosaics, allochemicals . Biological Reviews 66: and nutrients. In P.S Martin & R. G. Klein 453-562. (eds.), Quaternary Extinctions, a Prehi- Stuart, A.J. 1999. Late Pleistocene megafaunal storic Revolution. Tucson: University of extinctions; a European perspective. In: Arizona Press. R.D.E. MacPhee (ed.), Extinctions in Near Guthrie, D.E. 1990. Late Pleistocene faunal Time; Causes, Contexts and Consequences. revolution: a new perspective on the extinc- New York: Kluwer Academic/Plenum tion debate. In L.E. Agenbroad, J.I. Mead & Publishers.