39 Current Anthropology Volume 41, Number 1, February 2000 q 2000 by The Wenner-Gren Foundation for Anthropological Research. All rights reserved 0011-3204/2000/4101-0003$3.50 The Tortoise and the Hare Small-Game Use, the Broad- Spectrum Revolution, and Paleolithic Demography 1 by Mary C. Stiner, Natalie D. Munro, and Todd A. Surovell This study illustrates the potential of small-game data for identi- fying and dating Paleolithic demographic pulses such as those as- sociated with modern human origins and the later evolution of food-producing economies. Archaeofaunal series from Israel and Italy serve as our examples. Three important implications of this study are that (1) early Middle Paleolithic populations were ex- ceptionally small and highly dispersed, (2) the first major popula- tion growth pulse in the eastern Mediterranean probably oc- curred before the end of the Middle Paleolithic, and (3) subsequent demographic pulses in the Upper and Epi-Paleolithic greatly reshaped the conditions of selection that operated on hu- man subsistence ecology, technology, and society. The findings of this study are consistent with the main premise of Flannery’s broad-spectrum-revolution hypothesis. However, ranking small prey in terms of work of capture (in the absence of special har- vesting tools) proved far more effective in this investigation of human diet breadth than have the taxonomic-diversity analyses published previously. mary c. stiner is Assistant Professor of Anthropology at the University of Arizona (Tucson, Ariz. 86731, U.S.A. [mstiner@ u.arizona.edu]). She received her Ph.D. in anthropology from the University of New Mexico in 1990. Her book Honor among Thieves: A Zooarchaeological Study of Neanderthal Ecology (Princeton: Princeton University Press, 1994) won the first Soci- ety of American Archaeology book prize in 1996. She has done archaeological fieldwork at Paleolithic sites in Italy, Israel, Tur- 1. We have many people to thank for the opportunity to conduct this research and for constructive criticisms. Steve Kuhn’s advice was instrumental for designing the simulations and for visualizing the larger implications of our study. We thank S. Heppel, Y. Werner, G. Hartman, and J. Behler for insights on the lives of tortoises, E. Tchernov, O. Bar-Yosef, J. D. Speth, R. Rabinovich, L. Horwitz,T. Dayan, G. Bar-Oz, and A. Belfer-Cohen for supplementary infor- mation on the faunas and sites of Israel, A. Segre, E. Segre-Naldini, A. Bietti, A. Recchi, and P. Cassoli for their help in accessing and documenting the collections from Italy, and H. Harpending, A. Ugan, O. Bar-Yosef, E. Bassett, J. Broughton, and anonymous re- viewers for many thoughtful comments on earlier drafts. This re- search was supported by grants from the National Science Foun- dation (to M.C.S., SBR-9511894) and the Levi Sala Care Foundation (to N.D.M.). key, and France and at sites of diverse ages in the United States. Her interests include coevolutionary processes involving hu- mans, forager economics, the forager-producer transition, popula- tion and behavioral ecology, zooarchaeology, and taphonomy. natalie d. munro is a Ph.D. candidate in anthropology at the University of Arizona. Her dissertation research focuses on the nature of Natufian subsistence at large and small sites in western Asia, testing hypotheses about the causes of economic and social changes immediately prior to the emergence of agri- culture. She received her M.A. degree from Simon Fraser Univer- sity in 1994, and she has done fieldwork in the southwestern United States, northern Europe, and western Asia. Her interests include Pleistocene and Holocene forager ecology, complex hunter-gatherers, early food production societies, zooarchaeology, and taphonomy. todd a. surovell completed his M.A. degree in anthropol- ogy at the University of Arizona in 1998 on perfecting infrared spectrographic measures of bone diagenesis in archaeological sites. His dissertation research focuses on Paleoindian resource economics and land use in the Americas. He has done fieldwork in western North America, northern Europe, and western Asia. His interests include Paleoindians, colonization of the Americas, simulation modeling, hunter-gatherer ecology and technology, and geoarchaeology. The present paper was submitted 16 x 98 and accepted 10 iii 99; the final version reached the Editor’s office 17 iii 99. Hominids have subsisted from some combination of plants and animals for more than 2.5 million years. How- ever, the nature of hominid diets, including the ways foods such as meat were obtained, the package sizes nor- mally acquired, and the manner in which nutrients were extracted and processed, has changed dramatically over this time span. Several shifts in the human predatory niche are reflected by ungulate archaeofaunal remains in particular. These include a probable transition some- time in the Plio/Pleistocene from hunting smaller prey and scavenging larger ones to an increasing emphasis on hunting, from tool-assisted extraction of consolidated bone marrow to full-scale butchering, processing, and storage of animal tissue, and, recently, from large-game hunting to animal husbandry. It is therefore not surpris- ing that most discussions of humans as predators focus on large game. There exists, however, another dimension of the ar- chaeofaunal record—small-game use—which provides unique information about the demographic conditions under which human predator-prey relations evolved. Al- though well short of dominant in most Paleolithic ar- chaeofaunal collections, small animals were important to human diets in the Mediterranean Basin from at least the early Middle Paleolithic onward. The total contri- bution of small game to Paleolithic diets and the diver- sity of species consumed did not change very much over this last 200,000 years, but the types of small game em- phasized certainly did. This is quite interesting if one considers the distinct biological properties of the small animals most commonly involved: littoral shellfish, tor- toises, partridges, rabbits, and hares. These animals differ greatly in their reproductive potentials, maturation rates, and capacities for population recovery under conditions of heavy exploitation. They also differ in the ease with