Nina G. Jablonski The influence of life history and diet on Department of Anthropology, the distribution of catarrhine primates California Academy of during the Pleistocene in eastern Asia Sciences, Golden Gate Park, San Francisco, CA 94118-4599, U.S.A. E-mail: Environmental changes during the Pleistocene in eastern Asia had [email protected] profound impacts on the distributions of mammalian groups. Critical for many mammals were the southward latitudinal shifts of the tropical and subtropical vegetational zones, and decreases in the areas Matthew J. Whitfort of these zones. Examination of the responses of members of a single Department of Anatomy clade, the Catarrhini, indicates that the main catarrhine genera of and Human Biology, eastern Asia responded individually to the environmental changes in The University of Western the Pleistocene. These responses were influenced by the life history Australia, Nedlands, parameters and diets of the genera involved. Those animals WA 6907, Australia. E-mail: (macaques, langurs) with shorter gestation times, shorter weaning [email protected] periods, shorter interbirth intervals, higher intrinsic rates of increase of population, and abilities to survive on a wider variety of vegetation Nola Roberts-Smith in seasonal habitats were less adversely affected than those (gibbons, ESRI Australia Pty. Ltd., orang-utans and the giant extinct hominoid, Gigantopithecus) with 345 Harborne St., Herdsman, more protracted reproductive schedules, lower intrinsic rates of WA 6017, Australia. E-mail: population increase and preferences for the higher quality foods [email protected] (especially ripe fruits) of less seasonal environments. Hominids, while displaying ‘‘hyper-ape’’ life history parameters, increasingly overcame the constraints of these parameters through extrasomatic means not Xu Qinqi available to other catarrhines. This ability made possible their colon- Institute of Vertebrate ization, by the Late Pleistocene, of highly seasonal habitats such as Paleontology and tundra, which were off-limits to non-culture-bearing catarrhines. Paleoanthropology, Chinese 2000 Academic Press Academy of Sciences, P.O. Box 643, Beijing, China 100044. E-mail: [email protected] Received 1 June 1999 Revision received 23 September 1999 and accepted 3 February 2000 Keywords: Homioidea, Cercopithecoidea, seasonality, life history parameters, reproduction, Journal of Human Evolution (2000) 39, 131–157 Quaternary environmental doi:10.1006/jhev.2000.0405 change, biogeography. Available online at http://www.idealibrary.com on Introduction The sometimes extreme climatic fluctu- ations of the Pleistocene destabilized species Natural populations of mammals respond interactions within communities, with the to the myriad stresses of climatic change result that species tended to respond indi- by latitudinal shifts in abundances and/or vidually to environmental stresses because geographic range boundaries (FAUNMAP they observed the environment with their Working Group, 1996; Roy et al., 1996). own unique suites of life history parameters. 0047–2484/00/080131+27$35.00/0 2000 Academic Press 132 . ET AL. Environmental changes during the Pleis- Turner, 1992; Reed, 1997) have observed tocene in East Asia were more marked than that different genera and species of homi- in other parts of the Northern Hemisphere nids tend to be associated with distinct because the local climatic effects of the assemblages of mammals. The nature of Himalayas and Qinghai-Xizang (Tibetan) these assemblages changed through time, Plateau tended to magnify the orbitally as environmental conditions changed, as induced climatic fluctuations associated migrations brought hominids into contact with glacials and interglacials worldwide with new species, and as shifts in the trophic (Liu et al., 1997). Among the most dramatic levels of hominid populations occurred. of these changes was the abrupt increase, Here we attempt to examine, through the at about the Gauss/Matuyama boundary course of the Pleistocene, the nature of the (2·5 Ma), in aeolian dust deposition occur- mammalian communities that included ring on the Loess Plateau due to an apparent hominids in east Asia to determine if the northward shift and intensification of the species that hominids were associated with Siberian High (Ding et al., 1997). Marked changed through time. increases in environmental seasonality at all A geographic information system (GIS) latitudes, increasing environmental hetero- was used to assemble, visually superimpose, geneity and fragmentation, an increasing and analyze data concerning past environ- potential for physical isolation of popu- ments and animal distributions. Paleo- lations as a result of habitat fragmentation, environmental data were compiled from the and changes in the configuration of bio- literature as described below and developed geographic corridors were the most im- into composite coverages representing the portant consequences of these changes for environmental situations of the Early, mammalian populations (Ferguson, 1993; Middle and Late Pleistocene, respectively. Jablonski, 1993). On these coverages were plotted data on Our primary goal in this study is to exam- catarrhine-bearing paleontological localities ine the geographical responses of the genera for the Pleistocene. Our geographical scope within a closely related mammalian clade in was limited to China because paleoenviron- order to shed light on the factors critical to mental information and paleontological determining taxon survival and movement occurrence data for the country were plenti- during times of environmental stress. The ful and better sampled than for other East Catarrhini or Old World anthropoids were Asian countries. chosen as the clade of interest because the phylogeny of the group is widely agreed Methods upon and the biology of most of the con- stituent genera is well known. In their land- Paleoenvironmental coverages and data on mark paper, Temerin & Cant (1983) called fossil mammal distributions for empirical evaluation of their hypotheses Paleoecological data were compiled from concerning the evolutionary divergence of the literature and developed into composite Old World monkeys and apes. This study paleoenvironmental maps or coverages rep- represents an attempt to answer this call resenting the environmental zones of the with a combination of paleontological and Early, Middle and Late Pleistocene, respect- neontological data. ively. All coverages were developed using Our secondary goal is to determine if Arc/InfoGIS software. The sources from changes may have occurred over time in the which paleoenvironmental data were pattern of associations between hominids obtained were as follows: (1) soils, loess, and and other mammals. Other workers (e.g., paleoshorelines: An et al., 1991; Liu & Ding, 133 1984; (2) vegetation (mostly pollen), mol- entered. In most of the original fossil or lusks, and lake levels: Liu & Ding, 1984; geological descriptions consulted, only Winkler & Wang, 1993; and (3) permafrost: approximate ages (e.g., ‘‘Middle Pleisto- Cui & Song, 1991; Zhou et al., 1991. The cene’’) were provided for the fossils, based boundary dates adopted for subdivision of on faunal correlation. Most east Asian Ter- the Pleistocene were: Early Pleistocene: tiary and Quaternary fossil sites have not 2·5Ma–780 ka; Middle Pleistocene: 780– been dated to a high level of accuracy due, in 128 ka; and Late Pleistocene: 128–11 ka. large part, to a paucity of sediments of The recognition of the Gauss/Matuyama volcanic origin associated with the fossils. boundary as the Plio-Pleistocene boundary The use of electron spin resonance and is followed by most geologists in China uranium series dating on sediments and because it corresponds to the striking con- speleothems from some cave sites (e.g., trast between the Red Clay, formed under Zhoukoudian) and the implementation of conditions of more or less continuous paleomagnetic stratigraphy on sequences warmth, and the loess (Ding et al., 1997). of continuous sedimentation (e.g., the Loess Although the 2·5 Ma age for the Pliocene– Plateau) have made possible the refinement Pleistocene boundary is not widely accepted of dates in recent years. The application of as a standard outside of China, it was used these techniques is still not widespread here because the Chinese concept of the enough, however, to have made possible ‘‘Early Pleistocene’’ was that used as an broad-scale correlations by ‘‘absolute’’ ages. approximate age assignment for fossil finds The accuracy of taxonomic assignments and reported in the primary Chinese literature age assessments was carefully checked for all sources consulted in the development of the database entries, and amendments were fossil mammal database. It is recognized made in these areas as necessary. This work that the paleoenvironmental coverages consisted mostly of consulting published developed for this study represent average updates of age and taxonomic determi- situations or generalizations over relatively nations for species from various localities, long periods of time and that they cannot based on detailed geological and systematic convey details of the major, and sometimes investigations published after the original abrupt, climatic changes which occurred report. One record in the database consists during the Pliestocene epoch (Roy et al., of the occurrence, at one specific time and 1996). place, of one species, and is thus best The distributions of the families and gen- referred