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Appendicular Robusticity and the Paleobiology of Modern Human Emergence (Paleoanthropology͞human Origins͞late Pleistocene͞postcrania)

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Appendicular Robusticity and the Paleobiology of Modern Human Emergence (Paleoanthropology͞human Origins͞late Pleistocene͞postcrania) Proc. Natl. Acad. Sci. USA Vol. 94, pp. 13367–13373, November 1997 Anthropology This contribution is part of the special series of Inaugural Articles by members of the National Academy of Sciences elected on April 30, 1996. Appendicular robusticity and the paleobiology of modern human emergence (Paleoanthropologyyhuman originsyLate Pleistoceneypostcrania) ERIK TRINKAUS* Department of Anthropology, Campus Box 1114, Washington University, St. Louis, MO 63141, and Unite´ de Recherche Associe´e 376 du Centre National de la Recherche Scientifique, Laboratoire d’Anthropologie, Universite´ de Bordeaux I, 33405 Talence, France ABSTRACT The emergence of modern humans in the ern Old World. However, sufficient remains are now known Late Pleistocene, whatever its phylogenetic history, was char- from less well represented areas to indicate that, once normal acterized by a series of behaviorally important shifts reflected stochastic and ecogeographical patterns of interregional vari- in aspects of human hard tissue biology and the archeological ation within species of large-bodied mobile terrestrial mam- record. To elucidate these shifts further, diaphyseal cross- mals are taken into account, the northwestern Old World is sectional morphology was analyzed by using cross-sectional generally representative of the more global patterns of human areas and second moments of area of the mid-distal humerus biology. and midshaft femur. The humeral diaphysis indicates a grad- With this information in mind, we have been investigating ual reduction in habitual load levels from Eurasian late patterns of Late Pleistocene hominid diaphyseal appendicular archaic, to Early Upper Paleolithic early modern, to Middle robusticity by using cross-sectional geometry (1–5). Given the Upper Paleolithic early modern hominids, with the Levantine high degree of plasticity of the mammalian diaphyseal cortical Middle Paleolithic early modern humans being a gracile bone, especially during development (2, 6, 7), this approach anomalous outlier. The femoral diaphysis, once variation in provides a paleobiological window on the habitual activity ecogeographically patterned body proportions is taken into levels of extinct hominid populations. Moreover, potentially account, indicates no changes across the pre-30,000 years B.P. contrasting patterns of upper vs. lower limb diaphyseal cortical samples in habitual locomotor load levels, followed by a hypertrophy allow insights into manipulative vs. locomotor modest decrease through the Middle Upper Paleolithic. activity levels, thus shedding light on two of the most important aspects of hominid behavioral evolution. The middle of the Late Pleistocene, between approximately 100,000 and 30,000 years B.P., saw the emergence and estab- A Genealogical Digression lishment of a novel constellation of human biological charac- teristics. This evolutionary process, known as the ‘‘origins of At the same time, the majority of the research on Late modern humans,’’ led to the presence across the Old World by Pleistocene hominid evolution beyond philatelic concerns has ca. 30 thousands of years (kyr) B.P. of a new biological complex been focused on the phylogenetic relationships of geographical to the exclusion of the one which, with evolutionary modifi- groups of late archaic and early modern humans. And despite cation, had been present throughout archaic Homo for the a century of debate on this issue with the progressive intro- previous 1.71 million years. With minor evolutionary changes duction of more diverse and higher-quality data and analytical through time and space, this biological pattern remains in place techniques, combined with more global approaches to the in the present world. problem, there is little consensus. Indeed, the current and Our comprehension of the evolutionary emergence of mod- ongoing debate on the phylogenetic aspects of modern human ern humans rests primarily on our ability to decipher from the emergence appears to be more concerned with hypothesis paleoanthropological record the patterns and processes of confirmation than with hypothesis testing. change, whether adaptive or stochastic, regional or global, The past decade has seen the increasing application of which enabled one biological pattern to replace a previously human molecular data to issues of modern human origins. highly successful one in a relatively short period of geological However, with one exception (8), the molecular data that have time. This paleobiological and paleoanthropological problem, been brought to bear on the issue have no empirical time although dependent on neontological uniformitarian patterns depth, only probabilistic inferential time depth dependent on for explanatory reference, can be resolved only through the both the nature of the data and the layered analytical assump- analysis of the prehistoric record, both paleontological and tions behind the various quantitative techniques used to archeological. process those data. Moreover, all of these analyses assume a With this problem in mind, paleoanthropological research highly uniform stochastic accumulation of genetic change (i.e., has included attempts to decipher the patterns and degrees of a molecular clock that keeps accurate time throughout the last y change of functionally relevant aspects of human biology half-million years) and or geographically uniform human de- during the Late Pleistocene. Given the nature of the hominid mographic stability throughout the Middle and Late Pleisto- fossil record with its abundance of fossils from Europe and cene. These assumptions are simply untenable. Any reasonable western Asia and the dearth of reasonably complete remains assessment of molecular data, analytical techniques, and pro- from elsewhere in the Old World, this research has been focused on the paleoanthropological record of the northwest- Abbreviations: kyr, thousands of years; BIB, bi-iliac breadth; FL, femoral length. *To whom reprint requests should be addressed at: Department of © 1997 by The National Academy of Sciences 0027-8424y97y9413367-7$2.00y0 Anthropology, Campus Box 1114, Washington University, St. Louis, PNAS is available online at http:yywww.pnas.org. MO 63130. e-mail: [email protected]. 13367 13368 Anthropology: Trinkaus Proc. Natl. Acad. Sci. USA 94 (1997) cesses makes it highly unlikely that the standard errors of bi-planar radiography for the parallax-adjusted determination estimates of divergence times are sufficiently small to be useful. of cortical thicknesses, from which the endosteal contours The geographical and demographic fluctuations of Pleistocene were interpolated. Cross-sectional parameters (total and cor- hominid populations, given both their foraging adaptive pat- tical area, anatomically oriented and maximum-minimum sec- terns and susceptibility to major Pleistocene climatic fluctua- ond moments of area, and the polar moment of area) were tions, make any assumptions of uniform population size and computed from digitized cross sections by using a PC version distribution implausible, even for short periods of the last of SLICE (12, 13). In this framework, cortical area represents half-million years. structural resistance to axial loading, second moments of area The fossil data as it pertains to strictly phylogenetic issues indicate resistance to bending in the plane in question, and are not much better except in Atlantic Europe, a peripheral polar moments of area approximate strength relative to tor- cul-de-sac where the transition was very late, relatively abrupt, sional forces. Furthermore, because the polar moment of area and probably unrepresentative of more global patterns. Else- is the sum of any two perpendicular second moments of area, where across the Old World the human paleontological evi- it also provides an indication of overall biomechanical struc- dence is sufficiently ambiguous to be interpretable as indicat- tural integrity. ing varying degrees of population continuity, replacement, The resultant parameters for the samples are compared andyor gene flow. Moreover, the biological bases and hence graphically, for the humerus and initially for the femur by using phylogenetic usefulness of most of the morphological traits lne-lne plots of the resultant values. For the humerus, given its commonly used are simply unknown, making it uncertain what normally non-weight-bearing role in humans, the logarithmic is being analyzed. transformation appears to be adequate to adjust for allometric The simple accumulation of additional neontological and effects, especially of cross-sectional measures vs. bone length. paleontological data and its analysis by current techniques are For the femur, however, load levels are dependent on both insufficient for the resolution of these phylogenetic issues. For body mass (weight and momentum) and beam characteristics. these reasons, it may well be scientifically more profitable, To correct for documented variance of ecogeographically once one reasonably can define paleontological samples and patterned Late Pleistocene human body proportions (9, 10, their distributions in time and space, to look at changing 14–16), bi-iliac breadth was used to represent variance in body patterns of biology and behavior, no matter what the original laterality, and femoral length was used for both beam length genealogical relationships were between the groups. and to represent stature. Given the relative constancy of bi-iliac breadth within ecogeographically defined human Materials and Methods groups (14), bi-iliac
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