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Ences Are Reflected in Bony Morphology, They Should Anatomy 8 5 Early Hominid Postcrania and Locomotor Adaptations Randall J. Thompkins The locomotor system of modern man is a complex ical characteristics of early hominid1 postcrania is in and unique adaptation, and the evolutionary origin considerable part a lack of agreement about approp- and history of bipedalism are of fundamental impor- riate techniques and relevant parameters. Much ofthe tance in understanding the pattern of human evolu- earlier literature was on a purely descriptive level, and tion. For this reason much attention has been focused later statistical treatment gave ambiguous results. This upon the postcranial skeleton of the early hominids. is partly because of different workers measuring the The fossil evidence ot Australopithecus fortunately in- same set of fossils and getting different results; also, cludes numerous postcranial remains, and these have access to the original specimens is not readily available been analyzed to determine locomotor capabilities. to all workers, who therefore must rely upon pub- Bipedal, erect posture inAustralopithecus has been dem- lished reports and casts. Many of the original speci- onstrated, but the full spectrum oflocomotion in these mens are not fully prepared for study and are in need hominids is not known. The use of extant models has of reconstruction. A general consensus on placed undue emphasis on the striding gait and the operationalism of method is needed with respect to: question of its presence in Australopithecus. (a) the basic parameters involved, (b) how to obtain the Methodological problems have hindered progress in necessary data from the fossils (what to measure on the reconstructive anatomy and biomechanical analysis. bone and how to measure it), (c) how to standardize for Since the locomotor system influences most activities effects of body size, and (d) the appropriate statistical of an organism, differences in locomotor capacities methodologies. Isolated traits must undergo extensive often have phylogenetic significance; if these differ- comparative study, and then biomechanical recon- ences are reflected in bony morphology, they should struction, to validate preliminary inferences. Ulti- be appropriate for taxonomic criteria. Discerning the mately one hopes to integrate fossil postcranial data range of locomotor behaviors of the aus- into what Zihlman (1977) calls a behavioral- tralopithecines is a challenging and important task for evolutionary framework which (a) relates the human evolutionary studies if we are to assimilate the locomotor morphology and behavior to an environ- anatomy, ecology, behavior, and systematics of Au- mental context, (b) correlates the locomotor behavior stralopithecus into a synthetic, holistic view of the evolu- with the morphology, using living models, and (c) at- tion of the Hominidae. tempts a synthetic limb/joint analysis. These goals may be reached via a combination of techniques, including Analysis of Postcranial Material stress analysis (Preuschoft 1971), cineradiographic There are a number of paleontological problems analysis of related extant forms (Jenkins 1972), central to the analysis of post-cranial remains; these biomechanical reconstructions, including bone load- include incomplete and damaged specimens plus the ing patterns (Zihlman and Hunter 1972; Lovejoy et al. problem ofvariation. Several critical parameters: body 1973; Lovejoy 1977), andjoint reconstruction analysis stature and weight, sex, and age, are often, by the (Zihlman 1977). nature of the hominid fossil record, indeterminable. Multivariate statistical approaches, while very prom- This frequent lack of data on gross body proportions ising, have yielded some contradictory results with re- and paleodemography of the sample hinders any at- gard to australopithecine postcranial anatomy. Using tempt at biomechanical analysis of the postcranial multivariate techniques (specifically canonical material. Another frequent problem is that of associa- analysis) on a number of postcranial fossils of Aus- tion; do skeletal parts found in close proximity actually tralopithecus, Oxnard (1975) has stressed the unique- belong to one individual? The discovery in recent ness ofthese fossil remains, their clear differences with years ofseveral partial or relatively complete skeletons fossils placed in Homo, and the elimination of Aus- ofPlio-Pleistocene Hominidae will go far in illustrating tralopithecus as a direct ancestor ofHomo. Other work- the morphological pattern ofAustralopithecus and early ers, using the same fossils and multivariate techniques Homo, in delineating any differences within these taxa, (principal coordinate and canonical analysis), have and particularly in yielding suitable material (of one concluded that early hominid lower limb (McHenry individual) for the analysis ofjoints. Such analysis has and Corruccini 1975a) and upper limb (Ciochon and been infrequently attempted yet is crucial to a compos- Corruccini 1976) morphology is clearly allied with ite biomechanical study. Hominidae, and approaches, but is not identical with, The current lack ofconsensus about the morpholog- that of Homo. 8 6 The problem of how to compensate for size effects should be accounted for by use of appropriate statisti- in both uni- and multivariate analysis is critical. A cal techniques, and finally a total biomechanical pat- frequent technique to eliminate size effects is to con- tern should be reconstructed. It is then possible to vert measurements to angles or indices, and create address the question of differences in locomotor be- ratios between the structure being compared and some haviors between a fossil taxon (Australopithecus) and the morphological feature used as a size standard. Ratio extant relative (Homo), as well as possible differences data often cannot be compared for lack of matching or within the fossil taxon itself (i.e., between gracile and suitably preserved body parts. Multivariate techniques robust Australopithecus). Then paleoecological, have the potential to avoid latent size effects. Corruc- phylogenetic, and taxonomic inferences may be de- cini (1975) has criticized Oxnard and stressed the rived, whether differences between the various groups necessary distinction between size and shape. He notes under consideration exist or not, since either result that statistics computed directly from raw measure- would have significance in elucidating the pattern of ments are dominated by simple size differences. There evolution of early Hominidae. has been a tendency to deal exclusively with isolated bones in such statistics, and to make conclusions solely Early Hominid Fossil Postcrania from those data (e.g., Oxnard 1968a & b). The isolated Early hominid postcranial fossils are relatively rare bone is not the functional unit in limb mrorphology, when compared to more abundant dental, gnathic, and analysis should incorporate available joints and cranial parts. This is unfortunate since locomotor (Ciochon and Corruccini 1976). capabilities are so important in any organism's be- Two important issues in statistical studies are what havioral repertoire, and especially so in the evolution measurements are to be employed and how to inter- of the Hominidae. Recently, however, the sample of pret the results. When isolated bones instead ofarticu- postcrania has become fairly complete and most body lated joint complexes are utilized the measurements parts are now represented, some by relatively com- used severely limit their utility, and factors such as plete and undistorted fossils, some by the remains of body weight are often ignored (Zihlman 1975). Great several partially complete skeletons (Table 1). A review care must be taken not to "load" the multivariate pro- of the fossil evidence will be offered for each major gram with redundant measurements. Lovejoy (1977) anatomical region, describing basic morphologies and has emphasized that this problem of the interrelation- delineating any biomechanical differences. ship of metrically assessable characters is the greatest limitation to the interpretation of multivariate results. Axial Skeleton As Day (1973) has pointed out, multivariate studies Early hominid vertebrae are relatively rare in the have in recent years been at best a complement to fossil record, and the best specimens are fromjust two anatomical description. With additional fossil material individuals, Sts 14 (Sterkfontein) and AL 288-1 (Afar). and progress in multivariate techniques, the full im- Both Day (1977) and Robinson (1972) stress the small pact of advanced biometrics should be realized. size of the vertebral bodies. Small size is also evident in An important consideration is the level of inference the two articulated vertebrae (in matrix) of KNM-ER which is most appropriate for reconstructing postcra- 164 (East Lake Turkana, Day 1977). A relatively well nial anatomy and locomotor capacities in early developed lumbar curvature is evidenced in Sts 14 hominids. Preliminary anatomical description should (Robinson 1972), which concurs with lower limb evi- not be used as a final analysis. The observation of dence of bipedal erect posture. morphological differences is not in itself evidence of The sacrum of Sts 14 is of typically hominid disposi- locomotor differences between two groups. The range tion and shape, but in overall size significantly smaller of variation within a population must be accounted than that of modern man. A smaller auricular surface, for; differences in morphology, if statistically signifi- smaller lumbosacral articular
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