Morphometric variation in Plio-Pleistocene hominid distal humeri.

The magnitude and meaning of morphological variation among Plio-Pleistocene hominid distal humeri have been recurrent points of disagreement among paleoanthropologists. Some researchers have found noteworthy differences among fossil humeri that they believe merit taxonomic separation, while others question the possibility of accurately sorting these fossils into different species and/or functional groups. Size and shape differences among fossil distal humeri are evaluated here to determine whether the magnitude and patterns of these differences can be observed within large-bodied, living hominoids. Specimens analyzed in this study have been assigned to various taxa (Australopithecus afarensis, A. africanus, A. anamensis, Paranthropus, and early Homo) and include AL 288-1m, AL 288-1s, AL 137-48a, AL 322-1, Gomboré IB 7594, TM 1517, KNM-ER 739, KNM-ER 1504, KMN-KP 271 (Kanapoi), and Stw 431. Five extant hominoid populations are sampled to provide a standard by which to consider differences found between the fossils, including two modern human groups (Native American and African American), one group of Pan troglodytes, and two subspecies of Gorilla gorilla (G.g. beringei, G.g. gorilla). All possible pairwise d values (average Euclidena distances) are calculated within each of the reference populations using an exact randomization procedure. This is done using both raw linear measurements as well as scale-free shape data created as ratios of each measurement to the geometric mean. Differences between each pair of fossil humeri are evaluated by comparing their d values to the distribution of d values found within each of the reference populations. Principal coordinate analysis and an unweighted pair group method with arithmetic averages (UPGMA) cluster analysis are utilized to further assess similarities and differences among the fossils. Finally, canonical variates analysis and discriminant analysis are employed using all hominoid samples in order to control for correlations among variables and to identify those variables that discriminate among groups; possible affinities of individual fossils with specific extant species are also examined. The largest size differences, those between the small Hadar specimens and the two largest fossils (KNM-ER 739, IB 7594), can be accommodated easily within the ranges of variation of the two Gorilla samples, but are extreme relative to the other reference samples. The d values between most of the fossils based on shape data, with the notable exception of those associated with KNM-ER 739 and KNM-ER 1504, can be sampled safely within all five reference samples. Subsequent analyses further support the inference that KNM-ER 739 and KNM-ER 1504 are different from the other hominid humeri and possess a unique total morphometric pattern. In overall shape, the distal humeri of the other fossils (non-Koobi Fora) are most similar to living chimpanzees. The distal humerus of Paranthropus from Kromdraai (TM 1517e) is most similar to one of the Hadar specimens of A. afarensis (AL 137-48a), whereas the first specimen of A. africanus from Sterkfontein (Stw 431) is not closely linked to any of the other australopithecines. The A. anamensis humerus from Kanapoi exhibits no special affinities to A. afarensis or to modern humans.