Petite Bodies of the “Robust” Australopithecines
HENRY M. McHENRY Department of Anthropology, University of California, Davis, California 95616

KEY WORDS Body weight, Australopithecus, Plio-Pleistocene hominids, Paranthropus, Swartkrans, Sexual dimorphism
ABSTRACT The “robust” australopithecines are often depicted as having large and powerfully built bodies to match their massive masticatory apparatus, but until 1988 the sample of postcranial remains attributed with certainty to this group was very limited. Almost nothing was known about the body of the East African “robust” australopithecine because taxonomic attribution of the postcrania was so uncertain. The body of the South African “robust” australopithecine had to be reconstructed from about a dozen isolated fragments of postcrania. Now a partial skeleton is attributed with confidence to the East African “robust” group along with several isolated bones. The South African sample has more than tripled. Analyses of this vastly expanded sample reveal that a large portion of postcrania attributed t o “robust”australopithecines from Swartkrans Member 1 (35%)are from extraordinarily small-bodied individuals similar in size to a modern Pygmy weighing as little as 28 kg. These small elements include parts from the forelimb, spine, and hindlimb. About 22% of these Swartkrans 1 “robust” australopithecines are about the same size as a modern human weighing about 43 kgs and about 43%are larger than this standard but less than or equal to a 54 kg modern human. Approximately the same pattern is true for the Swartkrans 2 hominids, but taxonomic attribution is less certain. All of the Member 3 specimens are similar in size to the 45 kg standard. The partial skeleton of the East African “robust” australopithecine (KNM-ER 1500)has hindlimb joints that would correspond to a modern human of 34 kgs although the actual weight may be 5 t o 10 kgs greater judging from shaft robusticity and forelimb size. The largest postcranial element attributed with some certainty to the East African “robust”australopithecine group (the talus, KNM-ER 1464) is about the same overall size as a modern human of 54 kgs, although its tibia1 facet is slightly smaller. Although many previous studies have hinted at the possibility that “robust” australopithecines had relatively small bodies, the new fossils provide substantial evidence that these creatures ranged from quite small to only moderate in body size relative to modern humans. These were the petite-bodied vegetarian cousins of our ancestors. Sexual dimorphism in body size appears to be greater than that in modern humans, similar to that in Pan, and less than that in Gorilla or Pongo, although such comparisons are of limited value given the small samples, poorly known body proportions, time averaging, and many other problems.
It has often been assumed that the “ro- thers the notion that petiteness characterbust” australopithecines had large and pow- ized the lineage leading to Homo as opposed erfully built bodies to match their enormous chewing apparatus. Using the term “gracile” to describe the non-“robust” hominids furReceived December 3,1990; accepted June 24,1991.


40-19.g. The largest of the isolated postcranial remains from East Africa were often attributed to the “robust” australopithecines (e.b. Four additional specimens are attributed to A. boisei (Grausz et al. boisei. the large ulna from Omo. old (Vrba. 1989). KNM-ER 1500. The only other South African “robust” hominid postcranial specimens are 4 fragments from Kromdraai and 3 of these are reported to be from the same individual (Robinson. Napier. in age (Brain et al. 1981).8 m. 1959. but the attribution was tenuous because all of them occurred in sediments containing craniodental elements of both Australopithecus boisei and Homo.. The focus is on the South African form because much of the East African material is still in taxonomic limbo.. it is likely that all of the postcrania belongs to that species as well although Susman (1989)notes that one manual proximal phalanx (SKX 27431) has a Homo-like characteristic. erectus..y. KNM-ER 1500. 1991. but uncertainty lurks behind any attempt to attribute isolated specimens because of the contemporaneity of one or more species of non-“robusts. 1988). 1989. 19891. McHenry. L. by Howell. There were more complete specimens in East Africa attributed to “robust’’ australopithecines. Robinson (1972) very tentatively attributed the SK 853 juvenile lumbar to H. the long and robust humerus KNM-ER 739. however.g. There are 13 specimens from Swartkrans Member 2. 1988). Walker. Because the craniodental material from Member 3 is entirely attributed to the “robust” species (Grine.6 to 1... 1978. Susman (1989) reviews this problem and notes that most of the postcranial sample could just a s likely be assigned to either Homo or the “robust” australopithecine. Trinkaus and Long (1990) point out that neither SK 84 nor SKX 5020 can be attributed with any certainty to one or the other hominid species. Walker et al. 1988a. Susman.g. 1976).. Johanson and Edey. Robinson( 1972) described the South African “robust” hominid a s being powerfully built and weighing 150-200 lbs.M. 1978. 1988. The East African sample of “robust” australopithecine postcrania is potentially very large. boisei of East Africa (Grausz et al. boisei because of their association with A. erectus despite its decidedly unHomo-like appearance that has led all other authors to attribute it to the “robust”species (e. SK 84. 1989). . 1988) and a partial skeleton and four isolated postcranial bones have been identified a s belonging to A. Howell. 1989. Susman and Brain. 1971. Leakey. Susman (1988a) attributes the thumb metacarpal. McHENRY to the massiveness of the “robust” hominids. Leakey et al. Robinson. 1973.y. and the four isolated postcranial pieces do provide some clue a s to body size of that species.b. by Howell. 1988a. Howell and Wood. Howell (1978) lists 1 4 . to H. The purpose of this paper is to assess the body size of the “robust” australopithecines base on the newly expanded sample..5 and 1 m. One specimen. Susman and Brain. Robinson.. the SK 315513 pelvic fragment.. Until 1988 the published sample of “robust” australopithecine postcranial specimens from Swartkrans consisted of 12 fragmentary fossils.446 H. is one of the few certainties since it is associated with a fragmentary mandible that can be identified as A. Fortunately the Swartkrans sample has almost quadrupled since 1988 (Grine and Susman. 1972). 1973.b). 1972. There are 23 specimens from Member 1of Swartkrans that are 1. (19741. Until recently body size reconstruction of the “robust” australopithecine has been hampered by a paucity of postcranial fossils. Most of these sepcimens are assigned to the “robust” australopithecine species (Day and Scheuer.” The partial skeleton. but the author agrees with McHenry (197513) that it should be attributed to Australopithecus robustus. The partial skeleton of A. the massive femoral shaft. McHenry et al. Grine (1989) assigns about 33%of the craniodental material from Member 2 to Homo and the rest to the “robust” australopithecine species. 1973. Rightmire. 1971. This makes the task of sorting the isolated postcrania very difficult. 1978. 1985a. KNM-ER 736 by Leakey. In this paper the East African material is treated only tangentially because of the difficulty in attribution. 1972. Swartkrans Members 2 and 3 are similar to Member 1 in fauna and may be between 1. 1972). 1972. was assigned to Homo erectus by Brain et al. MATERIALS AND METHODS The South African fossil sample consists of 57 postcranial specimens from Swartkrans and 4 from Kromdraai.. Susman.Popular artistic reconstructions often pictured the “robust” australopithecines as having larger and more massive bodies than the “gracile” forms (e. Walker et al. 1988. boisei craniodental material. 1974.

is smaller than the equivalent vertebrae in the diminunitive Sts 14 skeleton (Robinson. 19831.99 and the relationship between these variables is described by the least squares equation logWt = 2. 1981) found enough difference between the Swartkrans and Kromdraai samples to justify the retention of a specific distinction between them and therefore used the species name crassidens for the Swartkrans “robust” australopithecine. 1986. The immature lumbar vertebra. and 2 Akka Pygmies (one male and one female). male North American. Although the facial architecture of all the “robust” hominids can be interpreted as different expressions of a similar basic plan (Rak. The medium-sized specimen is a 35 year old female South African Bantu with a n estimated weight of 45 kgs. robustus. and 3 male and 3 female Hylobates lar specimens. It is also possible that the craniodental similarity between East and South African “robust”australopithecines is not matched by postcranial similarity. SK 853. What is astounding to this author is the fact that so many of the Member 1 hominids are this small. the correlation between the log of weight and the log of femoral head size is 0. is slightly smaller in all dimensions (e.8903. and female North American averages is 0. 1978. Out of 23 Member 1 postcranial fossils. 1950a) placed the Swartkrans hominids in the taxon Paranthropus crassidens. 8 male and 4 female Gorilla gorilla specimens. SKX 5017. For several decades most authors agreed with Campbell (1963) and others that they should be referred to a s A.9844 logFem-2. the discovery of KNM-WT 17000 (Walker et al. The term “robust)’may simply be a description of evolutionary grade distinguished by megadontia. 6 skeletons of the diminutive Khoisan people (of unknown sex). 5 male and 7 female P. although some (e. The largest is a 29 year old female from the Schultz collection with a recorded weight a t death of 54 kgs.PETITE BODIES OF “ROBUST AUSTRALOPITHECINES 447 The human comparative sample consists of 32 males and 23 females from North America of known body weight. The smallest is a female Akka Pygmy with a n estimated weight of 28 kg.048. some of these specimens are slightly smaller than the standard. and East African megadonts are not clearly understood. Grine. length. 8 male and 10 female Pongopygmaeus specimens. The well-preserved first metatarsal. Generalizations about South African “robust” australopithecine postcrania may not be applicable to the East African sample. Howell. Using the male and female ape averages. the major axis and reduced major axis equations produce the same predictions a s are derived from these least squares formulae. There are 6 male and 9 female Pan troglodytes specimens. Three human skeletons serve a s size standards. These skeletons define three size categories to compare each fossil specimen.. paniscus specimens. The last thoracic (SK 3981a) and .g. 1988) showed that a t least some and perhaps many cranial and dental features that appear to unite the “robust” species as a monophyletic clade distinct from all other hominids are probably derived from parallel evolution related to the mechanical demands of heavy chewing. The correlation in the human sample using the Pygmy. The original describer (Broom.g. If this is true. All of these apes are associated with wild-collected body weights. Kromdraai. Although dozens of measurements were taken on this comparative sample. The term “robust”austra1opithecineis useful because the taxonomy and phylogenetic relationships of the Swartkrans.98 and the regression formula is logWt = 1. The small category consists of fossils that are approximately the same size as the female Pygmy skeleton with a calculated live body weight of 28 kgs. Walker and Leakey. 1972). only the vertical diameter of the femoral head is used in the present study. and tibia1 lengths following Olivier’s (1976) correlation axis and by deriving weight from stature using the power curve given in Jungers and Stern (1983). 1988) agree with Robinson (1972) that the “robust” australopithecines are generically distinct from other Plio-Pleistocene hominids and should be placed in the genus Paranthropus. With such high correlations. The body weights for the Khoisan and Pygmy samples are estimated by calculating stature using humeral. then it may be improper to group the “robust)’ australopithecines in a genus that is distinct from all other hominids. In fact... Khoisan. femoral. Several authors (reviewed in Grine. RESULTS AND DISCUSSION Table 1 presents the classification of the Swartkrans hominid postcrania into three size categories.71251ogFem-1. 8 (35%)are as small as the Pygmy. diameters of the proximal and distal ends) except the shaft is slightly more robust.

although SKX 5018. SK 84. last lumbar (SK 398113) are much smaller than the Pygmy's. Approximate size categories of the Swartkrans hominid postcranium Small' SKX 5016 Distal ohalanx. Five of the 23 Member 1fossils are about the same size as the 45 kg skeleton. 19. 11.8 . thumb SKX 5018 Proximal phalanx. 3Larger than the 45 kg standard skeleton but smaller than the 64 kg female North American skeleton. but SK 50 has a thicker iliac blade anteriorly (14. hand 'Range in size from slightly smaller than a Pygmy skeleton (with a n estimated weight of 28 kg) to slightly smaller than a small female Bantu skeleton (with a n estimated weight of 45 kg).5 mm).2 vs. 1in Susman. 1989). 5019. is about the same size as the Kromdraai ulna (TM 1517. hand SKX 27431 Proximal phalanx. is about 1mm longer and slightly thicker than the Pygmy.5 mm) and thickness of the iliac blade in the region of the sciatic notch (14. but considerably broader in the mediolateral direction (see Fig.0 and 36. 4Susman 11988a) attributes to Homo. The pollicial metacarpal. The species attribution of this specimen is in some dispute (Trinkaus and Long. and 8963 appear to be approximately the same size as the phalanges of the 45 kg skeleton. zApproximately equal in size to a small female Bantu skeleton (with a n extimated weight of 45 kg). foot SKW 2954 Metacarpal IV SKW 3646 Metacarpal 111 SKX 19576 Proximal phalanx. but the reconstructed acetabular diameters are smaller than the standard. the fossil is smaller than the standard in its minimum width of the ilium (50. Susman. SK 3155b.448 H. Ten of the Member 1 postcranial fossils (43%) are larger than the 45 kg standard skeleton but in most dimensions these are smaller than the 54 kg skeleton. The immature pelvic fragment.2 vs. hand SK 50 0 s coxae I I1 SK 84 Metacarpal I4 SK 853 Immature lumbar ver t ebr a SK 3155b Immature 0s coxae SK 3981a Thoracic XI1 SK 3981h Lumbar V SK 45690 Proximal phalanx. have smaller femoral heads than the 54 kg human (34.5 mm).1 vs. hand SKX 36712 Middle phalanx. Size attribution of the hand phalanges is difficult because it is not certain from which ray the specimens come. foot SKX 3498 Triquetral SK 82 Proximal femur SK 97 Proximal femur SK 854 Axis SKW 14147 Metacarpal V SKX 5020 Metacarpal I SKX 5021 Middle phalanx. 1989) and the middle-sized comparative skeleton.Table 2). hallux SKX 5017 Metatarsal I SKX 5022 Mid phalanx. The SK 50 pelvic fragment is too crushed and distorted to compare easily. The proximal ulna. hand SKX 31117 Medial cuneiform SKX 33355 Middle phalanx hand SKX 33380 Metatarsal V SKX 35439 Middle phalanx. has a smaller acetabulum than the Pygmy's but its iliac blade is slightly higher and broader. 1990). hand SKX 2511 Proximal phalanx. The proximal femora.M. hand SKX 27504 Distal phalanx. hand SKX 8761 Proximal ulna SKX 8963 Distal phalanx. hand SKW 1261 Middle phalanx. McHENRY TABLE 1. hand SKX 22741 Proximal phalanx. hand (IV?) SKX 5019 Middle phalanx. Of the 3 unreconstructed measurements reported by McHenry (1975b. hand SKX 3602 Distal radius SKX 3774 Distal humerus SKX 34805 Distal humerus SK 18b Proximal radius SKX 1084 Patella SKX 3062 Middle phalanx. The beautifully preserved distal pollicial phalanx. hand SKX 35822 Proximal phalanx. SK 82 and 97. SKX 5016. SKX 8761. is about 1mm shorter than this standard skeleton. 56. hand SKX 3699 Proximal radius 11 1 SK 85 Metacarpal IV SKX 247 Metatarsal 111 SKX 3342 Thoracic vertebra SKW 344 Middle phalanx.

then these large hindlimbs would be matched by much larger forelimbs than are present in the Member I sample. then one can show with a chi-square test that the two criteria (size and body part) are independent (i.and hindlimb parts in Member 1 as a contingency table. there are not more large forelimbs and small hindlimbs than one would expect if ape-like proportions prevailed). and references therein). 288-1. 25. 1974) although it is possible that the bones are from two or more individuals.A. 1983a.. a slightly more robust shaft. then the small forelimb elements should be matched by hindlimb elements much smaller than the smallest ones recovered from Swartkrans. If the Swartkrans hominids did have foreto hindlimb proportions that were intermediate between modern humans and apes. because modern humans are used as the standards and early hominids had body proportions quite different from those of modern humans. For example. Some early hominid species certainly did appear to have unhumanlike fore. SKW 14147. The Member 2 remains from Swartkrans fall within the same range of size variation. the Member 1 craniodental sample is composed of over 95% “robust” hominids. This is not the case. Perhaps sampling error explains this size homogeneity. referred to Paranthropus robustus by its discoverer (Broom. 1978. TM hindlimb size quite different from modern humans (Johanson et al. The partial skeleton from Olduvai. The two smallest specimens are between the Pygmy and Bantu standard skeletons in size.8 mm).4 and 32. O. is slightly shorter than the standard.b.0 and 24. The triquetral (SKX 3498) and foot phalanx (SKW 1261)are intermediate in size between the two standards. 1983. At least 9 and perhaps 11different individuals are represented by the craniodental remains. but the Member 3 hominid sample is entirely composed of the “robust” australopithecines. Member 2 has 12 to 17 “robust” individuals and 4 to 5 members of the genus Homo. referred to H. however. has a shorter femur than one would expect from the length of her humerus if scaled-down modern proportions are compared (Johanson. what size forelimb matches the large hindlimbs such as SK hindlimb. habilis. however. if one treats the distribution of small and large elements relative to fore. SKX 5021. the bias should result in too many fossil hindlimb elements in the small-bodied category. Perhaps the lack of size variability is partly the result of the fact that no Homo specimens are mixed in with the “robust”hominids. McHenry. however (Grine. 1982. KNM-ER 1500. possesses fore.3 vs. however. There are 6 specimens that match the 45 kg skeleton in size. The largest of the forelimb remains in the Member 1sample is the pollicial metacarpal. And the reverse is true: if they had ape-like proportions. has fore. ufarensis.. although the foot phalanx (SKW 344) is slightly shorter but more robust. boisei. has a ratio of fore.7 mm). Perhaps Table 1 is of limited value. 38. The partial skeleton of A. These large transverse shaft diameters do not imply a larger overall body size. 1988.L. In this case.b. because all of the non-Homo femora in the Plio-Pleistocene hominid collection appear to have disproportionately expanded transverse shaft diameters (McHenry. 62. As Susman (1988a. are slightly smaller than they are in the 54 kg human.PETITE BODIES OF “ROBUST AUSTRALOPITHECINES 449 vs. the hands and feet of the Swartkrans . The partial skeleton hindlimb proportions intermediate between modern humans and apes (Grausz et al. 1989) shows. 25.. because only ten specimens are represented and many of these might be from the same individual.82. falls within the range of lengths in the human skeleton. for a dissenting view). All 10 postcranial fossils from Member 3 are similar in size to the 45 kg standard. According to Grine (1989) and Susman (19891.3 mm).. Jungers. 1982. hindlimb proportions. 1938) and to A. proportions intermediate between modern humans and apes (McHenry.6 vs. et al. SKX 5020. Another test is to mix and match. 1989).e. 3774. and 97? If ape-like proportions are assumed. but larger transverse shaft diameters (30. 1991). For example. but see Wolpoff. The fifth metacarpal. The humeral and radial specimens. robustus by many (see Howell. and 34805. 1987). then the use of human standards in Table 1 should produce biased results. Why there is so little size variation in the Member 3 sample is difficult to explain. There are 4 specimens in the large category and none that are larger than the 54 kg standard skeleton. SKX 3602. 1978). which has the same length as the 54 kg human. The length of the middle phalanx of the hand. slightly smaller shafts in the anteroposterior direction (25. and a slightly narrower base. The partial skeleton of Kromdraai.H.

Napier.6 and 55.3 30. Even the forelimb elements are smaller than this human standard (Table 1)so they are much smaller than a 60 kg ape.3 35. 1964).6 33. McHenry and Corruccini. but the hips ofthe“robust”austra1opithecines were unlike either.9 60. 1988a). From the TABLE 2. 1991).6 and 43. 1978.0 kgs using the human formula and 47.7 kgs for SK 82 and SK 97 given in McHenry (1976) are probably too large because they are derived from multiple regression fromulae that use shaft diameters and neck length.0 30.0 SK 82 SK 97 SK 31551.2 kgs using a formula that included apes only. SK 3155b. The values of49. 1975a. SKX 5020. 1988)and the thumb metacarpal. Jungers (1988b) reviews this issue and points out that hip-joint size was not relatively expanded in the non-Homo early hominids. Assuming human proportions.0 mm by the method described in McHenry. show a human-like pattern that is uniquely different from any pongid (Susman. corresponds to a weight of 30. Based on the overall size distribution of all the hominid postcrania now known from Swartkrans. it appears that neither modern apes nor humans provide the ideal guide to body weight prediction from the size of the hip. While this specimen is a subadult with an unfused iliac epiphysis. and TM 1605) and two proximal femora (SK 82 and 97) it is clear that the hip was adapted to human-like bipedalism (McHenry. 1976. The big toe metatarsal.3 kg (its femoral head size is predicted to be 30. but perhaps the actual weight of the larger morph is somewhat greater than 40. But compared toHomo. of course. The predictions derived from the human formula presented in Table 2 appear to be too small when compared to the overall size comparisons of Table 1.9 and 60. and distal phalanx. ischial. Table 2 presents the estimated body weight derived from hip-joint size in the three best preserved hips in the Swartkans sample. correspond to weights of 37. 197513). 1975a).6 kg) that seems too large considering the distribution of all the other postcranial elements that are smaller than the 54 kg human standard. For this bipedal hominid with so many human-like attributes to its postcrania. SKX 5017. The estimates of body weight presented in Table 2 should be regarded as only the roughest of approximations. Using the ape formula its weight is 33.0 43. They are based on hi-joint size in modern humans and apes. 1972). There is some intuitive and empirical evidence that suggests that at least some early hominids had hip joints the size one would expect from modern humans of the same body weight. 1975.7 54. Robinson.R. Certainly the shafts are much thicker than would be expected from a hominid this small. Jungers (1988a) predicted weights of 45. SKX 5016. 1989).6 kgs using the ape formula. 1975a. its acetabulum had probably reached adult size since its ilial. is designed for human-like walking except for the dorsally narrowed distal articular surface (Susman and Brain. The two proximal femora.3 40. the hip joint itself appears to be small relative to other dimensions of the 0s coxa or femur (McHenry. although shaft diameter may be a poor predictor of body weight in early hominids (see above).. Estimated body weight of member 1 Swartkraus hominids from hip-joint size Specimen Predicted Body Weight (kg) Human Formula Ape Formula 37. But the formula based on modern apes gives a value for SK 97 (60.3 47. He presents evidence that suggests that body mass predictions in non-Homo early hominids are more reasonable when the predicting formulae are based on hominoid samples that exclude Homo sapiens (Jungers.1 and 65.8 and 52. These values also appear to be somewhat too high from the perspective of the current study.3 33.8 kgs from these two femora using a formula derived from all hominoid species and 52. Average Average large morph (male?) Average small morDh (female?) three South African “robust” australopithecine 0s coxae (SK 50. the predictions based on the human formula may be closer to the mark. Perhaps the best estimates of body weight for the Swartkrans hominids lie . and pubic components had nearly completely fused (McHenry.M. however (McHenry.450 H. the acetabulum. SK 315513. J.0 kg.6 43. MrHENRY hominids are remarkably human-like without the long and curved phalanges associated with ape-like bodies.3 kgs (the average estimate from the human formula for SK 82 and 97). SK 82 and 97. 1988a.

many of the elements display a degree of robusticity that is greater than that seen in modern humans. The remaining hominid postcrania are about the same size as the 45 kg standard human skeleton. First.. As noted above. 1985. 1972). While the lengths of the metacarpals.between 25 and 35 kgs). then several interesting generalizations emerge. appears to be about the same size as the more complete specimen from Koobi Fora Area 123 (KNM-ER 1504) which in trochlear width. Sites that have yielded taxonomically identifiable craniodental material of “robust”hominids also contain specimens attributable to Homo. and Olduvai (O. this specimen falls within the small category of Table 1(i. and A. The average endocranial volume for the 2-1.. Both of these humeri are unusually flat in anteroposterior dimensions. although its small size implies that it was a juvenile. KNM-ER 1825. erectus (KNM-WT15000. There is evidence that early H. Only one endocranial volume is recorded for the Swartkrans “robust” hominid.y... 1985. probably belonged to an individual that was equally small. these are above the value for A. erectus was considerably larger than the South and East African “robust”hominid (Jungers. is about the same size as the large skeletal standard ofTable 1(i. Johanson and Edey.H.McHenry. and Omo L 338y-6 at 427 cc: Holloway. Walker et al. 36) ulnae? These specimens are far too big to fit such petite hominids. The fragmentary atlas vertebra from Ileret 6A.e. Judging from the size of the humerus and ulna of early H. although the postcrania is fundamentally human-like. The other three postcranial pieces from Ileret 6A are from one or more largerbodied hominids. metatarsals. 1988. The East African sample of “robust” australopithecine postcranial remains is not well known because of the problem of taxonomic attribution. Leakey and Walker. is too fragmentary to determine its age at death. old “robust” australopithecine of East Africa is 510 cc (McHenry.the shaft diameters are often greater than the human equivalent. (1989) reports that the KNM-ER 1500 partial skeleton is only slightly larger than the diminunitive A. ufarensis (415 cc. In the present study. 1989) so their larger endocranial volumes may be relatively closer . KNM-ER 17000 at 410 cc: Walker and Leakey. 1981. these specimens do not fit within the expected range for the male H. The pelvic fragment. To what species. Walker et al. africanus 442 cc. One of the four postcranial specimens from Ileret 6A.e. the Omo (L40-19)ulna. 1989). Taken together.Brown et al. It is not difficult to imagine enormous within-species size variability when one reflects on body size differences within African populations today. 1988. KNM-ER 1500. 1988) also similar in size to this large standard. Brown et al. The talus. should one assign the massive forelimbs such as the KNM-ER 739 humerus. relative brain size appears t o have increased from the size seen in earlier hominids. Simons. In absolute terms.and ape-based predictions. early East African “robust” australopithecines (i. 1989.. 1970). The size of its knee and ankle joints corresponds to a human of about 34 kgs. The very fragmentary distal humerus. 19811. and 4 elements from Ileret Area 6A (Grausz et al. TM 1605. 19891. equivalent to a 54 kg modern human). 1988). 1988a.e.7 m.PETITE BODIES OF “ROBUST AUSTRALOPITHECINES 45 1 between the human.L. capitular area. 1988. and in most mediolateral dimensions is similar to the 54 kg standard skeleton. There is little or no evidence suggesting that these species were significantly smaller bodied than the “robust” hominids of South Africa. very little is known about the postcranium of the Kromdraai “robust”australopithecines.. Such greater robusticity implies that the human-based predictions may underestimate body weight. Attempts t o classify isolated postcranial specimens have not been successful (McHenry. The only postcrania known to belong to the East African “robust” australopithecines are the partial skeleton. a proximal fragment of a third metatarsal (KNM-ER 18231. especially in the minimum a-p diameter of the trochlea. KNM-ER 1464. SK 1585. then. Perhaps these large forelimbs represent part of the wide range in variability in size within East African early hominid species. and phalanges correspond t o the categories established in Table 1.Leakey and Walker. erectus. 288-1 partial skeleton. Although the sample is wonderfully expanded from Swartkrans. the available evidence suggests that the East African “robust” australopithecines also had petite bodies relative t o their hypertrophied masticatory apparatus. Holloway. with a value of 530 (Holloway. If it is true that the “robust” Plio-Pleistocene hominids were relatively small in body size. KNM-ER 1824. then.

Berkeley. A modern hominoid species with a n average endocranial volume of 500 cc will include normal variants as small as 410 cc (Walker and Leakey. for permission to study the comparative material in their charge and for many kindnesses.452 H. 4. and the staff of the Department of Anthropology. and the staff of the Musee d’Afrique Centrale. Thorington and the staff of the Division of Mammology. W. sample sizes are so small that much uncertainty remains.S.R. for permission to study the original fossil material in their charge and for numerous kindnesses. and the staff of the Powell-Cotton Museum. Although all of the postcrania attributed to “robust” australopithecines derive from bodies that are relatively small compared to modern humans. Brain and the staff of the Transvaal Museum and R. Pilbeam. At least a third ofthese newly attributed . Powell-Cotton. Biegert. Sexual dimorphism in body size among “robust” australopithecines is difficult to measure. It seems unlikely on available postcranial evidence. The variability in the crosssectioned area of the corpus in this species is as great a s the highly dimorphic modern gorilla (Walker and Leakey. The largest specimens are about the size of a 54 kg modern human. But such a generalization is uncertain because one cannot control for time averaging. D.F. Leakey. Howells. R. Smithsonian Institution. I also thank C. van den Audenaerde. This implies a degree of sexual dimorphism that is greater than that seen in modern H. however. The cross-sectioned area of the mandible of A. that the dimorphism in body size in this species was a s great a s that seen in mandibular corpus size.W. Grine FE. 5.B. Partial funding was provided by the Committee on Research. The degree of postcranial size variation in both East and South African “robust” australopithecines may imply a level of body size sexual dimorphism considerably greater than in modern Homo sapiens. D. boisei. boisei is similar to that of Members 1 and 2 Swartkrans. Until 1988 very little postcrania had been attributed with any certainty to the “robust” australopithecines of East and South Africa and therefore doubts remained as to the body size of these hominids.T. 2. McHENRY to those of the “robust” australopithecines. Schultz.D. the late J. Leakey. M. but not a s great as that seen in modern gorillas or orangutans.R. Certainly the range of size variation with the Swartkrans Member 1 and 2 samples is wide (ranging from smaller than the 28 kg Pygmy standard to equal to or larger than the 54 kg standard). Zurich. for example. D. the late A. for example. but the sample size is too small to be certain. F. Many specimens are smaller than the equivalent element in a 28 kg modern human Pygmy. supiens and species of Pun but less than that of species of Gorilla and Pongo.F.M. 3. The variability in body size documented by the postcranial bones attributable to A. Shipman P. Leakey. Lovette. R. their culture and environ- 1. Harvard University. LITERATURE CITED Brain CK. Harvard University. the late L. Howlett. 1988). Even if apelike proportions are assumed. Clark JD. Turner A. Susman RL. Churcher CS. The dimorphism in craniodental remains of A. M. Martin. and C. and the staff of the National Museums of Kenya. The sample from Member 1 Swartkrans. boisei is very strong and implies that body size may also have been dimorphic.E. the estimated body size is relatively small among the known sample of “robust” australopithecines.D. Rutzmoser and the staff of the Museum of Comparative Zoology. Since 1988 a partial skeleton and 4 isolated postcranial elements have been attributed to the East African “robust” australopithecine and the sample of postcrania from Swartkrans has almost quadrupled. Davis. University of California. CONCLUSIONS specimens derive from surprisingly smallbodied creatures. is approximately the same as that seen in modern gorillas whose body weight ranges from 75 to 180 kgs. most shafts are more robust relative to their lengths or size of their articular surfaces. However. might include many variants of the same species that died near the cave at very different times. The massiveness of their masticatory apparatus might imply a n equally massive body.K. ACKNOWLEDGMENTS I thank C. and Watson V (1988) New evidence of early hominids. Norick and the staff of the Lowie Museum. M. and the staff of the Anthropologisches Institut. 1988). Edelstamm and the staff of the Natur Historiska Riksmuseet.A. Most are smaller than or equal to the size of a 45 kg modern human. Stockholm.E.

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