Relationship of squamosal suture to asterion on external skull surfaces versus endocasts of pongids: Implications for Hadar early hominid AL 162-28

The relationship between the squamosal suture and asterion was quantified in 15 hemispheres of eight chimpanzee endocasts that were aligned in the conventional lateral view (i.e., with frontal pole [FP]–occipital pole [OP] horizontal). Using a three-dimensional digitizer, x, y, and z coordinates were collected for the highest and lowest points of the squamosal suture, and the most rostral point of the suture approximate to the coronal suture. Our results were compared to a similar study of the squamosal suture on the external surfaces of chimpanzee skulls that were oriented in the Frankfurt horizontal (Holloway and Shapiro, 1992). The relationship between the squamosal suture and asterion differs markedly between the outsides of skulls and endocasts. Whereas the squamosal suture is very rarely below asterion on the external skull, we found that most of the squamosal suture is located inferior to asterion on endocasts. We also found that the squamosal suture courses approximately 2.0 mm lower on the right side than the left. (An asymmetry of the same magnitude was reported for the external skull but, curiously, in the opposite direction.) It may be that a lowered right squamosal endosuture on chimpanzee endocasts is associated with earlier closure on that side. The discrepancy in results for the external skull versus endocast is partially attributable to orienting chimpanzee skulls in the Frankfurt horizontal, which usually results in the endocasts being tilted so that FP is above OP, i.e., FP-OP is not parallel with the Frankfurt horizontal. Falk's (1985) orientation of the early hominid endocast from Hadar (AL 162-28) is consistent with data determined from endocasts of chimpanzees. © 1994 Wiley-Liss, Inc.     

Some additional morphological and metrical observations on Pan brain casts and their relevance to the Taung endocast

Using an increased sample of chimpanzee brains and brain casts, 32 hemispheres were measured to determine the variability of OP-FP (occipital-pole) and OP-LS (occipital pole-lunate sulcus) arc dimensions and their ratios. The Taung endocast was remeasured to test Falk's assertions that the lunate sulcus is in a pongid position. The average ratio for the chimpanzee brains was 0.218, a value more than 2+ S.D.'s posterior to Falk's placement of the lunate sulcus on the Taung specimen. It is suggested that the chimpanzee and Taung occipital poles have a different morphometric pattern, the former being coincident with the caudal end of the LC (lateral calcarine) fissure. The Taung OP-LS arc consistently measured at least 42 mm, and not 40 mm as claimed by Falk.     

Brain expansion and rotation during hominid evolution

The question of how an endocast (or brain) is oriented within a skull that is positioned in the Frankfurt plane is investigated for African great apes, early hominids STS 71, KNM-ER 1813 and KNM-ER 1470, and modern humans using a 3SPACE digitizer. Our results suggest that, rather than being positioned in the orientation in which isolated brains (endocasts) are conventionally illustrated, brains within skulls that are oriented in the Frankfurt plane tend to be inclined so that the frontal pole is higher than the occipital pole, especially inHomo. These preliminary findings have implications for interpreting early hominid endocasts such as that of AL 162-28.     

Apples, oranges, and the lunate sulcus

Holloway (1984) used a method of direct tape-arc measurements on chimpanzee brain casts to reject the hypothesis that the lunate sulcus is located in an anterior position in the Taung endocast. However, Holloway neglected to measure the occipital pole-lunate sulcus (OP-LS) arc directly on the Taung endocast as he did on chimpanzee brain casts (a crucial part of his methodology); instead, he determined the relative position of Taung's lunate sulcus on the basis of a calculation that confounds direct measurements and measurements from photographs. When arc OP-LS is measured directly on Taung according to Holloway's methods, the feature that has been identified as the medial end of the lunate sulcus is shown to be located within the range that Holloway determined for chimpanzees. Thus Holloway's methodology and data support rather than refute the claim that the lunate sulcus is located in a pongid-like position in australopithecines.     

Hominid carpal,metacarpal, and phalangeal bones recovered from the Hadar formation: 1974–1977 collections

The fossil hominid hand bone collection from the Pliocene Hadar Formation, Ethiopia, is described anatomically. These hand bones, all from A.L. (Afar Locality) 333 and 333w, constitute the largest sample of hominid manus remains thus far recovered from the Plio-Pleistocene of Africa.     

Laetoli toes andAustralopithecus afarensis

The probable misfit between feet, particularly toes II–V, of 3.0-million-year-oldAustralopithecus afarensis from Hadar, Ethiopia, and the 3.5-million-year-old hominid footprints at Site G, Laetoli, Tanzania, casts doubt thatA. Afarensis made the Laetoli trails. We suggest that another species ofAustralopithecus or an anonymous genus of the Hominidae, with remarkably humanoid feet, walked at Laetoli. It would be imprudent to declare thatHomo was present at Laetoli 3.5 million years ago (my) because there is no evidence of brain expansion, advanced tool manufacture, or other non-locomotor hallmarks of the human condition at Site G.     

On the squamosal suture of KNM-WT 17000.

The extensive overlap of the temporal squama on the parietal in KNM-WT 17000 is restricted to a narrow segment of the arc of the squamosal suture. It appears as a long, narrow, posterosuperior extension onto the calvarial wall. The long axis of this extended strip is aligned with what appears to be the most hypertrophied section of the temporalis muscle fan. The substantial differences between the squamosal suture and associated masticatory elements of KNM-WT 17000 and those of Australopithecus boisei provide an opportunity to evaluate the various forces molding the anatomy of the squamosal suture of A. boisei. It is suggested that the extreme flare of the zygomatic arches in the latter accounts for these differences.     

Some comments on the case for Early Pleistocene hominids in South-Eastern Spain

Several sites in the Orce Basin have revealed evidence of the presence of hominids in the Early Pleistocene. These remains are dated to over 1.0 million years, while they may be as old as 1.6 million years. The skeletal remains from Venta Micena in the Orce Basin show a molecular “fossil protein” pattern which aligns them with hominids, but not with equids. This is supported by the anatomical evidence of the two humeral shafts from theEstrato Blanco in the Venta Micena deposits. The biparieto-occipital partial calvaria shows some unusual features if VM-0 is a hominid specimen. the presence of a prominent crest on the internal surface of the occipital fragment adjacent to the point lambda is decidedly unusual for a modern human calvaria. Moreover, theimpressions gyrorum, in the region where the superior parietal lobule of the cerebral hemisphere abutted against the calvaria, point to a bipartite superior parietal lobule with anterior and posterior moieties which, on the endocast, are clearly separated by a depression that represents a sulcus. These morphological traits are rather puzzling if VM-0 is a hominid, and at first they led me to hesitate over the anatomical identification of VM-0. However, the studies of Campillo (1989) and of Campillo and Barcelo (1986) suggest that the features of the fragment VM-0 are compatible with those of a hominid. Because I believe that we do not possess sufficient information on the variability of the endocranial and ectocranial manifestations of the sagittal suture and of its variance with age of the individual, in different hominid species and different equid species, I have not adduced this pattern as evidence in support or rebuttal of the hominid status of VM-0.     

Dental metric assessment of the omo fossils: implications for the phylogenetic position of Australopithecus africanus

The discovery of Australopithecus afarensis has led to new interpretations of hominid phylogeny, some of which reject A. africanus as an ancestor of Homo. Analysis of buccolingual tooth crown dimensions in australopithecines and Homo species by Johanson and White (Science 202:321-330, 1979) revealed that the South African gracile australopithecines are intermediate in size between Laetoli/hadar hominids and South African robust hominids. Homo, on the other hand, displays dimensions similar to those of A. afarensis and smaller than those of other australopithecines. These authors conclude, therefore, that A. africanus is derived in the direction of A. robustus and is not an ancestor of the Homo clade. However, there is a considerable time gap (ca. 800,000 years) between the Laetoli/Hadar specimens and the earliest Homo specimens; "gracile" hominids from Omo fit into this chronological gap and are from the same geographic area. Because the early specimens at Omo have been designated A. afarensis and the later specimens classified as Homo habilis, Omo offers a unique opportunity to test hypotheses concerning hominid evolution, especially regarding the phylogenetic status of A. africanus. Comparisons of mean cheek teeth breadths disclosed the significant (P less than or equal to 0.05) differences between the Omo sample and the Laetoli/Hadar fossils (P4, M2, and M3), the Homo fossils (P3, P4, M1, M2, and M1), and A. africanus (M3). Of the several possible interpretations of these data, it appears that the high degree of similarity between the Omo sample and the South African gracile australopithecine material warrants considering the two as geographical variants of A. africanus. The geographic, chronologic, and metric attributes of the Omo sample argue for its lineal affinity with A. afarensis and Homo. In conclusion, a consideration of hominid postcanine dental metrics provides no basis for removing A. africanus from the ancestry of the Homo lineage.     

Australopithecus afarensis and the single species hypothesis

Ferguson (1989) has recently argued that the variability seen in the fossils assigned toA. afarensis is far more than expected for a single hominid species, and therefore proposes they represent multiple taxa. In particular, he utilizes data on variation in dental metrics and in premolar morphology in support of this hypothesis. A re-evaluation of these data finds the above conclusion to be unwarranted. Variation in dental metrics providesno basis for separating this sample into multiple taxa, regardless of the analog that is used (i.e. modern primate species or fossil hominid species). Additionally, data on P₃ morphology indicate that thepattern of variation seen in the Laetoli/Hadar sample is comparable to the sexual variation seenwithin a single hominoid species. Overall, the balance of the evidence at present indicates that the fossils from Laetoli and Hadar represent a single hominid species,A. afarensis.     

The capitate of Australopithecus afarensis and A. africanus

The capitates of Australopithecus afarensis (AL 288-lw and AL 333–40) and A. africanus (TM 1526) have the identical combination of modern pongid, modern hominid, and unique characteristics. These traits include the combination of a length that is proximodistally shortened (Homo sapiens-like), a facet for the second metacarpal that is distolaterally facing (unique), the reduced styloid process on the third metacarpal (pongidlike), a dorsally placed trapezoid facet (pongidlike), mediolaterally constricted metacarpal III facet (pongidlike), a prominent palmar beak (pongidlike), a single elongated facet for the second metacarpal (H. sapiens-like), a waisted neck (pongidlike), and a reduced amount of “cupping” in the third metacarpal facet (H. sapiens-like). In overall shape the bones are more like H. sapiens than other extant hominids, although they are uniquely different. The two A. afarensis capitates provide no evidence that there are two postcranial morphotypes at Hadar. Available evidence shows that A. afarensis and A. africanus are strikingly similar postcranially. The morphological differences between the capitate of Australopithecus and H. sapiens may relate to the retention of climbing ability and an absence of certain grip capabilities in these early hominids.     

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 possiblity 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 Euclidean 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. © 1996 Wiley-Liss, Inc.     

French Contribution to the study of Human Origins: The case ofAustralopithecus afarensis

For a long time, French scientists have been involved in the study of human evolution and especially of human origins. Their key works in Eastern Africa have led to the discovery of major fossil hominid sites, especially in the Afar region in Ethiopia, where numerous remains ofAustralopithecus afarensis have been unearthed. The major contribution of the French scholars to the interpretation of the Hadar sample was to demonstrate the impact of the postcranial features on taxonomy and phylogeny. Two groups were identified in the sample and by comparison with modern populations of wild primates, these groups are assigned to different taxa. The other major impact was to show that early hominid bipedalism was an exact replica of modern human bipedality.     

The Belohdelie frontal: new evidence of early hominid cranial morphology from the Afar of Ethiopia

Study of the Belohdelie frontal has demonstrated that this four-million-year-old specimen belongs to a very generalized hominid that may be close to the divergence point of the hominid and African ape clades. Features associated with the temporalis muscle in the Belohdelie frontal and other new hominids from Hadar (AL 333-125) and West Turkana (KNM-ER 17000) suggest that the earliest hominids shared a large anterior component of this muscle relative to the extinct and extant apes. Results of this study support the phylogenetic hypothesis put forward by many workers that A. afarensis gave rise to the “robust” Australopithecus and A. africanus clades.     

The locomotor anatomy of Australopithecus afarensis

The postcranial skeleton of Australopithecus afarensis from the Hadar Formation, Ethiopia, and the footprints from the Laetoli Beds of northern Tanzania, are analyzed with the goal of determining (1) the extent to which this ancient hominid practiced forms of locomotion other than terrestrial bipedality, and (2) whether or not the terrestrial bipedalism of A. afarensis was notably different from that of modern humans. It is demonstrated that A. afarensis possessed anatomic characteristics that indicate a significant adaptation for movement in the trees. Other structural features point to a mode of terrestrial bipedality that involved less extension at the hip and knee than occurs in modern humans, and only limited transfer of weight onto the medial part of the ball of the foot, but such conclusions remain more tentative than that asserting substantive arboreality. A comparison of the specimens representing smaller individuals, presumably female, to those of larger individuals, presumably male, suggests sexual differences in locomotor behavior linked to marked size dimorphism. The males were probably less arboreal and engaged more frequently in terrestrial bipedalism. In our opinion, A. afarensis from Hadar is very close to what can be called a "missing link." We speculate that earlier representatives of the A. afarensis lineage will present not a combination of arboreal and bipedal traits, but rather the anatomy of a generalized ape.     

The Taung endocast: a reply to Holloway

Indices of rostrality (ir, ir') are developed to assess the extent to which the medial end of the lunate sulcus (L) is rostrally positioned in photographs and figures of lateral views of primate brains and endocasts, and indices are determined for chimpanzees, SK 1585 and the Taung endocast. Ir quantifies the extent of rostrality as it has traditionally been viewed (in A-P projections) while ir' takes dorsal curvature into account. The ir of the feature that I have identified as the lunate sulcus of Taung is within one standard deviation of the mean ir for Pan and its ir' is within 1.5 standard deviations from the mean ir' for Pan. Both findings are compatible with my earlier statement that the medial end of the lunate sulcus of the Taung endocast is in a pongid-like position. Use of stereoplotting to transfer the position of L from chimpanzee endocasts and brains to australopithecine endocasts is critically assessed: Holloway stereoplotted five chimpanzee brains and then transferred their mean coordinates that describe the lunate sulcus to the Taung endocast. If stereoplotting successfully transfers the extent to which L is rostrally located, one would expect the mean L of Pan and its transferred counterpart in Taung to have identical index values of rostrality. However, the ir of the lunate sulcus that Holloway located on Taung is over two standard deviations lower than the mean ir for the five chimpanzees he stereoplotted to determine its angular coordinates, and Holloway's ir' for Taung is one standard deviation lower than the five chimpanzees' mean ir'. These discrepancies are shown to be due to shape differences, and it is concluded that stereoplotting should not be used to transfer sulci between differently shaped endocasts without correcting for these differences. I also reply to Holloway's criticisms of my use of L/H indices, palpation, techniques for sampling endocasts, and illustration of the Taung endocast. It is shown that there is room on the Taung specimen for the lateral end of L, and the pongid-like sulcal pattern of Taung is reaffirmed. Thus, we do not yet know when human-like sulcal patterns first appeared in the hominid fossil record.     

Subnasal alveolar morphology and the systematic position of Sivapithecus

Recent collecting in the Potwar Plateau of Pakistan has produced several new maxillae attributable to Sivapithecus. Since the subnasal region is preserved in most of these specimens, comparisons with early Miocene hominoid and Pliocene hominid maxillae become possible. On the basis of these comparisons, it has become clear that subnasal/premaxillary morphology distinguishes Asian and African hominoids. Ramapithecus and Sivapithecus share with Pongo an "Asian" subnasal pattern. The Proconsul species from the early Miocene of western Kenya and Australopithecus afarensis from the Hadar Formation of Ethiopia present two subsets of an "African" subnasal pattern. We think it likely that Ramapithecus and Sivapithecus represent a lineage that postdates the last common ancestor of African and Asian hominoids.     

New australopithecine endocast,SK 1585, from Swartkrans,South Africa

The new SK 1585 endocast, found by Dr. Brain at Swartkrans, 1966, is that of a robust australopithecine, matching the endocast of the Olduvai Hominid 5 in volume, and being almost identical to it in morphology. Aside from Olduvai Hominid 5 it is the only robust australopithecine endocast complete enough to permit easy reconstruction, as only a small portion of the frontal lobe is missing. While the gyral and sulcal patterns are not clear, there are a number of features indicating that the brain is not that of a pongid, but that is has been reorganized to a hominid pattern, particularly the occipital, parietal, and temporal lobes.     

Modeling three-dimensional sculptures of australopithecines (Australopithecus afarensis) for the Museum of Natural History of Vienna (Austria): the post-cranial hypothesis.

In March 1999, E. Daynes, a sculptor specializing in fossil hominid reconstruction, asked C. Berge to take over the scientific supervision of the reconstruction of two australopithecine post-crania. The heads had been modeled from two skulls found in Hadar (AL 444-2, AL 417). The sculptures were to be represented in a walking stance. The female proportions (AL 417) are estimated from the skeleton of 'Lucy' (AL 288), and the male proportions (AL 444-2) extrapolated from the female ones. Biomechanical and anatomical data (comparison with great apes and humans) are used to reconstruct both dynamic equilibrium and muscular systems. The reconstruction suggests that the fossils moved the pelvis and shoulders extensively when they walked. The hindlimb muscles (such as adductors, gluteal muscles and calf) are fleshy and not or very little tendinous. As indicated by the Laetoli step prints (belonging to a close and contemporaneous species), the foot is adducted during the walk and the support is internal just before take off. In spite of inevitable approximations, such a reconstruction appears to be particularly helpful to bring out morphological and functional traits of the first hominids which are both close to and different from modern humans.     

Early hominid brain evolution: a new look at old endocasts

Early hominid brain morphology is reassessed from endocasts of Australopithecus africanus and three species of Paranthropus, and new endocast reconstructions and cranial capacities are reported for four key specimens from the Paranthropus clade. The brain morphology of Australopithecus africanus appears more human like than that of Paranthropus in terms of overall frontal and temporal lobe shape. These new data do not support the proposal that increased encephalization is a shared feature between Paranthropus and early Homo. Our findings are consistent with the hypothesis that Australopithecus africanus could have been ancestral to Homo, and have implications for assessing the tempo and mode of early hominid neurological and cognitive evolution.