Comparison of cranial ontogenetic trajectories among great apes and humans

Molecular data suggest that humans are more closely related to chimpanzees than either is to the gorillas, yet one finds the closest similarity in craniofacial morphology to be among the great apes to the exclusion of humans. To clarify how and when these differences arise in ontogeny, we studied ontogenetic trajectories for Homo sapiens, Pan paniscus, Pan troglodytes, Gorilla gorilla and Pongo pygmaeus. A total of 96 traditional three-dimensional landmarks and semilandmarks on the face and cranial base were collected on 268 adult and sub-adult crania for a geometric morphometric analysis. The ontogenetic trajectories are compared by various techniques, including a new method, relative warps in size-shape space. We find that adult Homo sapiens specimens are clearly separated from the great apes in shape space and size-shape space. Around birth, Homo sapiens infants are already markedly different from the great apes, which overlap at this age but diverge among themselves postnatally. The results suggest that the small genetic differences between Homo and Pan affect early human ontogeny to induce the distinct adult human craniofacial morphology. Pure heterochrony does not sufficiently explain the human craniofacial morphology nor the differences among the African apes.     

Identifying metameric variation in extant hominoid and fossil hominid mandibular molars

Landmark data were collected from cross sections and occlusal images of mandibular molar crowns, and Euclidean distance matrix analysis (EDMA) was used to identify metameric morphological variation between the first and second mandibular molars of living taxa: Gorilla gorilla (n = 30), Pan troglodytes (n = 34), and Homo sapiens (n = 26). Two patterns of metameric variation were identified, one unique to humans and the other shared by chimpanzees and gorillas. In order to assess the utility of this type of analysis for the interpretation of the hominid fossil record, 19 mandibular molars from Sterkfontein Member 4, South Africa, were examined. The pattern of metameric variation of the Sterkfontein molars resembled that of the African great apes, and differed from the modern human pattern. These results demonstrate that data on metameric variation may provide information regarding function or developmental processes previously indiscernible from fossil material.     

The articular surface of the temporal bone in certain fossil hominoids

Although quantitative variations exist between living Man ( Homo sapiens sapiens ) and the extant great apes ( Pongo, Pan, Gorilla ) in such features of the articular surface of the temporal bone (a part of the temporomandibular joint) as the proportionate development of the postglenoid tubercle, the relative prominence of the articular tubercle and the slope of its posterior face, these do not individually effect a clear differentiation between the four extant genera. But in multivariate combination of these features, although Pan and Pongo are relatively closely associated, Gorilla and Homo sapiens sapiens are distinct, and also clearly differentiated from each other. The differences between genera of extant apes are, on average, as great as those between extant Man and individual apes.
As portrayed by such multivariate compound, this anatomical region in four fossil groups displays a unique configuration differentiating Homo sapiens neanderthalensis, Homo erectus pekinensis, Australopithecus africanus and Australopithecus robustus both from one another and from extant types. The differences are such that the fossil species lie uniquely and not intermediate between extant groups.
Definable age changes in this multivariate compound occur in both Man and apes but neither these, nor overall differences between adults, appear to be associated with marked contrasts in the pattern of jaw movement. It would thus seem improbable that inferences can be made from these features about the type of jaw movement that characterized the several fossil groups.     

African ape ancestry

It is commonly believed that the australopithecines are more closely related to humans than to African apes. This view is hardly compatible with the biomolecular data which place theHomo/Pan split at the beginning of the australopithecine period. Nothing in the fossil hominid morphology precludes the possibility that some australopithecines were ancestral to gorillas or chimpanzees and others to humans.     

Cortical bone distribution in the femoral neck of hominoids: Implications for the locomotion of Australopithecus afarensis

Contiguous high resolution computed tomography images were obtained at a 1.5 mm slice thickness perpendicular to the neck axis from the base of the femoral head to the trochanteric line in a sample of 10 specimens each of Homo sapiens, Pan troglodytes, and Gorilla gorilla, plus five specimens of Pan paniscus. Superior, inferior, anterior, and posterior cortical thicknesses were automatically measured directly from these digital images. Throughout the femoral neck H. sapiens displays thin superior cortical bone and inferior cortical bone that thickens distally. In marked contrast, cortical bone in the femoral neck of African apes is more uniformly thick in all directions, with even greater thickening of the superior cortical bone distally. Because the femoral neck acts as a cantilevered beam, its anchorage at the neck-shaft junction is subjected to the highest bending stresses and is the most biomechanically relevant region to inspect for response to strain. As evinced by A.L. 128-1, A.L. 211-1 and MAK-VP-1/1, Australopithecus afarensis is indistinguishable from H. sapiens, but markedly different from African apes in cortical bone distribution at the femoral neck-shaft junction. Cortical distribution in the African ape indicates much greater variation in loading conditions consistent with their more varied locomotor repertoire. Cortical distribution in hominids is a response to the more stereotypic loading pattern imposed by habitual bipedality, and thin superior cortex in A. afarensis confirms the absence of a significant arboreal component in its locomotor repertoire. Am J Phys Anthropol 104:117–131, 1997. © 1997 Wiley-Liss, Inc.     

Moment coefficients of skewness in the femoral neck cortical bone distribution of BAR 1002’00

The cortical bone distributions in the femoral necks of apes and humans differ as a result of different loading environments caused by the realignment of the hip abductor apparatus. Femoral neck cortical bone in extant humans is very thin superiorly and thicker inferiorly, while the cortical bone in apes tends to be more uniformly thick. The unique internal anatomy of extant humans allows inferences to be made about primary locomotor function from incomplete femora. Here the differences in cortical bone distributions are quantified using moment coefficient of skewness. Skewness coefficients at two locations along the neck of the 6 million years old African femoral specimen BAR 1002’00 were compared to samples of 9 extant adult humans and 10 adult chimpanzees. The skewness coefficients of cortical bone in the femoral neck of BAR 1002’00 are more similar to those of chimpanzees than to humans, although the contrast is less pronounced in the region closer to the neck-shaft junction than more proximally toward the femoral head; this pattern indicates that in at least one respect this specimen attributed to Orrorin tugenensis manifests structural features suggesting influences of a hip abductor apparatus that had not yet evolved to the same extent as in extant humans.     

Postnatal temporal bone ontogeny in Pan,Gorilla, and Homo,and the implications for temporal bone ontogeny in Australopithecus afarensis

Assessments of temporal bone morphology have played an important role in taxonomic and phylogenetic evaluations of fossil taxa, and recent three‐dimensional analyses of this region have supported the utility of the temporal bone for testing taxonomic and phylogenetic hypotheses. But while clinical analyses have examined aspects of temporal bone ontogeny in humans, the ontogeny of the temporal bone in non‐human taxa is less well documented. This study examines ontogenetic allometry of the temporal bone in order to address several research questions related to the pattern and trajectory of temporal bone shape change during ontogeny in the African apes and humans. We further apply these data to a preliminary analysis of temporal bone ontogeny in Australopithecus afarensis. Three‐dimensional landmarks were digitized on an ontogenetic series of specimens of Homo sapiens, Pan troglodytes, Pan paniscus, and Gorilla gorilla. Data were analyzed using geometric morphometric methods, and shape changes throughout ontogeny in relation to size were compared. Results of these analyses indicate that, despite broadly similar patterns, African apes and humans show marked differences in development of the mandibular fossa and tympanic portions of the temporal bone. These findings indicate divergent, rather than parallel, postnatal ontogenetic allometric trajectories for temporal bone shape in these taxa. The pattern of temporal bone shape change with size exhibited by A. afarensis showed some affinities to that of humans, but was most similar to extant African apes, particularly Gorilla. Am J Phys Anthropol 151:630–642, 2013. © 2013 Wiley Periodicals, Inc.     

Effects of size and locomotor adaptations on the hominid pelvis: evaluation of australopithecine bipedality with a new multivariate method

Three pelves and eight innominate bones belonging to the fossil species, Australopithecus africanus, Australopithecus robustus, Homo erectus, and Homo sapiens, have been studied biometrically and compared with those of recent humans and apes. A new method of logarithmic factorial analysis suppresses both the size effects and the size reference on pelvic proportions. In combination with principal component analysis it allows specializations to be dissociated from allometrical variations. Some morphological differences on the hominid pelvis prove to be mainly allometric. However, the pelvic morphology of australopithecines is clearly differentiated from that of the genus Homo (including H. erectus, OH 28, KNMER 3227). A. africanus (Sts 14, MLD 7, AL 288) is nearer the humans than is A. robustus (SK 50, SK 3155), which appears to be more specialized in the australopithecine lineage. The pelvic morphology of A. africanus, as integrated with the articular pelvic-femoral link, appears to be biometrically equivalent to that of humans.     

The anomalous archaic Homo femur from Berg Aukas, Namibia: a biomechanical assessment.

The probably Middle Pleistocene human femur from Berg Aukas, Namibia, when oriented anatomically and analyzed biomechanically, presents an unusual combination of morphological features compared to other Pleistocene Homo femora. Its midshaft diaphyseal shape is similar to most other archaic Homo, but its subtrochanteric shape aligns it most closely with earlier equatorial Homo femora. It has an unusually low neck shaft angle. Its relative femoral head size is matched only by Neandertals with stocky hyperarctic body proportions. Its diaphyseal robusticity is modest for a Neandertal, but reasonable compared to equatorial archaic Homo femora. Its gluteal tuberosity is relatively small. Given its derivation from a warm climatic region, it is best interpreted as having had relatively linear body proportions (affecting proximal diaphyseal proportions, shaft robusticity, and gluteal tuberosity size) combined with an elevated level of lower limb loading during development (affecting femoral head size and neck shaft angle).     

Cranial base morphology and temporal bone pneumatization in Asian Homo erectus

The external morphological features of the temporal bone are used frequently to determine taxonomic affinities of fossils of the genus Homo. Temporal bone pneumatization has been widely studied in great apes and in early hominids. However, this feature is rarely examined in the later hominids, particularly in Asian Homo erectus. We provide a comparative morphological and quantitative analysis of Asian Homo erectus from the sites of Ngandong, Sambungmacan, and Zhoukoudian, and of Neandertals and anatomically modern Homo sapiens in order to discuss causes and modalities of temporal bone pneumatization during hominid evolution. The evolution of temporal bone pneumatization in the genus Homo is more complex than previously described. Indeed, the Zhoukoudian fossils have a unique pattern of temporal bone pneumatization, whereas Ngandong and Sambungmacan fossils, as well as the Neandertals, more closely resemble the modern human pattern. Moreover, these Chinese fossils are characterized by a wide midvault and a relatively narrow occipital bone. Our results support the point of view that cell development does not play an active role in determining cranial base morphology. Instead, pneumatization is related to available space and to temporal bone morphology, and its development is related to correlated morphology and the relative disposition of the bones and cerebral lobes. Because variation in pneumatization is extensive within the same species, the phyletic implications of pneumatization are limited in the taxa considered here.     

云南丽江古人类股骨的形态结构

1960年,在云南省丽江市发现了三根古人类股骨,通过地层观察,仅PA108可归为更新世晚期。前人对PA108做了初步报导,为了进一步了解丽江人股骨的演化分类地位和东亚早期现代人股骨形态变异,本文对PA108的内外结构进行了详尽的分析。研究发现,PA108具有明显的早期现代人特征,即明显的股骨粗线、骨干中部后侧骨密质最厚和中部横断面轮廓形状偏椭圆。PA108标本也有一定的特殊性,体现在骨干中近端和中部骨密质厚度分布上,这可能与其股骨嵴发育较弱有关,这一特征也导致了PA108与其他东亚早期现代人之间的形态差异,这些形态变异进一步扩大了目前已知的东亚地区早期现代人变异范围。同时,在采用骨密质厚度分布模式进行分类时,建议关注股骨骨干中部骨密质最厚部位。     

What's Your Angle? Size Correction and Bar–Glenoid Orientation in “Lucy” (A.L. 288-1)

There has been much debate as to the locomotor repertoire of Lucy (A.L. 288-1) and other specimens of Australopithecus afarensis, ranging from fully committed bipeds to species that spent a significant time in the trees as well as on the ground. We examined the bar–glenoid angle, a character purported to indicate arboreal propensities, and its implications for this specific debate and the more general challenge of extracting behavioral information from fossils. We examined the bar–glenoid angle in ontogenetic samples of Pan paniscus, Pan troglodytes, Gorilla gorilla gorilla, Gorilla gorilla beringei, Pongo pygmaeus, Homo sapiens, and A.L. 288-1 (Lucy). We found that there is no allometry in the bar–glenoid angle for the great apes, but a weak correlation for humans. Moreover, the data scatters for the African apes and humans converge at the smaller size ranges, and Lucy's value for bar–glenoid angle falls precisely in this area of overlap. Therefore, we conclude that the bar–glenoid angle is not tightly correlated with function and, as such, cannot be used as a morphological signal of arboreal behavior, especially in the smaller size ranges, at which arboreal and nonarboreal species overlap. Our work does not resolve issues concerning Lucy's precise locomotor repertoire but adds new information to consider. The total morphological pattern, plus an appreciation of the underlying variance in morphological and behavioral characters in extant species, is key for making functional inferences from the morphology of fossils.     

Electromyography of pronators and supinators in great apes.

We obtained electromyographic recordings from the supinator, biceps brachii, pronator quadratus, and pronator teres muscles of a chimpanzee and a gorilla and from the supinator, pronator quadratus, and biceps brachii muscles of an orangutan as they stood and walked quadrupedally on horizontal and inclined surfaces, engaged in suspensory behavior, reached overhead, and manipulated a variety of foods and artifacts. In Pan troglodytes and Pan gorilla, as in Homo sapiens, the supinator muscle is the prime supinator, with the biceps brachii muscle serving to augment speed or force of supination. Primary of the pronator quadratus muscle over the pronator teres muscle during pronation is less clear in the African apes than in humans. Possibly, pongid radial curvature or forelimb elongation or both factors are related to the somewhat different patterns of activity that we observed in the pronator muscles of Pan versus those reported for Homo sapiens. In Pongo pygmaeus, as in P. troglodytes and P. gorilla, the pronator quadratus muscle acts as a pronator and the supinator muscle acts to supinate the hand at the radioulnar joints. The biceps brachii muscle is active at low levels as the orangutan supinates its hand with the elbow flexed.     

Brief Communication: Shape analysis of the MT 1 proximal articular surface in fossil hominins and shod and unshod Homo

As a follow-up study to Proctor et al. (Am J Phys Anthropol 135 (2008) 216-224), this study quantifies the first metatarsal proximal articular surface using three-dimensional morphometrics to test for differences in articular surface shape between habitually shod and habitually unshod humans. In addition, differences in shape between Homo, Pan, Gorilla, and Hylobates are compared to the fossil hominin specimens A. L. 333-54, Stw 562, Stw 573 ("Little Foot"), OH 8, SKX 5017, and SK 1813. No difference in surface shape was found between habitually shod and habitually unshod humans. There is a clear quantitative division in articular surface shape between humans and apes that is more pronounced than a previous study by Proctor et al. (Am J Phys Anthropol 135 (2008) 216-224), due to additional landmarks present in this study. The specimen OH 8 is indistinguishable from modern Homo. The fossils A. L. 333-54, Stw 562, and Stw 573 are intermediate in shape between humans and apes. The specimens SKX 5017 and SK 1813 have a more apelike articular surface. When combined with other characteristics, this trait suggests that Paranthropus used a degree of abduction during locomotion that was much less than that in extant apes, but greater than that in Australopithecus, allowing for some small degree of grasping ability.     

Head posture in the Hominoidea

The angle between the antero-posterior plane of the occipital condyles and a vertical axis at right angles to the Frankfort Horizontal was measured in Homo sapiens, Gorilla, Pan, Pongo and casts of two Neanderthal skulls, the Rhodesian skull and three australopithecine skulls. The angle was much greater in adult Homo sapiens and in the Neanderthal and australopithecine casts than in the adult groups of the three apes. In the immature groups, the angle underwent little change with age in Homo sapiens but in Gorilla and Pan the angle decreased markedly during the growth period. These findings can be readily correlated with the habitual bodily posture of each of the extant genera. In Homo sapiens , an upright posture is adopted early in life while in the African apes the young tend to move by brachiation and thus have an habitual posture of the spine closer to the vertical than in the "knuckle walking" adults. The large value of the angle in the Neanderthal casts also correlates well with the now widely held view that this group has a fully upright posture. However, the finding of a relatively low value for the angle in adult Pongo —a brachiator—runs counter to the general thesis that the angle is a direct reflection of overall posture and casts some doubt upon a conclusion that the large value of the angle in the australopithecine fossils necessarily indicates that these creatures stood upright.     

华南化石猩猩前部牙齿釉面横纹与牙冠形成时间研究

釉面横纹的分布与数目可以反映牙齿生长发育的时间和速率变化, 在化石研究中能为复原个体生活史提供重要依据。本研究运用扫描电子显微镜观察华南化石猩猩门齿、犬齿釉面横纹分布与数目, 并估算门齿和犬齿牙冠形成时间, 结果如下: 牙冠从牙尖至牙颈方向釉面横纹分布密度有疏密变化, 牙尖釉面横纹密度小于10条/mm, 中间至牙颈釉面横纹密度较尖部增大, 大约10-15条/mm; 犬齿釉面横纹数目多于门齿, 雄性犬齿釉面横纹数目多于雌性; 根据釉面横纹计数及其生长周期的组织切片观察结果, 估算门齿牙冠形成时间大约为2.97-6.66年, 犬齿雄性长于雌性, 分别为6.25-11.31年和4.28-7.29年。与一些古猿、早期人类、现代人以及现生大猿比较, 华南化石猩猩釉面横纹整体密度稍大于南方古猿和傍人, 小于黑猩猩、大猩猩、现代人和禄丰古猿; 除侧门齿外, 华南化石猩猩釉面横纹数目明显多于南方古猿、傍人和现代人, 与大猩猩接近; 华南猩猩前部牙齿牙冠形成时间与现生大猿、禄丰古猿差别不大, 与现生猩猩最相近, 长于南方古猿和傍人。     

Growth changes in internal and craniofacial flexion measurements.

Growth changes in both internal and craniofacial flexion angles are presented for Pan troglodytes, Gorilla gorilla, and modern humans. The internal flexion angle (IFA) was measured from lateral radiographs, and the craniofacial flexion angle (CFA) was calculated from coordinate data. Stage of dental development is used as a baseline for examination of growth changes and nonparametric correlations between flexion angles and dental development stage are tested for significance. In Gorilla, the IFA increases during growth. The IFA is relatively stable in Pan and modern humans. Pan and Gorilla display an increase in the CFA. However, this angle decreases during growth in modern humans. Flexion angles were derived from coordinate data collected for several early hominid crania. Measurements for two robust australopithecine crania indicate strong internal flexion. It has been suggested that cerebellar expansion in this group may relate to derived features of the posterior cranial base. In general, australopithecine crania exhibit craniofacial flexion intermediate between great apes and modern humans. The "archaic" Homo sapiens specimen from Kabwe is most similar to modern humans.     

Quantitative three-dimensional shape analysis of the proximal hallucial metatarsal articular surface in Homo, Pan, Gorilla, and Hylobates

Multidimensional morphometrics is used to compare the proximal articular surface of the first metatarsal between Homo, Pan, Gorilla, Hylobates, and the hominin fossils A.L. 333-54 (A. afarensis), SKX 5017 (P. robustus), and OH 8 (H. habilis). Statistically significant differences in articular surface morphology exist between H. sapiens and the apes, and between ape groups. Ape groups are characterized by greater surface depth, an obliquely curved articular surface through the dorso-lateral and medio-plantar regions, and a wider medio-lateral surface relative to the dorso-plantar height. The OH 8 articular surface is indistinguishable from H. sapiens, while A.L. 333-54 and SKX 5017 more closely resemble the apes. P. robustus and A. afarensis exhibit ape-like oblique curvature of the articular surface.     

The taxonomic implications of cranial shape variation in Homo erectus

The taxonomic status of Homo erectus sensu lato has been a source of debate since the early 1980s, when a series of publications suggested that the early African fossils may represent a separate species, H. ergaster. To gain further resolution regarding this debate, 3D geometric morphometric data were used to quantify overall shape variation in the cranial vault within H. erectus using a new metric, the sum of squared pairwise Procrustes distances (SSD). Bootstrapping methods were used to compare the H. erectus SSD to a broad range of human and nonhuman primate samples in order to ascertain whether variation in H. erectus most clearly resembles that seen in one or more species. The reference taxa included relevant phylogenetic, ecological, and temporal analogs including humans, apes, and both extant and extinct papionin monkeys. The mean cranial shapes of different temporogeographic subsets of H. erectus fossils were then tested for significance using exact randomization tests and compared to the distances between regional groups of modern humans and subspecies/species of the ape and papionin monkey taxa. To gauge the influence of sexual dimorphism on levels of variation, comparisons were also made between the mean cranial shapes of single-sex samples for the reference taxa. Results indicate that variation in H. erectus is most comparable to single species of papionin monkeys and the genus Pan, which included two species. However, H. erectus encompasses a limited range of variation given its extensive geographic and temporal range, leading to the conclusion that only one species should be recognized. In addition, there are significant differences between the African/Georgian and Asian H. erectus samples, but not between H. ergaster (Georgia+Africa, excluding OH 9 and Daka) and H. erectus sensu stricto. This finding is in line with expectations for intraspecific variation in a long-lived species with a wide, but probably discontinuous, geographic distribution.