A complete second metatarsal (StW 89) from Sterkfontein Member 4, South Africa

The functional anatomy of the hominin foot has played a crucial role in studies of locomotor evolution in human ancestors and extinct relatives. However, foot fossils are rare, often isolated, and fragmentary. Here, we describe a complete hominin second metatarsal (StW 89) from the 2.0-2.6 million year old deposits of Member 4, Sterkfontein Cave, South Africa. Like many other fossil foot bones, it displays a mosaic of derived human-like features and primitive ape-like features. StW 89 possesses a domed metatarsal head with a prominent sulcus, indicating dorsiflexion at the metatarsophalangeal joint during bipedal walking. However, while the range of motion at the metatarsophalangeal joint is human-like in dorsiflexion, it is ape-like in plantarflexion. Furthermore, StW 89 possesses internal torsion of the head, an anatomy decidedly unlike that found in humans today. Unlike other hominin second metatarsals, StW 89 has a dorsoplantarly gracile base, perhaps suggesting more midfoot laxity. In these latter two anatomies, the StW 89 second metatarsal is quite similar to the recently described second metatarsal of the partial foot from Burtele, Ethiopia. We interpret this combination of anatomies as evidence for a low medial longitudinal arch in a foot engaged in both bipedal locomotion, but also some degree of pedal, and perhaps even hallucal, grasping. Additional fossil evidence will be required to determine if differences between this bone and other second metatarsals from Sterkfontein reflect normal variation in an evolving lineage, or taxonomic diversity.     

A hominoid distal tibia from the Miocene of Pakistan

A distal tibia, YGSP 1656, from the early Late Miocene portion of the Chinji Formation in Pakistan is described. The fossil is 11.4 million years old and is one of only six postcranial elements now assigned to Sivapithecus indicus. Aspects of the articular surface are cercopithecoid-like, suggesting some pronograde locomotor activities. However, YGSP 1656 possesses an anteroposteriorly compressed metaphysis and a mediolaterally thick medial malleolus, ape-like features functionally related to orthograde body postures and vertical climbing. YGSP 1656 lacks specializations found in the ankle of terrestrial cercopithecoids and thus Sivapithecus may have been primarily arboreal. Nevertheless, the morphology of this tibia is unique, consistent with other interpretations of Sivapithecus postcranial functional morphology that suggest the locomotion of this ape lacks a modern analog. Based on the limited postcranial remains from S. indicus, we hypothesize that this taxon exhibited substantial body size dimorphism.     

Little Foot and big thoughts—a re-evaluation of the Stw573 foot from Sterkfontein, South Africa

The part of the fossil assemblage Stw573 consisting of some medial foot bones was initially reported by Clarke & Tobias (Science 269 (2002) 521). They found it to have both ape- and human-like qualities, being human-like proximally and ape-like distally. We have undertaken a re-examination of this pedal assemblage using a multivariate analysis; while we also found ape- and human-like qualities, they are in direct conflict with the original findings of Clarke and Tobias. We report an essentially ape-like morphology proximally and a human-like morphology distally; the talus and navicular were found to be ape-like and the medial cuneiform human-like. We also undertook a morphometric analysis of the medial cuneiform from the fossil assemblage OH8, as this was not included in the original OH8 study of Kidd et al (J Hum Evol 31 (1996) 269); this cuneiform was found to have a human-like morphology. Thus, the medial column findings from the two assemblages are very similar. This finding, coupled with the re-evaluation of the stratigraphy at Sterkfontein (Am J Phys Anthrop 119 (2002) 192), suggests that the two may have been contemporaneous.

We also note that three broad patterns of modification have been identified, equating to proximal–distal lateral–medial (cranio–caudal) and dorsal–plantar (posterior–anterior). It has not escaped our notice that these patterns are each controlled by specific genes or growth factors; we thus see a morphometric expression of our developmental past.     

化石人类脑演化研究概况

脑演化是人类演化的一个重要组成部分,其研究可以为人类起源、演化、人群关系及语言、智力等方面提供重要的信息。脑演化的主要证据是通过研究颅内模（endocast）及颅骨的形态得到的。颅内模是从颅骨内表面得到的脑的外部形态。有时颅骨的内腔充满泥沙,并且为钙质所结固,可以自然形成颅内模。也可以人工制作颅内模。颅内模和颅骨的内表面能够提供人类脑髓及神经进化方面的直接证据。对化石人类脑演化的研究主要包括以下几个方面的内容：测量或估计脑量的大小及其和身体大小之间的关系;研究脑量随时间的变化过程;通过对早期人类颅内模表面沟回形态特征的研究,探索脑功能区在早期人类和猿类的区别及在演化上的变化;左右大脑不对称性与一侧优势关系,探讨语言的起源和惯用手的脑功能基础等;脑膜中动脉系统、静脉窦系统及与血液循环相关的排泄孔的变化,探讨大脑各部分比例的变化和功能的日益复杂对供血需求的影响;通过对人类脑演化的研究,探讨人类进化的原因。本文通过对以上几个方面及其中国化石脑演化研究的介绍,对化石人类脑演化的研究概况作综合论述和简单回顾。     

The biological and chronological maturation of early hominids

Recent studies on the rate and pattern of dental development indicate that the growth and maturation of early hominids were more similar to the extant apes than to modern humans. This contrasts with the previously held opinion derived from combined dental development, pattern and attrition studies claiming that early hominids were more hominine in their development (Mann, 1975). This paper explores the origin of this difference of opinion and reviews immature hominid dentitions with the benefit of improved radiographs and new data on the pattern and rate of pongid dental development. Paranthropus and Australopithecus specimens are shown to possess an ape-like development pattern but incisor development is specialized in the former and superficially human-like in pattern. The present and recent studies on dental development rate and pattern justify the position that early hominids were more ape-like in their growth and development. Therefore, ages at death calculated from pongid dental development schedules are provided for most immature early hominids. More detailed studies of early hominid developmental biology are now possible. It is suggested that divergent heterochronic processes characterize changes in brain/body proportions during hominid evolution. Relative rates of bone remodeling processes can now be identified on early hominid skeletons. The paleodemographic analysis of early hominids is little changed by the developmental model one chooses.     

Further observations on a method for estimating hominoid dental developmental patterns

Several recent studies have concluded that early hominines resembled apes in patterns of dental ontogeny and have inferred from this conclusion that they were ape-like in other aspects of growth and development as well. Prior to their employment on fossil junveniles, the comparative methods used in these studies were not first tested for their predicitive accuracy in distinguishing modern humans from apes on the basis of proposed patterns of dental ontogeny. We find that 92% of a smaple of 48 modern children are classified as ape-like or nonhuman by the criteria employed in one of these studies and overlap entirely with fossil juvenile dental patterns. The use of these methods to characterize early hominines as “ape-like” or distinctly divergent from humans is thus shown to be unwarranted. © 1993 Wiley-Liss, Inc.     

Crocodylian and mammalian carnivore feeding traces on hominid fossils from FLK 22 and FLK NN 3, Plio-Pleistocene, Olduvai Gorge, Tanzania

Taphonomic analysis of the Olduvai Hominid (OH) 8 left foot from FLK NN Level 3 and the OH 35 left leg from FLK Level 22 (Zinjanthropus level) in Middle Bed I, Olduvai Gorge, indicates that both were fed upon by crocodiles. Both bear extensive tooth marking, including bisected tooth marks diagnostic of crocodylian feeding. The location of the bisected tooth marks on the distal tibia and the talus indicates disarticulation of the foot by crocodiles. The broken proximal ends of the tibia and fibula are more typical of feeding by a leopard-like carnivore, as is damage to the OH 7 mandible and parietals that are associated with and may derive from the same individual as OH 8. Previous work showing a close articulation of the foot and the leg has been used to suggest that the two specimens belong to the same individual despite deriving from sites separated by 200 m and slightly different stratigraphic levels according to previous work. The location and agent of tooth marking and the nature of gross damage do not refute this hypothesis, but the punctures on the talus and distal tibia differ in size and sharpness. Recent work shows that the stratigraphic discrepancy between OH 8 and OH 35 is greater than previously thought, refuting the single-individual hypothesis. Although seemingly unlikely, this denotes that two hominids represented by rarely found leg and foot elements both lost their left foot to crocodiles at nearby sites within a 6,000 year interval. We cannot determine if the hominids were preyed upon by crocodiles or mammalian carnivores. However, the carnivore damage to them and associated faunal remains suggests that high predation risk constrained hominid activities involving discard of the stone artifacts found at these sites. This finding is inconsistent with the interpretation of the sites as home bases or living floors.     

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.     

The rise of the hominids as an adaptive shift in fallback foods: plant underground storage organs (USOs) and australopith origins

We propose that a key change in the evolution of hominids from the last common ancestor shared with chimpanzees was the substitution of plant underground storage organs (USOs) for herbaceous vegetation as fallback foods. Four kinds of evidence support this hypothesis: (1) dental and masticatory adaptations of hominids in comparison with the African apes; (2) changes in australopith dentition in the fossil record; (3) paleoecological evidence for the expansion of USO-rich habitats in the late Miocene; and (4) the co-occurrence of hominid fossils with root-eating rodents. We suggest that some of the patterning in the early hominid fossil record, such as the existence of gracile and robust australopiths, may be understood in reference to this adaptive shift in the use of fallback foods. Our hypothesis implicates fallback foods as a critical limiting factor with far-reaching evolutionary effects. This complements the more common focus on adaptations to preferred foods, such as fruit and meat, in hominid evolution.     

Phylogeny of early Australopithecus: new fossil evidence from the Woranso-Mille (central Afar,Ethiopia)

The earliest evidence of Australopithecus goes back to ca 4.2 Ma with the first recorded appearance of Australopithecus ‘anamensis’ at Kanapoi, Kenya. Australopithecus afarensis is well documented between 3.6 and 3.0 Ma mainly from deposits at Laetoli (Tanzania) and Hadar (Ethiopia). The phylogenetic relationship of these two ‘species’ is hypothesized as ancestor–descendant. However, the lack of fossil evidence from the time between 3.6 and 3.9 Ma has been one of its weakest points. Recent fieldwork in the Woranso-Mille study area in the Afar region of Ethiopia has yielded fossil hominids dated between 3.6 and 3.8 Ma. These new fossils play a significant role in testing the proposed relationship between Au. anamensis and Au. afarensis. The Woranso-Mille hominids (3.6–3.8 Ma) show a mosaic of primitive, predominantly Au. anamensis-like, and some derived (Au. afarensis-like) dentognathic features. Furthermore, they show that, as currently known, there are no discrete and functionally significant anatomical differences between Au. anamensis and Au. afarensis. Based on the currently available evidence, it appears that there is no compelling evidence to falsify the hypothesis of ‘chronospecies pair’ or ancestor–descendant relationship between Au. anamensis and Au. afarensis. Most importantly, however, the temporally and morphologically intermediate Woranso-Mille hominids indicate that the species names Au. afarensis and Au. anamensis do not refer to two real species, but rather to earlier and later representatives of a single phyletically evolving lineage. However, if retaining these two names is necessary for communication purposes, the Woranso-Mille hominids are best referred to as Au. anamensis based on new dentognathic evidence.     

Ontogeny and phylogeny of the pelvis in Gorilla, Pongo, Pan, Australopithecus and Homo

To examine the evolutionary differences between hominoid locomotor systems, a number of observations concerning the growth of the pelvis among the great apes as compared to modern and fossil hominids are reported. We are interested in the size and shape of the coxal bones at different developmental stages across species that may elucidate the relationship between ontogeny and phylogeny (i.e., heterochrony) in the hominoid pelvis. Our hypotheses are: (1) do rates of absolute growth differ?, (2) do rates of relative growth differ?, and (3) does heterochrony explain these differences? Bivariate and multivariate analyses of pelvic dimensions demonstrate both the diversity of species-specific ontogenetic patterns among hominoids, and an unequivocal separation of hominids and the great apes. Heterochrony alone fails to account for the ontogenetic differences between hominids and the great apes. Compared to recent Homo,Australopithecus can be described as 'hyper-human' from the relative size of the ischium, and short but broad ilium. Australopithecus afarensis differs from Australopithecus africanus by its relatively long pubis. In multivariate analyses of ilium shape, the most complete coxal bone attributed to Homo erectus, KNM-ER 3228, falls within the range of juvenile and adult Australopithecus, whereas Broken Hill falls within the range of modern Homo, suggesting that the modern human ilium shape arose rather recently. Among the great apes, patterns of pelvic ontogeny do not exclusively separate the African apes from Pongo.     

Axial and appendicular skeleton of Homo antecessor.

The human trunk and limb bones recovered from the Gran Dolina site, in the Sierra de Atapuerca (Burgos, Spain) are studied. All these fossils were excavated at the level called TD6 between 1994 and 1995 and have been dated in excess of 780,000 years ago. These remains have been recently attributed to a new Homo species named Homo antecessor. Axial (vertebrae and ribs) and part of the appendicular (clavicles, radii, femur and patellae) skeleton are studied here. Hand and foot bones have been studied elsewhere (Lorenzo et al., 1999). Four is the minimum number of individuals represented by the postcranial remains recovered up to now. All elements are briefly described anatomically, measured and compared with other fossil hominids and modern humans in order to establish, as far as possible, what postcranial morphology characterized this new species of our genus.The H. antecessor postcrania, generally, display a set of morphological traits that are more similar to modern humans than to the Middle and Upper Pleistocene European hominids. Our results do not contradict the previous phylogenetic analysis, i.e., that H. antecessor represents the last common ancestor for H. sapiens (modern humans) and H. neanderthalensis (Neandertals).     

Footprint Clues in Hominid Evolution and Forensics: Lessons and Limitations

In 1978 and 1979 at Laetoli, Northern, Tanzania, Mary D. Leakey and assistants excavated the most compelling evidence for the existence of bipedalism in Pliocene (3.5 Ma) hominids. They have stimulated controversy over the extent to which the three individuals had feet and gaits that are like those of humans versus having ape-like features and gaits. A short trail of bipedal tracks discovered in 1977 at Laetoli site A are probably not those of a hominid and more closely resemble the prints of bears. One of the original researchers on the Laetoli prints claimed that she could identify individual modern humans from their footprints, partial footprints and shoe prints because each individual has distinctive foot morphology on a par with the individuality of fingerprints. Courts of law should not have allowed her unproven (and now discredited) method to be used forensically prior to thorough review by scientific peers.     

Morphological correlates of the first metacarpal proximal articular surface with manipulative capabilities in apes,humans and South African early hominins

This study quantifies the metacarpal 1 (MC 1) proximal articular surface using three-dimensional morphometrics in extant hominids and fossil hominins (SK 84, cf. Paranthropus robustus/Homo erectus and StW 418, Australopithecus africanus) to understand which characteristics of the proximal metacarpal 1 are potentially correlated with human manipulative abilities and if they can be used in a paleoanthropological setting. A principal components (PC) analysis was used to compare MC 1 proximal articular surface shape and ANOVA and Tukey's HSD post-hoc tests were conducted to determine differences among groups. Homo is significantly different from nonhuman hominids having a less radioulnarly and dorsovolarly curved articular surface. All nonhuman hominids have more curved articular surface with Gorilla showing the most curved joint. Moreover, this study highlights the presence of a radially extended surface in Homo that may be related to the greater thumb abduction in human manipulation activities. Both fossils analyzed show a great ape-like MC 1 proximal articular surface which, associated with recent trabecular and archaeological evidence, may indicate that the ability to make/use stone tools preceded the morphological adaptations associated today with such behavior.     

Paleoanthropology of the Kibish Formation, southern Ethiopia: Introduction

Cranial and skeletal remains of modern humans, Homo sapiens, were discovered in the Kibish Formation in 1967 by a team from the Kenya National Museums directed by Richard Leakey. Omo I, from Kamoya's Hominid Site (KHS), consists of much of a skeleton, including most of the cranial vault, parts of the face and mandible, and many postcranial elements. Omo II, from Paul's Hominid Site (PHS), is a virtually complete calvaria. Only a limited fauna and a few stone artifacts attributed to the Middle Stone Age were recovered in conjunction with the fossil hominids. The available dating techniques suggested a very early age, over 100 ka, for Member I, from which the Omo I and Omo II fossils were recovered. However, in subsequent decades, the reliability of the dates and the provenance of the Kibish hominids were repeatedly questioned. The papers in this volume provide a detailed stratigraphic analysis of the Kibish Formation and a series of new radiometric dates that indicate an age of 196 +/- 2 ka for Member I and 104 +/- 1 for Member III, confirming the antiquity of the lower parts of the Kibish Formation and, in turn, the fossils from Member I. Studies of the postcranial remains of Omo I indicate an overall modern human morphology with a number of primitive features. Studies of an extensive lithic record from Members I and III indicate a Middle Stone Age technology comparable to assemblages of similar age elsewhere in Ethiopia. Studies of the mammalian, avian, and fish faunas indicate overall similarities to those found in the region today, with a few distinctive differences.     

Inferring hominoid and early hominid phylogeny using craniodental characters: the role of fossil taxa

Recent discoveries of new fossil hominid species have been accompanied by several phylogenetic hypotheses. All of these hypotheses are based on a consideration of hominid craniodental morphology. However, Collard and Wood (2000) suggested that cladograms derived from craniodental data are inconsistent with the prevailing hypothesis of ape phylogeny based on molecular data. The implication of their study is that craniodental characters are unreliable indicators of phylogeny in hominoids and fossil hominids but, notably, their analysis did not include extinct species. We report here on a cladistic analysis designed to test whether the inclusion of fossil taxa affects the ability of morphological characters to recover the molecular ape phylogeny. In the process of doing so, the study tests both Collard and Wood's (2000) hypothesis of character reliability, and the several recently proposed hypotheses of early hominid phylogeny. One hundred and ninety-eight craniodental characters were examined, including 109 traits that traditionally have been of interest in prior studies of hominoid and early hominid phylogeny, and 89 craniometric traits that represent size-corrected linear dimensions measured between standard cranial landmarks. The characters were partitioned into two data sets. One set contained all of the characters, and the other omitted the craniometric characters. Six parsimony analyses were performed; each data set was analyzed three times, once using an ingroup that consisted only of extant hominoids, a second time using an ingroup of extant hominoids and extinct early hominids, and a third time excluding Kenyanthropus platyops. Results suggest that the inclusion of fossil taxa can play a significant role in phylogenetic analysis. Analyses that examined only extant taxa produced most parsimonious cladograms that were inconsistent with the ape molecular tree. In contrast, analyses that included fossil hominids were consistent with that tree. This consistency refutes the basis for the hypothesis that craniodental characters are unreliable for reconstructing phylogenetic relationships. Regarding early hominids, the relationships of Sahelanthropus tchadensis and Ardipithecus ramidus were relatively unstable. However, there is tentative support for the hypotheses that S. tchadensis is the sister taxon of all other hominids. There is support for the hypothesis that A. anamensis is the sister taxon of all hominids except S. tchadensis and Ar. ramidus. There is no compelling support for the hypothesis that Kenyanthropus platyops shares especially close affinities with Homo rudolfensis. Rather, K. platyops is nested within the Homo + Paranthropus + Australopithecus africanus clade. If K. platyops is a valid species, these relationships suggest that Homo and Paranthropus are likely to have diverged from other hominids much earlier than previously supposed. There is no support for the hypothesis that A. garhi is either the sister taxon or direct ancestor of the genus Homo. Phylogenetic relationships indicate that Australopithecus is paraphyletic. Thus, A. anamensis and A. garhi should be allocated to new genera.     

Compact bone distribution and biomechanics of early hominid mandibles.

This investigation explores the effects of compact bone distribution on the biomechanical properties of the postcanine mandibular corpus of the fossil hominid taxa Australopithecus africanus and Paranthropus robustus. The mandibles of extant great apes, modern humans, and the fossil hominids are examined by computed tomography (CT), and compact bone contours are used to calculate cross-sectional biomechanical properties (cortical area, second moments of area, and Bredt's formula for torsional strength). The relative amount of compact bone is comparable in the modern and fossil mandibles, but the mechanical properties of A. africanus and P. robustus jaws are distinct in terms of the ratio of minimum to maximum second moments of area. This difference most likely represents a structural response to elevated torsional moments in the fossil hominids. Although the relative amount of compact bone in cross-section does not differ significantly between taxa by statistical criteria, A. africanus utilizes less cortical bone than P. robustus in the same manner in which Pongo is separated from the condition in other extant large-bodied hominoids. It has been suggested that the phenomenon of mandibular "robusticity" (expressed as an index of corpus breadth/corpus height) may be an effect of postcanine megadontia and/or reduced canine size in the australopithecines. Results presented here, however, indicate that it is unlikely that either factor adequately accounts for mandibular size and shape variation in early hominids.     

Comparative myology of the hominoid cranial base. I. The muscular relationships and bony attachments of the digastric muscle

This paper aims to document accurately the soft tissue anatomy and bony attachments of the posterior belly of the digastric muscle and other closely related muscles in the mastoid region of extant hominoids and fossil hominids. Five wet specimens including individuals of Pan, Gorilla and Pongo were dissected and described. Eight casts of fossil hominid cranial bases were also studied along with measurements and notes made from the same original fossil hominid specimens to assess their soft tissue markings in the light of the findings for the three great apes. The results indicate that whereas the attachment of the posterior belly of the digastric muscle in Homo sapiens is associated with a deep groove or fossa, it originates from a widened area and leaves no bony markings on the cranial base of the three great apes. Following a change in the position of the foramen magnum and the occipital condyles in hominids and H. sapiens the insertion of the posterior belly of the digastric has remained posteriorly positioned but has become compressed into a deep groove. It is likely that this has come about by the displacement of the more medial soft tissue structures which have been moved laterally away from the occipital condyles.