Body weight prediction in early fossil hominids: Towards a taxon-“independent” approach

The choice of a model taxon is crucial when investigating fossil hominids that clearly do not resemble any extant species (such as Australopithecus) or show significant differences from modern human proportions (such as Homo habilis OH 62). An “interhominoid” combination is not adequate either, as scaling with body weight is strongly divergent in African apes and humans for most skeletal predictors investigated here. Therefore, in relation to a study of seven long bone dimensions, a new taxon-“independent” approach is suggested. For a given predictor, its taxonomic “independence” is restricted to the size range over which the body weight-predictor relationship for African apes and humans converges. Different predictors produce converging body weight estimates (BWEs) for different size ranges: taxon-“independent” estimates can be calculated for small- and medium-sized hominids (e. g., for weights below 50 kg) using femoral and tibial dimensions, whereas upper limb bones provide converging results for large hominids (above 50 kg). If the remains of Australopithecus afarensis really belong to one species, the relationship of male (above 60 kg) to female body weight (approximately 30 kg) does not fall within the observed range of modern hominoids. Considering Sts 14 (22 kg) to represent a small-sized Australopithecus africanus, the level of encephalization lies well above that of extant apes. If OH 62 (approximately 25 kg), with limb proportions less human-like than those of australopithecines, indeed represents Homo habilis (which has been questioned previously), an increase in relative brain size would have occurred well before full bipedality, an assumption running counter to current assumptions concerning early human evolution. © 1993 Wiley-Liss, Inc.     

Model of a hypothetical protohominid dentition and its implications for hominid phylogeny

The complete dentition of the common ancestor ofAustralopithecus andHomo, intermediate between that of a pongid and a hominid, is virtually unknown. The maxillary dentition (P3-M2) ofRamapithecus brevirostris Lewis, 1934, a pongid from the Early Pliocene, and that of hominids from the Late Pliocene and Plio/Pleistocene is known. SinceR. brevirostris is probably ancestral to the hominids, a model of intermediate maxillary dentition (P3-M2) is extrapolated and described. The model represents a hypothetical protohominid dentition. It does not conform with the teeth ofAustralopithecus, but shows greater morphological affinity to hominine dentition and to 5 myo hominids. TheHomo lineage, therefore, may go back to the Middle Pliocene. According to the normal sequence of evolution, it is most unlikely thatAustralopithecus gave rise toHomo, but much more probable that a very early, generalizedHomo evolved into an advanced, specializedAustralopithecus.     

Relative cheek-tooth size in Australopithecus

Until the discovery of Australopithecus afarensis, cheek-tooth megadontia was unequivocally one of the defining characteristics of the australopithecine grade in human evolution along with bipedalism and small brains. This species, however, has an average postcanine area of 757 mm², which is more like Homo habilis (759 mm²) than A. africanus (856 mm²). But what is its relative cheek-tooth size in comparison to body size? One approach to this question is to compare postcanine tooth area to estimated body weight. By this method all Australopithecus species are megadont: they have cheek teeth 1.7 to 2.3 times larger than modern hominoids of similar body size. The series from A. afarensis to A. africanus to A. robustus to A. boisei shows strong positive allometry indicating increasing megadontia through time. The series from H. habilis to H. erectus to H. sapiens shows strong negative allometry which implies a sharp reduction in the relative size of the posterior teeth. Postcanine megadontia in Australopithecus species can also be demonstrated by comparing tooth size and body size in associated skeletons: A. afarensis (represented by A.L. 288–1) has a cheek-tooth size 2.8 times larger than expected from modern hominoids; A. africanus (Sts 7) and A. robustus (TM 1517) are over twice the expected size. The evolutionary transition from the megadont condition of Australopithecus to the trend of decreasing megadontia seen in the Homo lineage may have occurred between 3.0 and 2.5 m.y. from A. afarensis to H. habilis but other evidence indicates that it is more likely to have occurred between 2.5 to 2.0 m.y. from an A. africanus-like form to H. habilis.     

The brain of Homo habilis: A new level of organization in cerebral evolution

New studies have been made on endocranial casts of Olduvai specimens of Homo habilis. The results have been compared with those on other East African H. habilis and other hominoids. The mean absolute endocranial capacity of H. habilis is appreciably larger than the mean for australopithecine species: on the new estimates, the H. habilis mean is 45·1% greater than the A. africanus mean and 24·8% greater than that of A. boisei. New data for relative brain size, expressed by Jerison's Nc and EQ and Hemmer's CC, strongly confirm that it was with H. habilis that there appeared the remarkable autapomorphy of Homo, disproportionate expansion of the brain. Encephalometric studies reveal marked transverse expansion of the cerebrum, especially the frontal and parieto-occipital parts, in H. habilis and increased bulk of the frontal and parietal lobes, a derived feature of Homo. There is moderate cerebral heightening, but little or no cerebral lengthening. The sulcal and gyral pattern of the lateral part of the frontal lobe of H. habilis differs from those of Australopithecus and resembles the derived pattern of Homo. The inferior parietal lobule is prominently developed—an autapomorphy of Homo. The two major cerebral areas governing spoken language in modern man are well represented in the endocasts of H. habilis, a functionally important autapomorphy of Homo. The pattern of middle meningeal vessels is more complex with more anastomoses than in australopithecines: H. habilis shares this derived feature with later forms of Homo. In all these features, the brain of H. habilis had made major advances, beyond the more ape-like australopithecine brain. With H. habilis, cerebral evolution had progressed beyond the stage of “animal hominids” (Australopithecus spp.) to that of “human hominids” (Homo spp.). In functional capacity, in particular, its possession of a structural marker of the neurological basis of spoken language, H. habilis had attained a new evolutionary level of organization.     

Revision of fossil hominid jaws from the plio/pleistocene of hadar,in ethiopia including a new species of the genusHomo (hominoidea: homininae)

The assignment of fossil hominoid jaws from the Plio/Pleistocene of Hadar to a single genus,Australopithecus Dart, 1925, is a misnomer. They are morphologically unrestricted to and inconsistent with the diagnosis and evolutionary trend ofAustralopithecus. The morphological pattern of four large jaws is indeed australopithecine and similar toA. africanus Dart, 1925, but six small jaws reveal a pre-habilis stage of dental development early in theHomo lineage. On the basis of their unique hominine dentition, they are reinterpreted as representing a new species,Homo antiquus n.sp.     

The gait of Australopithecus

A biomechanical analysis of the pelvic and femoral samples available for Australopithecus is presented. No feature of these samples was found to distinguish their gait pattern from that of modern man or to differ in the two presently recognized allomorphs of Australopithecus. Morphological differences between Australopithecus and modern man appear to be the result of different degrees of encephalization rather than any difference in locomotor adaptation.     

New evidence of the genus Homo from East Rudolf, Kenya. I

Three new fossil hominid specimens that were recovered in 1970 from the Plio-Pleistocene sediments to the east of Lake Rudolf are described. They include the left side of the body and the symphyseal region of an adult mandible that contains three molar teeth (KNM-ER 730), an edentulous left-sided mandibular fragment (KNM-ER 731) and the shaft of a left femur (KNM-ER 737). The specimens are described in anatomical detail, illustrated and selected measurements given. It is concluded that they should be attributed to the genus Homo sp. indet. Detailed comparative studies will be published in due course.     

Quantitative and qualitative trends in human sapientization

Sapientization is envisaged as a process leading from the earliest representatives of the genus Homo to the shape and dynamism of Homo sapiens (sapiens). Taking into account the manifestation of the changes occurring in the Homo brain-case, two evolutionary trends can be distinguished: the expansion of the cranial capacity (quantitative sapientization) and the attainment of the recent shape (qualitative sapientization). Evidently, both trends cooperate towards a single objective. The writer suggests that they may come into play in an alternating way.The major changes from the psychic and ethological standpoint seem to be related to stages in qualitative change, namely, to the transition both from Australopithecus to Homo and from H. neanderthalensis to Homo sapiens (sapiens).     

Liang Bua Homo floresiensis mandibles and mandibular teeth: a contribution to the comparative morphology of a new hominin species

In 2004, a new hominin species, Homo floresiensis, was described from Late Pleistocene cave deposits at Liang Bua, Flores. H. floresiensis was remarkable for its small body-size, endocranial volume in the chimpanzee range, limb proportions and skeletal robusticity similar to Pliocene Australopithecus, and a skeletal morphology with a distinctive combination of symplesiomorphic, derived, and unique traits. Critics of H. floresiensis as a novel species have argued that the Pleistocene skeletons from Liang Bua either fall within the range of living Australomelanesians, exhibit the attributes of growth disorders found in modern humans, or a combination of both. Here we describe the morphology of the LB1, LB2, and LB6 mandibles and mandibular teeth from Liang Bua. Morphological and metrical comparisons of the mandibles demonstrate that they share a distinctive suite of traits that place them outside both the H. sapiens and H. erectus ranges of variation. While having the derived molar size of later Homo, the symphyseal, corpus, ramus, and premolar morphologies share similarities with both Australopithecus and early Homo. When the mandibles are considered with the existing evidence for cranial and postcranial anatomy, limb proportions, and the functional anatomy of the wrist and shoulder, they are in many respects closer to African early Homo or Australopithecus than to later Homo. Taken together, this evidence suggests that the ancestors of H. floresiensis left Africa before the evolution of H. erectus, as defined by the Dmanisi and East African evidence.     

Pan paniscus and human evolution

Detailed comparisons of the postcranium, cranium, and dentition of Pan paniscus, Pan troglodytes, and Homo reveal that except for slight differences in fore- and hindlimb proportions and the morphology of the shoulder, the postcranium of the two species of Pan are allometrically scaled variants of the same animal and one does not resemble Homo more than the other. Nor does the postcranium of one species of Pan resemble Australopithecus more closely than the other when the effects of body size are controlled. The over all morphological pattern of the skull and teeth of the two chimpanzees is clearly different, however, but both are about equally distinct from the earliest known members of the family Hominidae.     

Dental development and the evolution of life history in Hominidae

Development of the dentition is critically integrated into the life cycle in living mammals. Recent work on dental development has given rise to three separate lines of evidence on the evolution of human growth and aging; these three, based on several independent studies, are reviewed and integrated here. First, comparative study of living primate species demonstrates that measures of development (e.g., age of emergence of the first permanent molar) are highly correlated with the morphological attributes brain and body weight (as highly as r = 0.98, N = 21 species). These data predict that small-bodied, small-brained Australopithecus erupted M₁ at 3–3.5 years and possessed a life span comparable to that of a chimpanzee. Second, chronological age at death for three australopithecines who died at or near emergence of M₁ is now estimated as ～3.25 years based on incremental lines in teeth; this differs substantially from expectations based on human growth schedules (5.5–6 years). Third, developmental sequences (assessed by the coefficient of variation of human dental age) observed in gracile Australopithecus and great apes diverge from those of humans to a comparable degree; sequences become more like modern humans after the appearance of the genus Homo. These three lines of evidence agree that the unique rate and pattern of human life history did not exist at the australopithecine stage of human evolution. It is proposed that the life history of early Homo matched no living model precisely and that growth and aging evolved substantially in the Hominidae during the last 2 million years.     

A SINGLE LINEAGE IN EARLY PLEISTOCENE HOMO: SIZE VARIATION CONTINUITY IN EARLY PLEISTOCENE HOMO CRANIA FROM EAST AFRICA AND GEORGIA

The relationship between Homo habilis and early African Homo erectus has been contentious because H. habilis was hypothesized to be an evolutionary stage between Australopithecus and H. erectus, more than a half‐century ago. Recent work re‐dating key African early Homo localities and the discovery of new fossils in East Africa and Georgia provide the opportunity for a productive re‐evaluation of this topic. Here, we test the hypothesis that the cranial sample from East Africa and Georgia represents a single evolutionary lineage of Homo spanning the approximately 1.9–1.5 Mya time period, consisting of specimens attributed to H. habilis and H. erectus. To address issues of small sample sizes in each time period, and uneven representation of cranial data, we developed a novel nonparametric randomization technique based on the variance in an index of pairwise difference from a broad set of fossil comparisons. We fail to reject the hypothesis of a single lineage this period by identifying a strong, time‐dependent pattern of variation throughout the sequence. These results suggest the need for a reappraisal of fossil evidence from other regions within this time period and highlight the critical nature of the Plio‐Pleistocene boundary for understanding the early evolution of the genus Homo.     

Postcranial robusticity in Homo. I: Temporal trends and mechanical interpretation

Temporal trends in postcranial robusticity within the genus Homo are explored by comparing cross-sectional diaphyseal and articular properties of the femur, and to a more limited extent, the humerus, in samples of Recent and earlier Homo. Using both theoretical mechanical models and empirical observations within Recent humans, scaling relationships between structural properties and bone length are developed. The influence of body shape on these relationships is considered. These scaling factors are then used to standardize structural properties for comparisons with pre-Recent Homo (Homo sp. and H. erectus, archaic H. sapiens, and early modern H. sapiens). Results of the comparisons lead to the following conclusions: 1) There has been a consistent, exponentially increasing decline in diaphyseal robusticity within Homo that has continued from the early Pleistocene through living humans. Early modern H. sapiens are closer in shaft robusticity to archaic H. sapiens than they are to Recent humans. The increase in diaphyseal robusticity in earlier Homo is a result of both medullary contraction and periosteal expansion relative to Recent humans. 2) There has been no similar temporal decline in articular robusticity within Homo–relative femoral head size is similar in all groups and time periods. Thus, articular to shaft proportions are different in pre-Recent and Recent Homo. 3) These findings are most consistent with a mechanical explanation (declining mechanical loading of the postcranium), that acted primarily through developmental rather than genetic means. The environmental (behavioral) factors that brought about the decline in postcranial robusticity in Homo are ultimately linked to increases in brain size and cultural-technological advances, although changes in robusticity lag behind changes in cognitive capabilities. © 1993 Wiley-Liss, Inc.     

Emergence of the genus Homo: From concept to taxonomy

The main goal of this paper is to present an overview of hypotheses concerning early Homo specimens and to discuss the definition of the genus Homo in the light of recent discoveries. For some authors, all the specimens attributed to early Homo belong to one unique species. For others, this group (Homo habilis sensu lato) is heterogeneous and could be splitted into two groups: H. habilis and Homo rudolfensis. Some researchers have also proposed to put the species habilis and rudolfensis into the genera Australopithecus or Kenyanthropus. Therefore, two scenarios concerning first humans seem to emerge. An emergence of the genus Homo, as early as 2.8 Ma, with Homo sp. specimens and the species H. habilis and H. rudolfensis, another at 1.9 Ma with Homo ergaster. According to the recent archaeological and paleoanthropological discoveries, these criteria often considered to be crucial for the definition of the genus Homo, as the cranial capacity, the humanlike manipulative abilities, the habitual erect posture and bipedal gait, the language ability and the capacity to make tools are now obsolete.     

How big were early hominids?

The recent discovery of new postcranial fossils, particularly associated body parts, of several Plio-Pleistocene hominids provides a new opportunity to assess body size in human evolution.¹ Body size plays a central role in the biology of animals because of its relationship to brain size, feeding behavior, habitat preference, social behavior, and much more. Unfortunately, the prediction of body weight from fossils is inherently inaccurate because skeletal size does not reflect body size exactly and because the fossils are from species having body proportions for which there are no analogues among modern species. The approach here is to find the relationship between body size and skeletal size in ape and human specimens of known body weight at death and to apply this knowledge to the hominid fossils, using a variety of statistical methods, knowledge of the associated partial skeletons of the of early hominids, formulae derived from a modern human sample, and, finally, common sense. The following modal weights for males and females emerge: Australopithecus afarensis, 45 and 29 kg; A. africanus, 41 and 30 kg; A. robustus, 40 and 32 kg; A. boisei, 49 and 34 kg; H. habilis, 52 and 32 kg. The best known African early H. erectus were much larger with weights ranging from 55 kg on up. These estimates imply that (1) in the earliest hominid species and the “robust” australopithecines body sizes remained small relative to modern standards, but between 2.0 and 1.7 m.y.a. there was a rapid increase to essentially modern body size with the appearance of Homo erectus; (2) the earliest species had a degree of body size sexual dimorphism well above that seen in modern humans but below that seen in modern gorillas and orangs which implies (along with other evidence) a social organization characterized by kin-related, multi-male groups with females who were not kin-related; (3) relative brain sizes increased through time; (4) there were two divergent trends in relative cheek-tooth size—a steady increase through time from A. afarensis to A. africanus to the “robust” australopithecines, and a decrease beginning with H. habilis to H. erectus to H. sapiens.     

Disproportionate Cochlear Length in Genus Homo Shows a High Phylogenetic Signal during Apes’ Hearing Evolution

Changes in lifestyles and body weight affected mammal life-history evolution but little is known about how they shaped species’ sensory systems. Since auditory sensitivity impacts communication tasks and environmental acoustic awareness, it may have represented a deciding factor during mammal evolution, including apes. Here, we statistically measure the influence of phylogeny and allometry on the variation of five cochlear morphological features associated with hearing capacities across 22 living and 5 fossil catarrhine species. We find high phylogenetic signals for absolute and relative cochlear length only. Comparisons between fossil cochleae and reconstructed ape ancestral morphotypes show that Australopithecus absolute and relative cochlear lengths are explicable by phylogeny and concordant with the hypothetized ((Pan,Homo),Gorilla) and (Pan,Homo) most recent common ancestors. Conversely, deviations of the Paranthropus oval window area from these most recent common ancestors are not explicable by phylogeny and body weight alone, but suggest instead rapid evolutionary changes (directional selection) of its hearing organ. Premodern (Homo erectus) and modern human cochleae set apart from living non-human catarrhines and australopiths. They show cochlear relative lengths and oval window areas larger than expected for their body mass, two features corresponding to increased low-frequency sensitivity more recent than 2 million years ago. The uniqueness of the “hypertrophied” cochlea in the genus Homo (as opposed to the australopiths) and the significantly high phylogenetic signal of this organ among apes indicate its usefulness to identify homologies and monophyletic groups in the hominid fossil record.     

First Early Hominin from Central Africa (Ishango,Democratic Republic of Congo)

Despite uncontested evidence for fossils belonging to the early hominin genus Australopithecus in East Africa from at least 4.2 million years ago (Ma), and from Chad by 3.5 Ma, thus far there has been no convincing evidence of Australopithecus, Paranthropus or early Homo from the western (Albertine) branch of the Rift Valley. Here we report the discovery of an isolated upper molar (#Ish25) from the Western Rift Valley site of Ishango in Central Africa in a derived context, overlying beds dated to between ca. 2.6 to 2.0 Ma. We used µCT imaging to compare its external and internal macro-morphology to upper molars of australopiths, and fossil and recent Homo. We show that the size and shape of the enamel-dentine junction (EDJ) surface discriminate between Plio-Pleistocene and post-Lower Pleistocene hominins, and that the Ishango molar clusters with australopiths and early Homo from East and southern Africa. A reassessment of the archaeological context of the specimen is consistent with the morphological evidence and suggest that early hominins were occupying this region by at least 2 Ma.     

On the taxonomic affinities of the Dmanisi mandible (Georgia)

The recent discovery of unexpectedly ancient human remains has fuelled interest about the first dispersion of Homo outside Africa. The Dmanisi mandible is perhaps one of the most interesting findings, as it supposedly represents one of the oldest hominids outside of Africa. Recently, different interpretations have been published about this specimen. Our comparison of the Dmanisi mandible with a large sample of mandibles and teeth has led us to a new interpretation. In our view, the Dmanisi mandible exhibits a unique combination of traits. Some of its features, taken in isolation, may be attributed to morphological extremes within the genus Homo. The architecture of the mandible as well as the morphology and dimensions of incisors, canines, and P3s are clearly primitive. However, dental traits such as the reduction of the talonid in the P4s and a distally decreasing molar series seems to be derived. Some combinations of these traits are found in specimens of Homo ergaster and differ from those generally present in later hominids. Thus, we propose that the Dmanisi mandible might be taxonomically classified as Homo sp. indet. (aff. ergaster). Furthermore, some aspects of the dentition in Dmanisi display close similarities to Asian Homo erectus. If the 1.8–1.6 Myr dating for the Dmanisi mandible is correct, the differentiation of the Asian branch of the genus Homo could be regarded as a very ancient event. Am J Phys Anthropol 107:145–162, 1998. © 1998 Wiley-Liss, Inc.     

A solution to the human paradox: fundamental ontogenies and heterochronies

Solving the human paradox means explaining how a genetic difference of a mere 1% can be consistent with 5 million years of anatomical transformation from great apes to present-dayHomo sapiens. The solution proposed here is that of the internal history of ontogenetic change. A concept of “fundamental ontogeny” is developed and deduced from comparison between living and fossil primates. The fossil human lineage can be summarized into five fundamental ontogenies corresponding to successive skull plans (bauplans) resulting from five major phases of craniofacial contraction: prosimians (adapiforms), monkey apes (propliopithecidae), great apes (dryopithecidae), australopithecines andHomo. The morphological areas defined by these skull plans include more-or-less numerous species. This concept leads to renewed debate about (i) the relationship between speciation and bauplans, and (ii) the mechanisms involved in the successive steps of cranio-facial contraction and the correlated morphological changes. It is suggested that, from great apes to modern man, numerous heterochronies (hypermorphosis, hypomorphosis and post-displacements) have occurred during ontogeny, allowing the acquisition of permanent bipedalism inAustralopithecus andHomo, the increased cranial capacity of primitive forms ofHomo, and the disappearance of simian characters associated with renewed increase in cranial capacity inH. sapiens.     

A reappraisal of variation in hominid mandibular corpus dimensions

The relationship between breadth and height of the mandibular corpus has been investigated in a sample of 77 hominid mandibles. An interspecific allometric increase in robusticity with size occurs between four taxonomic subgroups of Australopithecus, but subgroups of Homo vary in robusticity while differing little in size. Within taxonomic subgroups, variation in breadth is not significantly related to variation in height among the “gracile” australapithecines; however, it is isometrically related to height in the “robust” australopithecines and bears an allometric relationship to height in Homo. Thus, robusticity, in conjunction with size, may provide a useful indicator of the taxonomic affinities of hominid mandibles.