Early Homo Specimens Research Articles

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.

Early Pleistocene hominin teeth from Meipu, southern China

The rarity and poor preservation of hominin fossils from the East Asian Early Pleistocene hamper our understanding of their taxonomy and possible phylogenetic relationship with other members of the genus Homo. In the 1970s, four isolated hominin teeth were recovered from the Meipu site, southern China, which biostratigraphic analysis placed in the late Early Pleistocene. Early reports assigned the teeth to late Homo erectus. Since then, the teeth have not been re-evaluated, nor has reliable dating been performed at the Meipu site. Here, biostratigraphic and paleomagnetic dating allow for a more precise chronological constraint of the Meipu hominins in the late Early Pleistocene, between 780 ka and 990 ka, making them one of the few known hominins for this time in mainland Asia. The comparison of the morphology of the Meipu teeth with other members of the genus Homo reveals that the Meipu teeth preserve traits such as moderate shoveling of the I1, the square crown contour of M1, and a buccolingually wider lingual cusp in P4 that make them closer to early Homo specimens from Africa and Homo ergaster from Dmanisi (Georgia). In addition, the Meipu teeth exhibit features that are more typical for late mainland East Asian H. erectus, such as the moderately convex I1 labial surface and a pronouncedly convex I2 labial surface. In these features, the Meipu hominins are morphologically intermediate between African/Dmanisi early Homo and East Asian Middle Pleistocene hominins. This study contributes to a better understanding of the morphologies and the taxonomic status of East Asian Early Pleistocene hominins, a time period for which the hominin evidence with secure stratigraphic context is scarce.

Testing the inhibitory cascade model in the Middle Pleistocene Sima de los Huesos (Sierra de Atapuerca, Spain) hominin sample.

The Middle Pleistocene Sima de los Huesos (SH) site has yielded more than 7.500 human fossil remains belonging to a minimum of 29 individuals. Most of these individuals preserve either the complete mandibular molar series or at least the first (M1 ) and second (M2 ) molars. The inhibitory cascade mathematical model was proposed by Kavanagh et al. (Nature, 449, 427-433 [2007]) after their experimental studies on the dental development of murine rodent species. The activator-inhibitor mechanism of this model has shown its ability for predicting evolutionary size patterns of mammalian teeth, including hominins. The main aim of this study is to test whether the size molar patterns observed in the SH hominins fit the inhibitory cascade model. With this purpose, we have measured the crown area of all SH molars in photographs, using a planimeter and following techniques used and well contrasted in previous works. Following one of the premises of the inhibitory cascade model, we expect that the central tooth (M2 in our case) of a triplet would have the average size of the two outer teeth. The absolute difference between the observed and the expected values for the M2 s ranges from 0.23 to 8.46mm2 in the SH sample. In terms of percentage, the difference ranges between 0.25% and 10.34%, although in most cases, it is below 5%. The plot of the estimated M3 /M1 and M2 /M1 size ratios obtained in the SH hominins occupies a small area of the theoretical developmental morphospace obtained for rodent species. In addition, the majority of the values are placed near the theoretical line which defines the relationship predicted by the inhibitory cascade model in these mammals. The values of the slope and intercept of the reduced major regression obtained for the SH individuals do not differ significantly from those obtained for rodent species, thus confirming that the size of the molars of the SH hominins fits the inhibitory cascade model. We discuss these results in terms of dental development. Despite the promising results in the SH sample, we draw the attention to the fact that most Early Pleistocene Homo specimens exhibit a pattern (M1 <M2 >M3 ), which is outside the expected theoretical morphospace predicted by the inhibitory cascade model. The shift from the M1<M2<M3 size relationship observed in early hominins (including H.habilis) to the M1>M2>M3 size relationship, which is predominant in modern humans, includes sequences that depart from predictions of the inhibitory cascade model. Additional studies are required to understand these deviations.

Contemporaneity of Australopithecus, Paranthropus, and early Homo erectus in South Africa.

Understanding the extinction of Australopithecus and origins of Paranthropus and Homo in South Africa has been hampered by the perceived complex geological context of hominin fossils, poor chronological resolution, and a lack of well-preserved early Homo specimens. We describe, date, and contextualize the discovery of two hominin crania from Drimolen Main Quarry in South Africa. At ~2.04 million to 1.95 million years old, DNH 152 represents the earliest definitive occurrence of Paranthropus robustus, and DNH 134 represents the earliest occurrence of a cranium with clear affinities to Homo erectus These crania also show that Homo, Paranthropus, and Australopithecus were contemporaneous at ~2 million years ago. This high taxonomic diversity is also reflected in non-hominin species and provides evidence of endemic evolution and dispersal during a period of climatic variability.

Maxillary molar enamel thickness of Plio-Pleistocene hominins

Enamel thickness remains an important morphological character in hominin systematics and is regularly incorporated into dietary reconstructions in hominin species. We expand upon a previous study of enamel thickness in mandibular molars by examining a large maxillary molar sample of Plio-Pleistocene hominins (n = 62) and a comparative sample of extant nonhuman apes (n = 48) and modern humans (n = 29). 2D mesial planes of section were generated through microtomography, and standard dental tissue variables were measured to calculate average enamel thickness (AET) and relative enamel thickness (RET). AET was also examined across the lingual, occlusal, and buccal regions of the crown. This study confirms previous findings of increasing enamel thickness throughout the Plio-Pleistocene, being thinnest in Australopithecus anamensis and peaking in Australopithecus boisei, with early Homo specimens, exhibiting intermediate enamel thickness. Agreeing with previous findings, 2D plane of section enamel thickness is found to be a poor taxonomic discriminator, with no statistically significant differences observed between fossil hominins. For fossil hominins, modern humans, and Pongo, the occlusal region of enamel was the thickest, and the lingual enamel thickness was greater than buccal. Pan and Gorilla present the opposite pattern with enamel being thinnest occlusally. Comparison at each molar position between the maxilla and mandible revealed very few significant differences in fossil hominins but some evidence of significantly thicker maxillary enamel (AET) in modern humans and thinner maxillary enamel in Pan (RET).

Endostructural morphology in hominoid mandibular third premolars: Geometric morphometric analysis of dentine crown shape

In apes, the mandibular third premolar (P3) is adapted for a role in honing the large upper canine. The role of honing was lost early in hominin evolution, releasing the tooth from this functional constraint and allowing it to respond to subsequent changes in masticatory demands. This led to substantial morphological changes, and as such the P3 has featured prominently in systematic analyses of the hominin clade. The application of microtomography has also demonstrated that examination of the enamel-dentine junction (EDJ) increases the taxonomic value of variations in crown morphology. Here we use geometric morphometric techniques to analyze the shape of the P3 EDJ in a broad sample of fossil hominins, modern humans, and extant apes (n = 111). We test the utility of P3 EDJ shape for distinguishing among hominoids, address the affinities of a number of hominin specimens of uncertain taxonomic attribution, and characterize the changes in P3 EDJ morphology across our sample, with particular reference to features relating to canine honing and premolar ‘molarization’. We find that the morphology of the P3 EDJ is useful in taxonomic identification of individual specimens, with a classification accuracy of up to 88%. The P3 EDJ of canine-honing apes displays a tall protoconid, little metaconid development, and an asymmetrical crown shape. Plio-Pleistocene hominin taxa display derived masticatory adaptations at the EDJ, such as the molarized premolars of Australopithecus africanus and Paranthropus, which have well-developed marginal ridges, an enlarged talonid, and a large metaconid. Modern humans and Neanderthals display a tall dentine body and reduced metaconid development, a morphology shared with premolars from Mauer and the Cave of Hearths. Homo naledi displays a P3 EDJ morphology that is unique among our sample; it is quite unlike Middle Pleistocene and recent Homo samples and most closely resembles Australopithecus, Paranthropus and early Homo specimens.

Intra-individual metameric variation expressed at the enamel-dentine junction of lower post-canine dentition of South African fossil hominins and modern humans.

The aim of this study is to compare the degree and patterning of inter- and intra-individual metameric variation in South African australopiths, early Homo and modern humans. Metameric variation likely reflects developmental and taxonomical issues, and could also be used to infer ecological and functional adaptations. However, its patterning along the early hominin postcanine dentition, particularly among South African fossil hominins, remains unexplored. Using microfocus X-ray computed tomography (µXCT) and geometric morphometric tools, we studied the enamel-dentine junction (EDJ) morphology and we investigated the intra- and inter-individual EDJ metameric variation among eight australopiths and two early Homo specimens from South Africa, as well as 32 modern humans. Along post-canine dentition, shape changes between metameres represented by relative positions and height of dentine horns, outlines of the EDJ occlusal table are reported in modern and fossil taxa. Comparisons of EDJ mean shapes and multivariate analyses reveal substantial variation in the direction and magnitude of metameric shape changes among taxa, but some common trends can be found. In modern humans, both the direction and magnitude of metameric shape change show increased variability in M2 -M3 compared to M1 -M2 . Fossil specimens are clustered together showing similar magnitudes of shape change. Along M2 -M3 , the lengths of their metameric vectors are not as variable as those of modern humans, but they display considerable variability in the direction of shape change. The distalward increase of metameric variation along the modern human molar row is consistent with the odontogenetic models of molar row structure (inhibitory cascade model). Though much remains to be tested, the variable trends and magnitudes in metamerism in fossil hominins reported here, together with differences in the scale of shape change between modern humans and fossil hominins may provide valuable information regarding functional morphology and developmental processes in fossil species.

Structural organization and tooth development in a Homo aff. erectus juvenile mandible from the Early Pleistocene site of Garba IV at Melka Kunture, Ethiopian highlands.

The immature partial mandible GAR IVE from the c. 1.7 Ma old Garba IV site at Melka Kunture (Upper Awash Basin, Ethiopia), the earliest human representative from a mountain-like environment, represents one of the oldest early Homo specimens bearing a mixed dentition. Following its first description (Condemi, ), we extended the analytical and comparative record of this specimen by providing unreported details about its inner morphology, tooth maturational pattern and age at death, crown size, and tooth tissue proportions. The new body of quantitative structural information and virtual imaging derives from a medical CT record performed in 2013. Compared to the extant human condition and to some fossil representatives of comparable individual age, the GAR IVE mandible reveals absolutely and relatively thick cortical bone. Crown size of the permanent lateral incisor and the canine fit the estimates of H. erectus s.l., while the dm2 and the M1 more closely approach those of H. habilis-rudolfensis. Molar crown pulp volumes are lower than reported in other fossil specimens and in extant humans. The mineralization sequence of the permanent tooth elements is represented four times in our reference sample of extant immature individuals (N = 795). The tooth developmental pattern displayed by the immature individual from Garba IV falls within the range of variation of extant human populations and is also comparable with that of other very young early fossil hominins. Taken together, the evidence presented here for mandibular morphology and dental development suggest GAR IVE is a robust 2.5- to 3.5-year old early Homo specimen.

Morphometric analysis of Atapuerca-Sima de los Huesos lower first molars

The study of dental remains is a wealthy and stable source of information about taxonomical relations between extinct hominin species. Both morphological and metric dental features can be a reliable indicator of the underlying genotype of the individual/species under study. First lower molars (M1s) are the most stable teeth within the lower molar series. Since the last systematic metric analyses of the Atapuerca-Sima de los Huesos (SH) lower molars (Bermudez de Castro and Nicolas, 1995) the dental sample has substantially increased from 133 to over 533 specimens, so an updated assessment was necessary. Here, we analyze the absolute and relative proportions of the crown and the cusps of the SH M1s and we compare it against a large fossil simple including Australopithecus, Paranthropus, early Homo sapiens, Homo neanderthalensis, Homo heidelbergensis, Homo antecessor and Asian and African early Homo specimens, as well as a wide sample of contemporary Homo sapiens. Our study reveals that concerning M1s absolute size SH does not differ significantly from contemporary Homo sapiens. Homo antecessor M1s fall near to Asian Early and Middle Pleistocene Homo and differ from SH and H. sapiens. Regarding the relative cusp proportion we find three patterns: 1) a pattern that characterizes non-Homo specimens and some African Early Pleistocene Homo specimens (metaconid > protoconid > hypoconid > entoconid > hypoconulid), 2) a pattern that characterizes Early and Middle Pleistocene Homo from Asia and Europe including early Homo sapiens (protoconid > metaconid > hypoconid > entoconid > hypoconulid), 3) a pattern represented by contemporary Homo sapiens and Neandertals (protoconid > metaconid > entoconid > hypoconid > hypoconulid) and 4) an exclusive pattern for SH (pro > met = ent = hyp > hld). The SH protoconid is the largest from all the studied groups. Interestingly, this result matches previous observations by Martinon-Torres et al. (2013) regarding the protocone in SH upper first molars. We suggest that this may be an exclusive feature for this population. Molar size reduction in SH and contemporary Homo sapiens suggests a likely case of evolutionary parallelism. However, the cusp pattern variation differ between SH and contemporary Homo sapiens, suggesting that the evolutionary mechanisms of dental reduction in each group are different. Analysis of other dental classes may be necessary.

Femoral neck structure and function in early hominins

All early (Pliocene-Early Pleistocene) hominins exhibit some differences in proximal femoral morphology from modern humans, including a long femoral neck and a low neck-shaft angle. In addition, australopiths (Au. afarensis, Au. africanus, Au. boisei, Paranthropus boisei), but not early Homo, have an "anteroposteriorly compressed" femoral neck and a small femoral head relative to femoral shaft breadth. Superoinferior asymmetry of cortical bone in the femoral neck has been claimed to be human-like in australopiths. In this study, we measured superior and inferior cortical thicknesses at the middle and base of the femoral neck using computed tomography in six Au. africanus and two P. robustus specimens. Cortical asymmetry in the fossils is closer overall to that of modern humans than to apes, although many values are intermediate between humans and apes, or even more ape-like in the midneck. Comparisons of external femoral neck and head dimensions were carried out for a more comprehensive sample of South and East African australopiths (n = 17) and two early Homo specimens. These show that compared with modern humans, femoral neck superoinferior, but not anteroposterior breadth, is larger relative to femoral head breadth in australopiths, but not in early Homo. Both internal and external characteristics of the australopith femoral neck indicate adaptation to relatively increased superoinferior bending loads, compared with both modern humans and early Homo. These observations, and a relatively small femoral head, are consistent with a slightly altered gait pattern in australopiths, involving more lateral deviation of the body center of mass over the stance limb.

Imitation and Shared Intentionality in the Acheulean

Imitation and shared intentionality are traits essential to the socio-cultural adaptation of Homo sapiens. Non-human apes display some capacity for imitation and shared intentionality, but are deficient in comparison to Homo sapiens. The Acheulean archaeological record provides evidence that imitation and shared intentionality were part of the behavioural repertoire of the hominins manufacturing that industry. The palaeoneurological record suggests some early Homo specimens possessed the neural architecture associated with modern human imitation. From this evidence it is suggested that a propensity for imitation and shared intentionality evolved soon after 2 million years ago.

Body size and body shape in early hominins – implications of the Gona Pelvis

Discovery of the first complete Early Pleistocene hominin pelvis, Gona BSN49/P27, attributed to Homo erectus, raises a number of issues regarding early hominin body size and shape variation. Here, acetabular breadth, femoral head breadth, and body mass calculated from femoral head breadth are compared in 37 early hominin (6.0–0.26 Ma) specimens, including BSN49/P27. Acetabular and estimated femoral head sizes in the Gona specimen fall close to the means for non- Homo specimens ( Orrorin tugenesis, Australopithecus africanus, Paranthropus robustus), and well below the ranges of all previously described Early and Middle Pleistocene Homo specimens. The Gona specimen has an estimated body mass of 33.2 kg, close to the mean for the non- Homo sample (34.1 kg, range 24–51.5 kg, n = 19) and far outside the range for any previously known Homo specimen (mean = 70.5 kg; range 52–82 kg, n = 17). Inclusion of the Gona specimen within H. erectus increases inferred sexual dimorphism in body mass in this taxon to a level greater than that observed here for any other hominin taxon, and increases variation in body mass within H. erectus females to a level much greater than that observed for any living primate species. This raises questions regarding the taxonomic attribution of the Gona specimen. When considered within the context of overall variation in body breadth among early hominins, the mediolaterally very wide Gona pelvis fits within the distribution of other lower latitude Early and Middle Pleistocene specimens, and below that of higher latitude specimens. Thus, ecogeographic variation in body breadth was present among earlier hominins as it is in living humans. The increased M-L pelvic breadth in all earlier hominins relative to modern humans is related to an increase in ellipticity of the birth canal, possibly as a result of a non-rotational birth mechanism that was common to both australopithecines and archaic Homo.

A new early Pleistocene hominin mandible from Atapuerca-TD6, Spain

We present the description of a new mandibular specimen, ATD6-113, recovered in 2006 from the TD6 level of the Gran Dolina cave site in Sierra de Atapuerca, northern Spain. A detailed study of the lithostratigraphy of the top sequence of this level, the section from where all human remains have been recovered so far, is also presented. We have observed that the hominin stratum, previously defined as Aurora Stratum, represents a condensed deposit of at least six layers, which could not be distinguished in the test pit made in 1994–95. Therefore, the human fossil remains were probably deposited during a discrete and undetermined time period. The new mandibular fragment exhibits a very similar morphology to that of the most complete specimen, ATD6-96, which was recovered in 2003 from a different layer. This suggests that both specimens represent the same biological population. The two mandibles, as well as the small mandibular fragment ATD6-5 (which constitutes part of the holotype of Homo antecessor), present a morphological pattern clearly derived with regard to that of the African early Homo specimens usually included in H. habilis and H. rudolfensis, the mandibles D211 and D2735 from Dmanisi, and most of the early Pleistocene mandibles from Sangiran. The TD6 mandibles also exhibit some derived features with regard to the African early Pleistocene specimens included in H. ergaster (or African H. erectus). Thus, the TD6 hominins seem to represent a lineage different from other African and Asian lineages, although some (metric in particular) similarities with Chinese middle Pleistocene mandibles are noted. Interestingly, none of the apomorphic mandibular features of the European middle and early late Pleistocene hominins are present in the TD6 mandibles.

A geometric morphometric analysis of hominin upper first molar shape

Recent studies have revealed interesting differences in upper first molar morphology across the hominin fossil record, particularly significant between H. sapiens and H. neanderthalensis. Usually these analyses have been performed by means of classic morphometric methods, including the measurement of relative cusp areas or the angles defined between cusps. Although these studies have provided valuable information for the morphological characterization of some hominin species, we believe that the analysis of this particular tooth could be more conclusive for taxonomic assignment. In this study, we have applied geometric morphometric methods to explore the morphological variability of the upper first molar (M 1) across the human fossil record. Our emphasis focuses on the study of the phenetic relationships among the European middle Pleistocene populations (designated as H. heidelbergensis) with H. neanderthalensis and H. sapiens, but the inclusion of Australopithecus and early Homo specimens has helped us to assess the polarity of the observed traits. H. neanderthalensis presents a unique morphology characterized by a relatively distal displacement of the lingual cusps and protrusion in the external outline of a large and bulging hypocone. This morphology can be found in a less pronounced degree in the European early and middle Pleistocene populations, and reaches its maximum expression with the H. neanderthalensis lineage. In contrast, modern humans retain the primitive morphology with a square occlusal polygon associated with a round external outline.

Les limites du Quaternaire : 1,8 ou 2,6 millions d’années? Apports de l’étude concernant les premiers représentants du genre homo

The Quaternary, which is sometimes also termed the Anthropogene Period, often is linked to the beginning of humankind. But what exactly is the basis in the fossil hominin record for this link, in particular what is the early Homo evidence? Since 1999, debate concerning the definition of the genus Homo and the identification of its first members has intensified. At the centre of recent debates, there are four hypotheses: (1) all early Homo specimens belong to the same species: Homo habilis sensu lato; (2) the early Homo hypodigm is heterogeneous, two species could be defined: Homo habilis sensu stricto and Homo rudolfensis; (3) these species do not belong to the genus Homo but to the genus Australopithecus; or (4) it would be more appropriate to put the specimens of Homo rudolfensis into the genus Kenyanthropus. The earliest appearance of the genus Homo will differ depending upon the favoured hypothesis: 2.45 Myr (hypotheses 1 and 2); 1.9 Myr (hypothesis 3); 1.9 or 2.4 Myr (hypothesis 4, depending on the genus attribution for habilis (2.4 Myr if Homo; 1.9 Myr if Australopithecus)). Our study, based on palaeoanthropological data, is focused on the 2.6-1.6 Myr period. In this paper, we propose to review and assess knowledge of early Homo taxonomy in the context of the definition of the Quaternary boundary.

Dental microwear and diets of African early Homo

Conventional wisdom ties the origin and early evolution of the genus Homo to environmental changes that occurred near the end of the Pliocene. The basic idea is that changing habitats led to new diets emphasizing savanna resources, such as herd mammals or underground storage organs. Fossil teeth provide the most direct evidence available for evaluating this theory. In this paper, we present a comprehensive study of dental microwear in Plio-Pleistocene Homo from Africa. We examined all available cheek teeth from Ethiopia, Kenya, Tanzania, Malawi, and South Africa and found 18 that preserved antemortem microwear. Microwear features were measured and compared for these specimens and a baseline series of five extant primate species ( Cebus apella, Gorilla gorilla, Lophocebus albigena, Pan troglodytes, and Papio ursinus) and two protohistoric human foraging groups (Aleut and Arikara) with documented differences in diet and subsistence strategies. Results confirmed that dental microwear reflects diet, such that hard-object specialists tend to have more large microwear pits, whereas tough food eaters usually have more striations and smaller microwear features. Early Homo specimens clustered with baseline groups that do not prefer fracture resistant foods. Still, Homo erectus and individuals from Swartkrans Member 1 had more small pits than Homo habilis and specimens from Sterkfontein Member 5C. These results suggest that none of the early Homo groups specialized on very hard or tough foods, but that H. erectus and Swartkrans Member 1 individuals ate, at least occasionally, more brittle or tough items than other fossil hominins studied.

Palaeopathological and variant conditions of the Homo heidelbergensis type specimen (Mauer, Germany).

Although early Homo specimens are now known from a number of African, Asian and European Middle Pleistocene sites, the taxon Homo heidelbergensis was initially introduced for the Mauer jaw recovered in 1907. Fossil hominids from the earlier Middle Pleistocene of Europe are very rare and the Mauer mandible is generally accepted as one of the most ancient, with an age of approximately 700 kyr. A new preparation of the mandible was conducted in 1996 and gave rise to the detailed palaeopathological examination which is presented here. Based on comparative analyses, the extreme breadth of the mandibular ramus and its flat intercondylar incision, in conjunction with the flattening and broadening of the coronoid process tip, results either from an idiosyncratic pattern of the course and insertion of the temporalis muscle on the coronoid process or from the temporalis possessing an accessory head. The incidence of periodontal pocketing, together with a vertical reduction of the alveolar margin to approximately 3.00 mm, and a slight protuberance formed in vicinity of the right M(2)can safely be interpreted as pathognomonic indications of periodontal disease. The short distance between the enamel-dentine junction of the teeth and the horizontal alveolar margins could either be an inherited variant or may result from incipient osteoporosis. In addition, an arthrotic condition with slight osteophytic peripheral exostoses and an arthrolit (i.e. an articular calculus or "joint mouse") on the left condylus articularisand a depression in the medial part of the left mandibular condyle extending into the inferior part of the ramus are present. These features are indicative of a trauma-induced osteochondrosis dissecans. The diagnosis therefore suggests that the observed depression results from a well-healed fracture. This traumatic event illustrates the demanding living conditions endured by humans during the European Middle Pleistocene. The variations and pathological conditions observed in Mauer do not question the mandible's role as type specimen for the taxon Homo heidelbergensis.

Implications of early hominid labyrinthine morphology for evolution of human bipedal locomotion.

The upright posture and obligatory bipedalism of modern humans are unique among living primates. The evolutionary history of this behaviour has traditionally been pursued by functional analysis of the postcranial skeleton and the preserved footprint trails of fossil hominids. Here we report a systematic attempt to reconstruct the locomotor behaviour of early hominids by looking at a major component of the mechanism for the unconscious perception of movement, namely by examining the vestibular system of living primates and early hominids. High-resolution computed tomography was used to generate cross-sectional images of the bony labyrinth. Among the fossil hominids the earliest species to demonstrate the modern human morphology is Homo erectus. In contrast, the semicircular canal dimensions in crania from southern Africa attributed to Australopithecus and Paranthropus resemble those of the extant great apes. Among early Homo specimens, the canal dimensions of Stw 53 are unlike those seen in any of the hominids or great apes, whereas those of SK 847 are modern-human-like.

Postcranial robusticity in Homo. III: Ontogeny.

The influence of developmental factors on long-bone cross-sectional geometry and articular size in modern humans is investigated using two approaches: (1) an analysis of the effects of increased mechanical loading on long-bone structure when applied during different developmental periods, using data collected for a study of upper limb bone bilateral asymmetry in professional tennis players; and (2) an analysis of the relative timing of age changes in femoral dimensions among juveniles from the Pecos Pueblo Amerindian archaeological sample. Results of these analyses are used to interpret the femoral morphology of three pre-Recent Homo juveniles--the H. erectus KNM-WT 15000 and the archaic H. sapiens La Ferrassie 6 and Teshik-Tash 1--as well as observed differences in postcranial morphology between adult Recent and earlier Homo (Ruff et al., 1993). Our findings indicate the following: (1) There are age-related changes in long-bone diaphyseal envelope sensitivity to increased mechanical loading, with the periosteal envelope more responsive prior to mid-adolescence, and the endosteal envelope more responsive thereafter. The periosteal expansion and endosteal contraction of the diaphysis documented earlier for adult pre-Recent Homo relative to Recent humans (Ruff et al., 1993) is thus consistent with a developmental response to increased mechanical loading applied throughout life. The relatively large medullary cavity in the 11-12-year-old KNM-WT 15000 femur is also consistent with this model. However, the two archaic H. sapiens juveniles show relatively small medullary cavities, possibly indicating a modified developmental pattern in this group. (2) Articulations follow a growth pattern similar to that of long-bone length (and stature), while cross-sectional diaphyseal dimensions (cortical area, second moments of area) show a contrasting growth pattern, with slower initial growth from childhood through mid-adolescence, followed by a "catch-up" period that continues through early adulthood. This latter pattern is more similar to the growth curve for body weight, and may in fact partially reflect adaptation of the diaphysis to increased weight bearing. Because of these different growth patterns, articulations appear relatively large, and diaphyseal breadths relatively small during late childhood to mid-adolescence (i.e., about 9-13 years), when compared to adults from the same population. KNM-WT 15000 shows this same proportional difference from adult early Homo specimens, which is therefore interpreted as simply a developmental consequence of his age at death.(ABSTRACT TRUNCATED AT 400 WORDS)

The ontogeny of Pan troglodytes craniofacial architectural relationships and implications for early hominids

Developmental constraints characterize functional growth boundaries of the mammalian face. Enlow and colleagues identified boundary conditions and planes of the midface in humans and other mammal species that are defined on the basis of important growth sites and the developmental disposition of neural and pharyngeal matrices. With few exceptions a conservative mammalian architecture is said to be achieved by the adult stage. Three aspects of this architecture are investigated here for a cross-sectional ontogenetic series of Pan troglodytes crania: (1) a line passing from the maxillary tuberosity through the junction of middle and anterior cranial fossae is perpendicular to the neutral horizontal axis of the orbit (PM-NHA angle), (2) an average 45° angle, whose origin is the external auditory meatus, separates the maxillary tuberosity from the midpoint of the orbital opening (meatus angle) and (3) the base of the brain, maxillary tuberosity and prosthion are on or close to the same plane (anterior maxillary hypoplasia). This investigation considers to what extent samples of chimpanzees and early hominids reflect Enlow's characterization of mammalian craniofacial architecture. Evidence derived from the chimpanzee sample bears out the perpendicularity of the PM-HNA angle and a near 45° meatus angle (though slightly higher overall), but does not conform to the absence of marked anterior maxillary hypoplasia seen in other mammals. Pan is instead characterized by considerable ontogenetic variation (noted by Enlow amongst anthropoids) influenced to some extent by sex. Preliminary early hominid data on the meatus angle indicates that Australopithecus specimens are similar to the widespread and probably primitive condition, while Paranthropus and early Homo specimens portray an uncharacteristic mammalian architectural relationship.