This study of the developing pongid dentition is based on cross-sectional radiographic data of juvenile Pan troglodytes, Gorilla gorilla, and Pongo pygmaeus skulls. Comparisons with developmental features of the human dentition are made, and possible explanations for the formation of larger teeth within the reduced pongid growth period are discussed. The data presented in this study provide an alternative method for ageing individual pongid crania in comparative cross-sectional growth studies. The advantages of this method are demonstrated by ageing individual Gorilla crania from radiographs and plotting relative dental age against length of the jaw.

Tooth fragments are an appreciable but neglected proportion of fossil hominid specimens. The present study on 47 naturally fractured enamel surfaces of premolar and molar teeth of Plio-Pleistocene East African hominids measured enamel thickness, slope of incremental lines (striae of Retzius), and the morphology of Hunter Schreger bands (HSBs). Specimens allocated to three categories--"robust" australopithecines (EAFROB), "early Homo" (EAFHOM), and "unknown"--were photographed in ethanol with polarised light. Enamel thickness was measured on the occlusal (OT), cuspal (CT), and lateral (LT) aspects. The angle of intersection of striae of Retzius (D) with the enamel-dentine junction (EDJ) was recorded, together with the degree of curvature and width of Hunter-Schreger bands (HSB). Absolute measurements of enamel thickness were scaled by using two allometry correction factors. Absolute thicknesses of all enamel measurements were significantly greater in the EAFROB (OT 3.1 mm; CT 3.3 mm; LT 2.4 mm) compared with EAFHOM (OT 1.4 mm; CT 1.6 mm; LT 1.6 mm) categories. Correction for size reduces the mean difference between the two taxa, but CT and OT thickness remain significantly different (P less than 0.05). HSBs in EAFROB were relatively straight and narrower (means = 52.8 micron) than in EAFHOM, which are more curved and wider (means = 62.0 micron), suggesting greater enamel prism decussation in early Homo. The slope of striae was less in EAFROB permanent molars (means = 23 degrees) compared with EAFHOM (means = 31 degrees), indicating faster rates of coverage during crown formation in "robust" australopithecines. We conclude that the study of fractured enamel surfaces can contribute to our understanding of the systematic relationships and patterns of enamel growth of early hominids.

A recent study of the surface manifestation of incremental lines associated with enamel formation suggested that the crowns of early hominid incisor teeth were formed more rapidly than those of modern humans. In the absence of comparative data, the authors were forced to assume that enamel increments in fossil teeth were similar to those in modern humans. We have used evidence from the fractured surfaces of molar teeth to deduce estimates for both long- and short-period incremental growth markers within enamel in east African 'robust' australopithecine and early Homo teeth. We conclude that in these early hominids, crown formation times in posterior teeth, particularly in the large thick enamelled molar teeth of the east African 'robust' australopithecines, were shorter than those of modern humans. This evidence, considered together with data on crown and root formation times in modern apes, suggests that the posterior teeth in these hominids both formed and erupted more rapidly than those of modern man. These results have implications for attempts to assess dental and skeletal maturity in hominids.

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We report here on early hominid facial diversity, as part of a more extensive morphometric survey of cranial variability in Pliocene and early Pleistocene Hominidae. Univariate and multivariate techniques are used to summarise variation in facial proportions in South and East African hominids, and later Quaternary groups are included as comparators in order to scale the variation displayed. The results indicate that "robust" australopithecines have longer, broader faces than the "gracile" form, but that all australopithecine species show comparable degrees of facial projection. "Robust" crania are characterised by anteriorly situated, deep malar processes that slope forwards and downwards. Smaller hominid specimens, formally or informally assigned to Homo (H. habilis, KNM-ER 1813, etc.), have individual facial dimensions that usually fall within the range of Australopithecus africanus, but which in combination reveal a significantly different morphological pattern; SK 847 shows similarly hominine facial proportions, which differ significantly from those of A. robustus specimens from Swartkrans. KNM-ER 1470 possesses a facial pattern that is basically hominine, but which in some respects mimics that of "robust" australopithecines. Early specimens referred to H. erectus possess facial proportions that contrast markedly with those of other Villafranchian hominids and which suggest differing masticatory forces, possibly reflecting a shift in dietary niche. Overall the results indicate two broad patterns of facial proportions in Hominidae: one is characteristic of Pliocene/basal Pleistocene forms with opposite polarities represented by A. boisei and H. habilis; the other pattern, which typifies hominids from the later Lower Pleistocene onwards, is first found in specimens widely regarded as early representatives of H. erectus, but which differ in which certain respects from the face of later members of that species.

We have examined the crowns of chimpanzee, australopith, and Paranthropus species and early Homo in order to investigate two different, widely recognized, dental trends in Plio-Pleistocene hominin evolution. They are a reduction in crown size and morphological complexity in Homo, and an increase in crown size and morphological complexity in Paranthropus. A phenetic assessment of maxillary and mandibular molar crown non-metrical traits revealed that two australopith species (Au. africanus and Au. afarensis) are much more similar to each other than either is to Paranthropus, and together all hominins are distinctively different from chimpanzees (P. troglodytes and P. paniscus). The difference between Paranthropus and australopith postcanine teeth was 20-30 times greater than that between the australopith species and the difference between the two australopith species was about half the difference between the two extant chimpanzee species. The characters that contribute to the increase in crown complexity seen in Paranthropus do not appear to be primitive retentions from a great ape ancestor, and there is 34Hominin Postcanine Occlusal Morphology some evidence that the same, or a very similar, trend towards trait intensification is already present in australopiths. These traits include additional cusps on the maxillary and mandibular molars, and the expanded P 4 talonid. Early Homo exhibits the primitive condition for many of the molar traits, but it has also lost many other primitive traits (upper molar anterior and posterior foveae, for example) that are present in the australopiths. Relative to Pan, and similar to the australopiths, early Homo possesses a larger P 4 with a somewhat expanded talonid, but this trend is subsequently reversed in later Homo. Our study reveals that some of the dental trends said to be characteristic of Homo actually appear relatively late in human evolution.

The venous valves are believed to play an important role in venous function, but their number, position, and spacing in limb veins are reported to be irregular. In this study, the relationship between the number of valves and the length of veins in which they occur was investigated for humans and nonhuman primates. In addition, valve distributions within the superficial veins of the human upper limb were compared with those of the lower limb. Upper and lower limb veins were dissected from adult humans, and forelimb veins were obtained from seven genera of primates. The mean valve index (number of valves per unit length of vein) of the forelimb lateral superficial vein exhibited relatively little variation between humans and primates of a wide range of body sizes and locomotor repertoires. The mean valve index for the lateral superficial vein of the human upper limb is similar to that of the short saphenous vein of the lower limb, and in both veins, the maximum valve index occurs in the most proximal portion of the vein. The relative constancy of the valve index across a range of primate species, and between the upper and lower limbs of humans, may reflect the known constancy of circulatory pressures in mammals. The minimum numbers of valves in superficial veins are only slightly greater than required to limit capillary pressure to below 10 mm Hg, above which pressure the tissue fluid flow would be compromised.

The ontogeny and comparative anatomy of the forelimb superficial veins were investigated in humans, non-human primates and other mammals. Adult humans and the orangutan (Pongo) possess two autonomous forelimb veins, one on the lateral (preaxial) margin of the limb, the other on the medial (postaxial) margin. All other adult primates and mammals examined possess a lateral vein alone. In African apes (Pan and Gorilla) and in 24% of human forelimbs the lateral vein is short, being essentially confined to the antebrachial region, whereas in other mammals and in 76 % of human limbs the lateral vein runs from the carpus to the clavicular region. In humans the medial vein develops before the lateral vein, whereas in the rabbit and the pig the medial vein is present in early embryos but is subsequently lost. We propose that in humans, and probably also in the orangutan, the possession of a medial vein is a neotenic retention of a primitive tetrapod condition. These animals, which retain their medial vein, are united by losing a late stage in their ontogeny. Other animals subsequently pass through a stage in which the medial vein is lost, but Pongo and Homo retain this vein to adulthood. The loss of an ontogenetic stage can arise independently, and the presence of a medial vein therefore affords only weak evidence for a close phylogenetic relationship between humans and the orangutan. The polymorphic lateral vein of humans may be a character state that is intermediate between the derived (short) lateral vein of the African apes and the primitive long lateral vein of other non-human primates and mammals.