Worldwide variation in hip fracture incidence weakly aligns with genetic divergence between populations.

UNLABELLED: This study investigates the influence of genetic differentiation in determining worldwide heterogeneity in osteoporosis-related hip fracture rates. The results indicate that global variation in fracture incidence exceeds that expected on the basis of random genetic variance. INTRODUCTION: Worldwide, the incidence of osteoporotic hip fractures varies considerably. This variability is believed to relate mainly to non-genetic factors. It is conceivable, however, that genetic susceptibility indeed differs across populations. Here, we present the first quantitative assessment of the effects of genetic differentiation on global variability in hip fracture rates. METHODS: We investigate the observed variance in publically reported age-standardized rates of hip fracture among 28 populations from around the world relative to the expected variance given the phylogenetic relatedness of these populations. The extent to which these variances are similar constitutes a "phylogenetic signal," which was measured using the K statistic. Population genetic divergence was calculated using a robust array of genome-wide single nucleotide polymorphisms. RESULTS: While phylogenetic signal is maximized when K > 1, a K value of only 0.103 was detected in the combined-sex fracture rate pattern across the 28 populations, indicating that fracture rates vary more than expected based on phylogenetic relationships. When fracture rates for the sexes were analyzed separately, the degree of phylogenetic signal was also found to be small (females: K = 0.102; males: K = 0.081). CONCLUSIONS: The lack of a strong phylogenetic signal underscores the importance of factors other than stochastic genetic diversity in shaping worldwide heterogeneity in hip fracture incidence.

Cochlear shape distinguishes southern African early hominin taxa with unique auditory ecologies.

Insights into potential differences among the bony labyrinths of Plio-Pleistocene hominins may inform their evolutionary histories and sensory ecologies. We use four recently-discovered bony labyrinths from the site of Kromdraai to significantly expand the sample for Paranthropus robustus. Diffeomorphometry, which provides detailed information about cochlear shape, reveals size-independent differences in cochlear shape between P. robustus and Australopithecus africanus that exceed those among modern humans and the African apes. The cochlea of P. robustus is distinctive and relatively invariant, whereas cochlear shape in A. africanus is more variable, resembles that of early Homo, and shows a degree of morphological polymorphism comparable to that evinced by modern species. The curvature of the P. robustus cochlea is uniquely derived and is consistent with enhanced sensitivity to low-frequency sounds. Combined with evidence for selection, our findings suggest that sound perception shaped distinct ecological adaptations among southern African early hominins.

Australopithecine enamel prism patterns.

Following a recent suggestion that tooth enamel prism shape differs within Hominoidea, the teeth of a number of extinct and extant hominoid species were analyzed by scanning electron microscopy. The enamel prism patterns of some gracile and robust australopithecine specimens from Sterkfontein, Swartkrans, and Kromdraai are recorded. The characteristic arrangements of enamel prisms in all modern and extinct hominoid species were found to be essentially similar. The implications of enamel prisms for phylogenetic deduction in Hominoidea are discussed.

A review of taurodontism with new data on indigenous southern African populations.

The prevalence and degree of taurodontism (enlargement of the pulp cavity) in the mandibular permanent molars of two recent population samples from southern Africa were investigated quantitatively from lateral radiographs. The degree of occlusal wear was scored and two measures of relative pulp cavity size were recorded for each tooth. There was a significant association between increased attrition and a reduction in the size of the pulp cavity when all of the teeth were considered together, but no correlation among individual molar types within each sample. In both the Zulu (n=68 individuals) and Khoisan (n=28 individuals) samples, third molars had the highest prevalence of taurodontism and first molars the lowest. The data for Zulus are similar to those recorded for other modern populations (e.g. white and black Americans, and Israelis), whereas the Khoisan data exhibit significantly higher frequencies. An increased appreciation of the distribution of this variant in modern human populations would contribute to an understanding of its possible evolutionary significance in the human fossil record.

Early hominid dental pathology: interproximal caries in 1.5 million-year-old Paranthropus robustus from Swartkrans.

Carious lesions on the mesial and distal aspects of a RM1 of a young adult 'robust' australopithecine. Paranthropus robustus, are described. The specimen is from Member 1 sediments of the Swartkrans cave, which are dated to between about 1.8 and 1.5 million years before present. The mesial lesion, located cervically, had progressed further than the two distal lesions, which were situated along the cervix and in the enamel of the interproximal contact facet. This is the third specimen of P. robustus to be discovered with dental caries, but the other two cases were associated with occlusal and buccal enamel hypoplasia. Caries is present in less than about 3.0% of the approx. 116-128 P. robustus specimens at Swartkrans, and this incidence is noticeably lower than in the small Homo erectus sample from the same site. While this difference may simply be a sampling artefact, it may reflect dietary differences between these two synchronic and presumably sympatric early hominid species.

An evaluation of dental radiograph accuracy in the measurement of enamel thickness.

Many studies have employed lateral radiographs to measure the thickness of tooth enamel in recent human and fossil hominid samples, but the accuracy of measurements obtained by this technique has not been assessed. In this study, 20 isolated human maxillary permanent molars were radiographed using the parallel film technique. The crowns were then sectioned longitudinally through the tips of the buccal cusps. Measurements of enamel cap area, and of linear enamel thickness in the occlusal basin and over the metacone apex, were made from the radiographs and corresponding sectioned surfaces. Comparisons of the two sets of values revealed that radiographs generally overestimated enamel thickness but there was considerable variability in the error by which measurements from radiographs either under- or overestimated the true value. Lateral radiographs may provide a rough visual impression of whether a tooth has thin or thick enamel but they do not generally provide for accurate measurement of enamel thickness. Quantitative data on enamel thickness from studies that have employed lateral radiographs should be viewed with circumspection.

A review of interproximal wear grooves on fossil hominin teeth with new evidence from Olduvai Gorge.

Interproximal (approximal) grooves at the cementum-enamel junction of premolar and molar teeth have been observed in a broad range of human ancestors and related extinct species from 1.84 million years ago to the present. Many hypotheses have been presented to explain the aetiology of these grooves, though their form and positioning are most consistent with tooth-picking behaviours. This paper reviews occurrences of interproximal grooves in the cheek teeth of modern and fossil humans, evaluates hypotheses on their cause, and reports on a previously undescribed groove found in OH 60, a molar tooth from Olduvai Gorge. This specimen is among the earliest to show such grooving, and is most likely attributable to Homo erectus. It is concluded that, because interproximal grooves have been observed only on Homo teeth, they probably reflect a behaviour or behaviours unique to that genus.

Error rates in dental microwear quantification using scanning electron microscopy.

There is a degree of correlation between dietary habits and dental microwear in extant primates, and this has enabled inferences to be made about prehistoric diets. Several techniques have been used to quantify microwear, but the comparability of results derived from each has not been demonstrated. Moreover, neither intra- nor interobserver error rates in microwear quantification have been documented to date. We here assess intra- and interobserver error using Microware 4.0, and evaluate intertechnique comparability using the three methods that have been most widely employed in the field. This study documents an overall intraobserver error rate of about 7%, and an overall interobserver error rate of some 9%. Both intra- and interobserver error appears to be influenced substantially by the nature of the micrograph being measured. In no instance did the results obtained by different observers using Microware 4.0 differ significantly, and there was a reasonable degree of interobserver consistency in the rank ordering of micrographs in relation to any given parameter. The results obtained through the use of different quantitative techniques differed significantly, with an overall intertechnique error rate of approximately 19%. Several variables, including differences in magnification factor, scanning electron microscope kV settings, and specimen-detector relationships undoubtedly contribute to the differences among the three methods, but we were not able to assess their relative importance. Microwear quantification permits distinctions between broad dietary categories, but the margin of intra- and interobserver error should be taken into account when defining pattern differences between populations (or species) or when documenting seasonally mitigated differences within a taxon. In view of the error introduced by the use of different methods, we suggest that a consistent technique, such as offered by the Microware software package, be adopted by current researchers to establish a common microwear database.

Age-related variation in limb bone diaphyseal structure among Inuit foragers from Point Hope, northern Alaska.

UNLABELLED: Age-related deterioration of limb bone diaphyseal structure is documented among precontact Inuit foragers from northern Alaska. These findings challenge the concept that bone loss and fracture susceptibility among modern Inuit stem from their transition away from a physically demanding traditional lifestyle toward a more sedentary Western lifestyle. INTRODUCTION: Skeletal fragility is rare among foragers and other traditional-living societies, likely due to their high physical activity levels. Among modern Inuit, however, severe bone loss and fractures are apparently common. This is possibly because of recent Western influences and increasing sedentism. To determine whether compromised bone structure and strength among the Inuit are indeed aberrant for a traditional-living group, data were collected on age-related variation in limb bone diaphyseal structure from a group predating Western influences. METHODS: Skeletons of 184 adults were analyzed from the Point Hope archaeological site. Mid-diaphyseal structure was measured in the humerus, radius, ulna, femur, and tibia using CT. Structural differences were assessed between young, middle-aged, and old individuals. RESULTS: In all bones examined, both females and males exhibited significant age-related reductions in bone quantity. With few exceptions, total bone (periosteal) area did not significantly increase between young and old age in either sex, nor did geometric components of bending rigidity (second moments of area). CONCLUSIONS: While the physically demanding lifestyles of certain traditional-living groups may protect against bone loss and fracture susceptibility, this is not the case among the Inuit. It remains possible, however, that Western characteristics of the modern Inuit lifestyle exacerbate age-related skeletal deterioration.

Three-dimensional molar enamel distribution and thickness in Australopithecus and Paranthropus.

Thick molar enamel is among the few diagnostic characters of hominins which are measurable in fossil specimens. Despite a long history of study and characterization of Paranthropus molars as relatively 'hyper-thick', only a few tooth fragments and controlled planes of section (designed to be proxies of whole-crown thickness) have been measured. Here, we measure molar enamel thickness in Australopithecus africanus and Paranthropus robustus using accurate microtomographic methods, recording the whole-crown distribution of enamel. Both taxa have relatively thick enamel, but are thinner than previously characterized based on two-dimensional measurements. Three-dimensional measurements show that P. robustus enamel is not hyper-thick, and A. africanus enamel is relatively thinner than that of recent humans. Interspecific differences in the whole-crown distribution of enamel thickness influence cross-sectional measurements such that enamel thickness is exaggerated in two-dimensional sections of A. africanus and P. robustus molars. As such, two-dimensional enamel thickness measurements in australopiths are not reliable proxies for the three-dimensional data they are meant to represent. The three-dimensional distribution of enamel thickness shows different patterns among species, and is more useful for the interpretation of functional adaptations than single summary measures of enamel thickness.

Late Pleistocene human skull from Hofmeyr, South Africa, and modern human origins.

The lack of Late Pleistocene human fossils from sub-Saharan Africa has limited paleontological testing of competing models of recent human evolution. We have dated a skull from Hofmeyr, South Africa, to 36.2 +/- 3.3 thousand years ago through a combination of optically stimulated luminescence and uranium-series dating methods. The skull is morphologically modern overall but displays some archaic features. Its strongest morphometric affinities are with Upper Paleolithic (UP) Eurasians rather than recent, geographically proximate people. The Hofmeyr cranium is consistent with the hypothesis that UP Eurasians descended from a population that emigrated from sub-Saharan Africa in the Late Pleistocene.

Enamel thickness of deciduous and permanent molars in modern Homo sapiens.

This study presents data on the enamel thickness of deciduous (dm2) and permanent (M1-M3) molars for a geographically diverse sample of modern humans. Measurements were recorded from sections through the mesial cusps of unworn teeth. Enamel is significantly thinner on deciduous than on permanent molars, and there is a distinct trend for enamel to increase in relative thickness from M1 to M3. The relatively thicker enamel of M2s and especially M3s can be related to the overall reduction in size of more distal molar crowns, which has been attained through a differential loss of the dentine component. Enamel tends to be thicker on the protocone than on the paracone, and thicker on the protoconid than on the metaconid, but its distribution is not wholly concordant with models that predict increased thickness as a means by which to counter heavier attritional loss on these "functional" cusps. Indeed, the thickness of enamel tends to be more variable on cusp tips and occlusal surfaces than over the lateral aspects of cusps. The proportionately thicker enamel over the lateral aspects of the protocone and protoconid more likely serves as a means to prolong functional crown life by preventing cusp fracture, rather than being an adaptation to increase the attritional longevity of wear facets. The present data suggest that the human dentition is not predisposed to develop a helicoidal wear plane through the disposition of molar enamel thickness.

Molar enamel thickness in the chacma baboon, Papio ursinus (Kerr 1792).

Modern humans exhibit increasing relative enamel thickness from M1 to M3. Some biomechanical (basic lever) models predict that the more distal molars in humans encounter higher occlusal forces, and it has been postulated that this provides a functional explanation for the observed gradient in relative enamel thickness. However, constrained three-dimensional models and experimental observations suggest that there is a reduction in bite force potential from M1 to M3, which would be consistent with the tendency for humans to reduce the size of the distal molars. In this regard, it has been postulated that the distal increase in enamel thickness is a consequence of crown size reduction; thus, it is unnecessary to invoke functional scenarios to explain this phenomenon. We assess these competing proposals by examining relative enamel thickness in a catarrhine primate (Papio ursinus) that exhibits crown size increase from M1 to M3. The molar row of P. ursinus is positioned relatively far forward of the temporomandibular joint, which results in the baboon being able to exert relatively greater muscle forces during posterior biting in comparison to modern humans. Thus, a significant distalward gradient of increasing enamel thickness would be expected in P. ursinus according to the hypothesis that posits it to be functionally related to bite force. The present study reveals no significant difference in relative enamel thickness along the molar row in P. ursinus. This finding lends support to the notion that the relatively thicker enamel of human distal molars is related primarily to their reduction in size. This carries potential implications for the interpretation of enamel thickness in phylogenetic reconstructions: the relatively thick molar enamel shared by modern humans and some of our fossil relatives may not be strictly homologous, in that it may result from different underlying developmental mechanisms.

On the eruption pattern of the permanent incisors and first permanent molars in Paranthropus.

It has been claimed recently that Australopithecus exhibited a pattern of permanent tooth eruption like that of extant great apes, whereas a significantly different pattern was shared by Paranthropus and Homo (Dean, 1985). More particularly, each of the four Paranthropus specimens examined in that study was held to show advanced development and eruption of the permanent incisors relative to the first molar. It is demonstrated here that the eruption sequence that was posited for at least one of these four Paranthropus specimens (SK 61) is clearly erroneous, while the developmental/eruption sequences manifested by the other three specimens would appear to be more ambiguous than was claimed. Another juvenile specimen of Paranthropus (KNM-ER 1820) that was not included in Dean's study also does not necessarily support the eruption pattern that was said to characterize that taxon.

Quantitative analysis of occlusal microwear in Australopithecus and Paranthropus.

Quantitative analysis of microwear features preserved on the occlusal surfaces of the M2s of southern African specimens of Australopithecus and Paranthropus (the so-called 'gracile' and 'robust' australopithecines) reveals that there is no striking relationship in either taxon between occlusal facet inclination and the incidence of wear features. Within each taxon, Phase I and Phase II facets tend to differ in a similar manner in the total number of wear features, the percentage frequency of pitting, and in the orientation of wear scratches. Nevertheless, Paranthropus molars tend to display significantly greater numbers of microwear features on both Phase I and II facets than do Australopithecus homologues, and Paranthropus molars also evince significantly higher proportions of occlusal pitting on these surfaces. Paranthropus and Australopithecus crowns also differ significantly in the degree by which the occlusal wear scratches vary in their orientation. On each facet, Australopithecus tooth scratches display a greater degree of directional similarity. The differences between the Phase I and Phase II facets of Australopithecus and Paranthropus M2s suggest that the dietary items involved in the production of these observed patterns differed also. The diets of these Plio-Pleistocene hominids appear to have been qualitatively dissimilar.

New hominid fossils from the Swartkrans formation (1979-1986 excavations): craniodental specimens.

Seventy-two individually numbered hominid craniodental fossils from recent excavations at Swartkrans are described. All derive from in situ decalcified breccia and/or unconsolidated sediments. A total of 20 specimens, representing 13 to 16 individuals derive from Member 1 "Lower Bank," two teeth derive from sediments along the Member 1-2 Interface, 38 fossils representing 19 to 24 individuals come from Member 2, and 12 teeth representing 9 to 11 individuals derive from Member 3. All but four of the specimens are attributable at the generic level; one specimen from Member 1 "Lower Bank" and five specimens from Member 2 are attributed to Homo, while the others represent Paranthropus. The proportional representation of Homo in the Swartkrans Formation is markedly higher in Member 2 (c. 33%) than in the Member 1 "Lower Bank" (c. 8%) and Member 1 "Hanging Remnant" (c. 5%) samples.

Middle Stone Age human fossils from Die Kelders Cave 1, Western Cape Province, South Africa.

Die Kelders Cave 1 (DK1) preserves a thick series of Middle Stone Age (MSA) horizons that date to a fairly short temporal interval sometime between about 60 and 80 ka ago. Twenty-seven human fossils, comprising 24 isolated teeth, a mandibular fragment, and two manual middle phalanges derive from seven of the 12 layers. The vast majority are children, and all may have come from sub-adult individuals. The entire assemblage may represent a minimum of ten individuals. As might be expected for teeth of such antiquity, most of the DK1 crowns tend to be large in comparison to recent African homologues. They tend to be smaller than, albeit more similar in size to, the teeth of penecontemporaneous archaic populations from Eurasia. The majority of morphological variants displayed by the DK1 crowns characterize the teeth of recent sub-Saharan Africans, and the DK1 crowns resemble those of recent Africans in a number of traits that have been used to define a sub-Saharan African regional complex. The morphological similarities between the DK1 MSA and recent African teeth, however, do not necessarily signify a close evolutionary relationship between them, because these crowns variants appear to be plesiomorphic.

Radius of Paranthropus robustus from member 1, Swartkrans formation, South Africa.

Recently recovered hominid postcrania from Member 1, Swartkrans Formation include the proximal and distal ends of a right radius attributed to a single individual of Paranthropus robustus. These fossils are essentially similar to Australopithecus afarensis, A. africanus, and P. boisei homologues. The head manifests an ape-like circumferentia articularis, and the distal end has prominent medial, dorsal, and lateral tubercles and a well developed brachioradialis crest, features also commonly exhibited by extant great apes. The volar set of the P. robustus radiocarpal joint, like that of Australopithecus homologues, more closely resembles the neutral condition exhibited by Homo than the greater flexion evinced by living apes. Compared with fossil and recent specimens of Homo, the configuration of the P. robustus radial head suggests enhanced stability against medial displacement during pronation and supination; the strong crest for the attachment of brachioradialis may attest to enhanced forearm flexor capability. In addition, this crest and the prominent dorsal tubercles may indicate enhanced hand extensor and, therefore, hand flexor capabilities. The differences in radial morphology between Paranthropus and Homo may relate to significant behavioral differences between these two synchronic taxa.

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.

Scanning electron microscopy of early dinosaur egg shell structure: a comparison with other rigid sauropsid eggs.

Fossil eggs attributable to dinosaur (probably prosauropod) parentage that have been recovered from the early Jurassic Elliot Formation sediments at the Rooidraai locality possess shells that are similar to those of birds and crocodilians, and distinctly unlike those of chelonians and gekkonids. The preserved shell is very thin, and distinct mammillary processes are lacking, although the inner surface displays an undulating contour. The absence of these processes may be attributable to the inner portion of the shell having been at least partially decalcified during incubation and not preserved in the fossil state. The shells appear to be composed of broadly wedge-shaped, albeit ill-defined calcareous units, and they are similar to those of birds and other dinosaurs in the pattern of cleavage shown by the tabular calcite crystals of the palisade layer, and in the absence of the dominant horizontal lamellae that characterize crocodilian shells. The differential resemblance of these early Jurassic shells to the eggs of other closely related sauropsid taxa may be pertinent to questions concerning the evolution of egg shell structure within this clade.