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.

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.

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.

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.

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.

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.

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 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.

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.

Human remains from Blombos Cave, South Africa: (1997-1998 excavations).

The Middle Stone Age (MSA) layers at Blombos Cave contain abundant bifacial Still Bay points, formal and ad hoc bone artefacts, and an intentionally incised bone piece. These artefacts add weight to arguments that some aspects of modern human behavior developed earlier in sub-Saharan Africa than elsewhere. Four human teeth were recovered from the MSA strata at Blombos during the 1997-1998 excavations. Two are heavily worn deciduous teeth, and two are incomplete permanent premolar crowns. The Blombos di(1)is comparatively large in relation to modern African homologues, falling within the lower part of the observed Neandertal range. The dm(1)and P(3)are comparable to modern teeth and smaller than most Neandertal crowns. The premolars preserve horizontal circum-cervical striae that suggest palliative toothpick use. The di(1)evinces labial scratches that resemble neither the "cutmarks" that have been observed on Neandertal incisors, nor the striae that have been recorded on modern human teeth.

Middle Stone Age stratigraphy and excavations at Die Kelders Cave 1 (Western Cape Province, South Africa): the 1992, 1993, and 1995 field seasons.

Die Kelders Cave 1, first excavated under the direction of Franz Schweitzer in 1969-1973, was re-excavated between 1992 and 1995 by a combined team from the South African Museum, SUNY at Stony Brook, and Stanford University. These renewed excavations enlarged the artefactual and faunal samples from the inadequately sampled and less intensively excavated lower Middle Stone Age (MSA) layers, increased our understanding of the complex site formation processes within the cave, enlarged the hominid sample from the MSA deposits, and generated ESR, TL, and OSL dates for the MSA layers. Importantly, these new excavations dramatically improved our comprehension of the vertical and lateral characteristics of the MSA stratigraphy. Surface plotting of the MSA layers has led to the identification of at least two major zones of subsidence that significantly warped the layers, draping some along the eroding surface contours of major blocks of fallen limestone roof rock. A third zone of subsidence is probably present in the older excavations. Dramatic roof falls of very large limestone blocks occurred at least twice-once in the middle of Layer 4/5 where the roof blocks were only slightly weathered after collapse, and at the top of Layer 6 where the blocks weathered heavily after collapse, producing a zone of decomposed rock around the blocks. Many of the sandy strata are cut by small and localized faults and slippages. All of the strata documented by Schweitzer's excavations are present throughout the exposed area to the west of his excavated area, where many of them thicken and become more complex. Layer 6, the thickest MSA layer, becomes less diagenetically altered and compressed to the west.

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.

Additional human fossils from Klasies River Mouth South Africa.

A fragmentary temporal bone and partial atlas from the Middle Stone Age (MSA) at Klasies River Mouth (KRM) are described and analyzed. The atlas (SAM-AP 6268) is comparable to Levantine "Early Modern", Neandertal and recent human vertebrae. The temporal (SAM-AP 6269) is similar to recent African homologues except that the posteromedial wall of the glenoid fossa is composed entirely of the squamous temporal, a situation that appears to be infrequent among other Pleistocene fossils. The KRM glenoid fossa is also mediolateraly broad and anteroposteriorly short in comparison with many, but not, all recent specimens. Nevertheless, the KRM temporal is decidedly modern, both morphologically and metrically, by comparison with other Pleistocene specimens. The limited evidence provided by this bone is consistent with that of other MSA cranial remains from this site in suggesting an overall, if somewhat ambiguous pattern of morphological modernity.

Re-analysis of the hominid radii from Cave of Hearths and Klasies River Mouth, South Africa.

Two of the few postcranial fragments from the late Early Stone Age and/or the Middle Stone Age of southern Africa are the proximal radii from the Cave of Hearths and Klasies River Mouth. The Cave of Hearths fossil is metrically indistinguishable from both archaic (e.g., Neandertals) and recent humans, and presents a mosaic of primitive and modern features. The primitive include a relatively slender neck and thick cortical bone (the latter of which distinguishes recent humans from archaic, Early Modern, and Upper Paleolithic hominids); the modern includes an anteromedially (rather than medially) facing radial tuberosity. Its extreme collo-diaphyseal angle is unusual, although it can be matched by modern homologues. The neck-shaft angle of some Neandetral and Early Modern radii also appears to match that of the Cave of Hearths specimen. The Klasies River Mouth radius also has thick cortical bone of the neck. It is morphologically indistinguishable from Early Modern and Neandertal homologues. These, and other fossils, suggest a mosaic pattern of evolution in the postcranial skeleton of the late Early Stone Age and/or Middle Stone Age inhabitants of sub-Saharan Africa.

A reappraisal of early hominid phylogeny.

We report here on the results of a new cladistic analysis of early hominid relationships. Ingroup taxa included Australopithecus afarensis, Australopithecus africanus, Australopithecus aethiopicus, Australopithecus robustus, Australopithecus boisei, Homo habilis, Homo rudolfensis, Homo ergaster and Homo sapiens. Outgroup taxa included Pan troglodytes and Gorilla gorilla. Sixty craniodental characters were selected for analysis. These were drawn from the trait lists of other studies and our own observations. Eight parsimony analyses were performed that differed with respect to the number of characters examined and the manner in which the characters were treated. Seven employed ordered characters, and included analyses in which (1) taxa that were variable with respect to a character were coded as having an intermediate state, (2) characters with variable states in any taxon were excluded; (3) a variable taxon was coded as having the state exhibited by the majority of its hypodigm, (4) variable taxa were coded as missing data for that character, (5) some characters were considered irreversible, (6) masticatory characters were excluded, and (7) characters whose states were unknown in some taxa were excluded. In the final analysis, (8) all characters were unordered. All analyses were performed using PAUP 3.0s. Despite the fact that the eight analyses differed with respect to methodology, they produced several consistent results. All agreed that the "robust" australopithecines form a clade, A. afarensis is the sister taxon of all other hominids, and the genus Australopithecus, as conventionally defined, is paraphyletic. All eight also supported trees in which A. africanus is the sister taxon of a joint Homo+ "robust" clade, although in one analysis an equally parsimonious topology found A. africanus to be the sister of the "robust" species. In most analyses, the relationships of A. africanus and H. habilis were unstable, in the sense that their positions vary in trees that are marginally less parsimonious than the favored one. Trees in which "robust" australopithecines are paraphyletic were found to be extremely unparsimonious.

Fossil Homo femur from Berg Aukas, northern Namibia.

The proximal half of a hominid femur was recovered from deep within a paleokarst feature at the Berg Aukas mine, northern Namibia. The femur is fully mineralized, but it is not possible to place it in geochronological context. It has a very large head, an exceptionally thick diaphyseal cortex, and a very low collodiaphyseal angle, which serve to differentiate it from Holocene homologues. The femur is not attributable to Australopithecus, Paranthropus, or early Homo (i.e., H. habilis sensu lato). Homo erectus femora have a relatively longer and AP flatter neck, and a shaft that exhibits less pilaster than the Berg Aukas specimen. Berg Aukas also differs from early modern femora in several features, including diaphyseal cortical thickness and the degree of subtrochanteric AP flattening. The massive diaphyseal cortex of Berg Aukas finds its closest similarity within archaic H. sapiens (e.g., Castel di Guido) and H. erectus (e.g., KNM-ER 736) samples. It has more cortical bone at midshaft than any other specimen, although relative cortical thickness and the asymmetry of its cross-sectional disposition at this level are comparable with those of other Pleistocene femora. The closest morphological comparisons with Berg Aukas are in archaic (i.e., Middle Pleistocene) H. sapiens and Neandertal samples.

Taxonomic affinity of the early Homo cranium from Swartkrans, South Africa.

A quantitative analysis that employs randomization methods and distance statistics has been undertaken in an attempt to clarify the taxonomic affinities of the partial Homo cranium (SK 847) from Member 1 of the Swartkrans Formation. Although SK 847 has been argued to represent early H. erectus, exact randomization tests reveal that the magnitude of differences between it and two crania that have been attributed to that taxon (KNM-ER 3733 and KNM-WT 15000) is highly unlikely to be encountered in a modern human sample drawn from eastern and southern Africa. Some of the variables that differentiate SK 847 from the two early H. erectus crania (e.g., nasal breadth, frontal breadth, mastoid process size) have been considered to be relevant characters in the definition of that taxon. Just as the significant differences between SK 847 and the two early H. erectus crania make attribution of the Swartkrans specimen to that taxon unlikely, the linkage of SK 847 to KNM-ER 1813, and especially Stw 53, suggests that the Swartkrans cranium may have its closest affinity with H. habilis sensu lato. Differences from KNM-ER 1813, however, hint that the South African fossils may represent a species of early Homo that has not been sampled in the Plio-Pleistocene of eastern Africa. The similarity of SK 847 and Stw 53 may support faunal evidence which suggests that Sterkfontein Member 5 and Swartkrans Member 1 are of similar geochronological age.