A new species of Homo from the Late Pleistocene of the Philippines.

A hominin third metatarsal discovered in 2007 in Callao Cave (Northern Luzon, the Philippines) and dated to 67 thousand years ago provided the earliest direct evidence of a human presence in the Philippines. Analysis of this foot bone suggested that it belonged to the genus Homo, but to which species was unclear. Here we report the discovery of twelve additional hominin elements that represent at least three individuals that were found in the same stratigraphic layer of Callao Cave as the previously discovered metatarsal. These specimens display a combination of primitive and derived morphological features that is different from the combination of features found in other species in the genus Homo (including Homo floresiensis and Homo sapiens) and warrants their attribution to a new species, which we name Homo luzonensis. The presence of another and previously unknown hominin species east of the Wallace Line during the Late Pleistocene epoch underscores the importance of island Southeast Asia in the evolution of the genus Homo.

Dental data challenge the ubiquitous presence of Homo in the Cradle of Humankind.

The origins of Homo, as well as the diversity and biogeographic distribution of early Homo species, remain critical outstanding issues in paleoanthropology. Debates about the recognition of early Homo, first appearance dates, and taxonomic diversity within Homo are particularly important for determining the role that southern African taxa may have played in the origins of the genus. The correct identification of Homo remains also has implications for reconstructing phylogenetic relationships between species of Australopithecus and Paranthropus, and the links between early Homo species and Homo erectus. We use microcomputed tomography and landmark-free deformation-based three-dimensional geometric morphometrics to extract taxonomically informative data from the internal structure of postcanine teeth attributed to Early Pleistocene Homo in the southern African hominin-bearing sites of Sterkfontein, Swartkrans, Drimolen, and Kromdraai B. Our results indicate that, from our sample of 23 specimens, only 4 are unambiguously attributed to Homo, 3 of them coming from Swartkrans member 1 (SK 27, SK 847, and SKX 21204) and 1 from Sterkfontein (Sts 9). Three other specimens from Sterkfontein (StW 80 and 81, SE 1508, and StW 669) approximate the Homo condition in terms of overall enamel-dentine junction shape, but retain Australopithecus-like dental traits, and their generic status remains unclear. The other specimens, including SK 15, present a dominant australopith dental signature. In light of these results, previous dietary and ecological interpretations can be reevaluated, showing that the geochemical signal of one tooth from Kromdraai (KB 5223) and two from Swartkrans (SK 96 and SKX 268) is consistent with that of australopiths.

Reassessment of the phylogenetic relationships of the late Miocene apes Hispanopithecus and Rudapithecus based on vestibular morphology.

Late Miocene great apes are key to reconstructing the ancestral morphotype from which earliest hominins evolved. Despite consensus that the late Miocene dryopith great apes Hispanopithecus laietanus (Spain) and Rudapithecus hungaricus (Hungary) are closely related (Hominidae), ongoing debate on their phylogenetic relationships with extant apes (stem hominids, hominines, or pongines) complicates our understanding of great ape and human evolution. To clarify this question, we rely on the morphology of the inner ear semicircular canals, which has been shown to be phylogenetically informative. Based on microcomputed tomography scans, we describe the vestibular morphology of Hispanopithecus and Rudapithecus, and compare them with extant hominoids using landmark-free deformation-based three-dimensional geometric morphometric analyses. We also provide critical evidence about the evolutionary patterns of the vestibular apparatus in living and fossil hominoids under different phylogenetic assumptions for dryopiths. Our results are consistent with the distinction of Rudapithecus and Hispanopithecus at the genus rank, and further support their allocation to the Hominidae based on their derived semicircular canal volumetric proportions. Compared with extant hominids, the vestibular morphology of Hispanopithecus and Rudapithecus most closely resembles that of African apes, and differs from the derived condition of orangutans. However, the vestibular morphologies reconstructed for the last common ancestors of dryopiths, crown hominines, and crown hominids are very similar, indicating that hominines are plesiomorphic in this regard. Therefore, our results do not conclusively favor a hominine or stem hominid status for the investigated dryopiths.

Estimates of absolute crown strength and bite force in the lower postcanine dentition of Gigantopithecus blacki.

Gigantopithecus blacki is hypothesized to have been capable of processing mechanically challenging foods, which likely required this species to have high dental resistance to fracture and/or large bite force. To test this hypothesis, we used two recently developed approaches to estimate absolute crown strength and bite force of the lower postcanine dentition. Sixteen Gigantopithecus mandibular permanent cheek teeth were scanned by micro-computed tomography. From virtual mesial cross-sections, we measured average enamel thickness and bi-cervical diameter to estimate absolute crown strength, and cuspal enamel thickness and dentine horn angle to estimate bite force. We compared G. blacki with a sample of extant great apes (Pan, Pongo, and Gorilla) and australopiths (Australopithecus anamensis, Australopithecus afarensis, Australopithecus africanus, Paranthropus robustus, and Paranthropus boisei). We also evaluated statistical differences in absolute crown strength and bite force between the premolars and molars for G. blacki. Results reveal that molar crown strength is absolutely greater, and molar bite force absolutely higher, in G. blacki than all other taxa except P. boisei, suggesting that G. blacki molars have exceptionally high resistance to fracture and the ability to generate exceptionally high bite force. In addition, G. blacki premolars have comparable absolute crown strength and larger bite force capabilities compared with its molars, implying possible functional specializations in premolars. The dental specialization of G. blacki could thus represent an adaptation to further facilitate the processing of mechanically challenging foods. While it is currently not possible to determine which types of foods were actually consumed by G. blacki through this study, direct evidence (e.g. dental chipping and microwear) left by the foods eaten by G. blacki could potentially lead to greater insights into its dietary ecology.

A reassessment of the distinctiveness of dryopithecine genera from the Iberian Miocene based on enamel-dentine junction geometric morphometric analyses.

A vast diversity of catarrhines primates has been uncovered in the Middle to Late Miocene (12.5-9.6 Ma) of the Vallès-Penedès Basin (northeastern Spain), including several hominid species (Pierolapithecus catalaunicus, Anoiapithecus brevirostris, Dryopithecus fontani, Hispanopithecus laietanus, and Hispanopithecus crusafonti) plus some remains attributed to 'Sivapithecus' occidentalis (of uncertain taxonomic validity). However, Pierolapithecus and Anoiapithecus have also been considered junior synonyms of Dryopithecus by some authors, which entail a lower generic diversity and an inflated intrageneric variation of the latter genus. Since the distinction of these taxa partly relies on dental features, the detailed and quantitative analysis of tooth shape might help disentangling the taxonomic diversity of these Miocene hominids. Using diffeomorphic surface matching and three-dimensional geometric morphometrics, we investigate the enamel-dentine junction shape (which is a reliable taxonomic proxy) of these Miocene hominids, with the aim of investigating their degree of intra- and intergeneric variation compared with that of extant great ape genera. We conducted statistical analyses, including between-group principal component analyses, canonical variate analyses, and permutation tests, to investigate whether the individual and combined (i.e., Dryopithecus s.l.) variation of the extinct genera exceeds that of the extant great apes. Our results indicate that Pierolapithecus, Anoiapithecus, Dryopithecus, and Hispanopithecus show morphological differences of enamel-dentine junction shape relative to the extant great apes that are consistent with their attribution to different genera. Specifically, the variation displayed by the Middle Miocene taxa combined exceeds that of extant great ape genera, thus undermining the single-genus hypothesis. 'Sivapithecus' occidentalis specimens fall close to Dryopithecus but in the absence of well-preserved comparable teeth for Pierolapithecus and Anoiapithecus, their taxonomic attribution remains uncertain. Among the Hispanopithecus sample, IPS1802 from Can Llobateres stands out and might either be an outlier in terms of morphology, or represent another dryopithecine taxon.

Early Pleistocene hominin teeth from Gongwangling of Lantian, Central China.

The fossil hominin individual from Gongwangling of Lantian, Central China, represents one of the earliest members attributed to Homo erectus in East Asia. Recent paleomagnetic analyses have yielded an age of 1.63 Ma for the Gongwangling hominin. The fossils from this site are critical to characterize the morphological features of early hominins in East Asia and to understand their relationships with other earlier and later members of the genus Homo. However, most morphological details of the Gongwangling cranium were obliterated due to postmortem erosion and deformation. Here we used high-resolution microcomputed tomography and three-dimensional virtual imaging techniques to extract the teeth and reconstruct the worn/damaged areas, describe the external morphology, measure crown diameters, record nonmetric traits of the crown and root, and investigate the shape of the enamel-dentine junction using geometric morphometrics. We compared the data obtained from the six teeth of the Gongwangling hominin with African early Homo, African and Georgian Homo erectus s.l., Asian Homo erectus, Homo antecessor, pre-Neanderthals, Neanderthals, and modern humans. Our results show that the Gongwangling specimens display affinities with other specimens attributed to H. erectus s.l. The highly divergent and noncoalesced three-root system in the Gongwangling specimens is comparable to that in the Early Pleistocene members of H. erectus s.l., and differs from Middle Pleistocene representatives of the species. The enamel-dentine junction shape of the Gongwangling molars prefigures the Asian H. erectus pattern later found in East Asian Middle Pleistocene H. erectus. The morphological comparisons between East Asian Early Pleistocene (e.g., Gongwangling, Meipu, and Quyuan River Mouth) and Middle Pleistocene H. erectus (e.g., Zhoukoudian, Hexian, and Yiyuan) suggest a potential temporal trend within this species in East Asia.

Further analyses of the structural organization of Homo luzonensis teeth: Evolutionary implications.

The species Homo luzonensis has recently been described based on a set of dental and postcranial elements found at Callao Cave (Northern Luzon, Philippines) and dated to at least 50-67 ka. Seven postcanine maxillary teeth are attributed to this taxon, five of them belonging to the same individual (CCH6) and representing the holotype of H. luzonensis, whereas the isolated upper premolar CCH8 and the upper third molar CCH9 are paratypes of the species. The teeth are characterized by their small dimensions associated with primitive features, as also found in Homo floresiensis, another hominin having evolved in an insular environment of Southeast Asia. Postcranial bones of the hands and feet of H. luzonensis and H. floresiensis show Homo habilis-like or australopith-like features, whereas cranial and dental morphology are more consistent with the Asian Homo erectus morphology. Due to this mosaic morphology, the origin and phylogenetic relationships of both H. luzonensis and H. floresiensis are still debated. To test the hypotheses that H. luzonensis derives from H. erectus or from an earlier small-brained hominin, we analyzed the µCT scans of the teeth. We investigated both external and internal tooth structure using morphometric methods including: crown outline shape, tooth crown tissue proportions, enamel-dentine junction shape, and pulp morphology. Homo luzonensis external crown morphology aligns more with H. erectus than with H. habilis/H. rudolfensis. The internal structural organization of H. luzonensis teeth exhibits more affinities with that of H. erectus and H. floresiensis than with Neanderthals and modern humans. Our results suggest that both H. floresiensis and H. luzonensis likely evolved from some H. erectus groups that dispersed in the various islands of this region and became isolated until endemic speciation events occurred at least twice during the Pleistocene in insular environments.

A comparative analysis of the vestibular apparatus in Epipliopithecus vindobonensis: Phylogenetic implications.

Pliopithecoids are an extinct group of catarrhine primates from the Miocene of Eurasia. More than 50 years ago, they were linked to hylobatids due to some morphological similarities, but most subsequent studies have supported a stem catarrhine status, due to the retention of multiple plesiomorphic features (e.g., the ectotympanic morphology) relative to crown catarrhines. More recently, some morphological similarities to hominoids have been noted, raising the question of whether they could be stem members of this clade. To re-evaluate these competing hypotheses, we examine the morphology of the semicircular canals of the bony labyrinth of the middle Miocene pliopithecid Epipliopithecus vindobonensis. The semicircular canals are suitable to test between these hypotheses because (1) they have been shown to embed strong phylogenetic signal and reliably discriminate among major clades; (2) several potential hominoid synapomorphies have been identified previously in the semicircular canals; and (3) semicircular canal morphology has not been previously described for any pliopithecoid. We use a deformation-based (landmark-free) three-dimensional geometric morphometric approach to compare Epipliopithecus with a broad primate sample of extant and extinct anthropoids. We quantify similarities in semicircular canal morphology using multivariate analyses, reconstruct ancestral morphotypes by means of a phylomorphospace approach, and identify catarrhine and hominoid synapomorphies based on discrete characters. Epipliopithecus semicircular canal morphology most closely resembles that of platyrrhines and Aegyptopithecus due to the retention of multiple anthropoid symplesiomorphies. However, Epipliopithecus is most parsimoniously interpreted as a stem catarrhine more derived than Aegyptopithecus due to the possession of a crown catarrhine synapomorphy (i.e., the rounded anterior canal), combined with the lack of other catarrhine and any hominoid synapomorphies. Some similarities with hylobatids and atelids are interpreted as homoplasies likely related to positional behavior. The semicircular canal morphology of Epipliopithecus thus supports the common view that pliopithecoids are stem catarrhines.

Trophic ecology of a Late Pleistocene early modern human from tropical Southeast Asia inferred from zinc isotopes.

Tam Pà Ling, a cave site in northeastern Laos, has yielded the earliest skeletal evidence of Homo sapiens in mainland Southeast Asia. The reliance of Pleistocene humans in rainforest settings on plant or animal resources is still largely unstudied, mainly due to poor collagen preservation in fossils from tropical environments precluding stable nitrogen isotope analysis, the classical trophic level proxy. However, isotopic ratios of zinc (Zn) in bioapatite constitute a promising proxy to infer trophic and dietary information from fossil vertebrates, even under adverse tropical taphonomic conditions. Here, we analyzed the zinc isotope composition (66Zn/64Zn expressed as δ66Zn value) in the enamel of two teeth of the Late Pleistocene (63-46 ka) H. sapiens individual (TPL1) from Tam Pà Ling, as well as 76 mammal teeth from the same site and the nearby Nam Lot cave. The human individual exhibits relatively low enamel δ66Zn values (+0.24‰) consistent with an omnivorous diet, suggesting a dietary reliance on both plant and animal matter. These findings offer direct evidence of the broad utilization of resources from tropical rainforests by one of the earliest known anatomically modern humans in Southeast Asia.

A deep-learning-based workflow to assess taxonomic affinity of hominid teeth with a test on discriminating Pongo and Homo upper molars.

OBJECTIVES: Convolutional neural network (CNN) is a state-of-art deep learning (DL) method with superior performance in image classification. Here, a CNN-based workflow is proposed to discriminate hominid teeth. Our hope is that this method could help confirm otherwise questionable records of Homo from Pleistocene deposits where there is a standing risk of mis-attributing molars of Pongo to Homo. METHODS AND MATERIALS: A two-step workflow was designed. The first step is converting the enamel-dentine junction (EDJ) into EDJ card, that is, a two-dimensional image conversion of the three-dimensional EDJ surface. In this step, researchers must carefully orient the teeth according to the cervical plane. The second step is training the CNN learner with labeled EDJ cards. A sample consisting of 53 fossil Pongo and 53 Homo (modern human and Neanderthal) was adopted to generate EDJ cards, which were then separated into training set (n = 84) and validation set (n = 22). To assess the feasibility of this workflow, a Pongo-Homo classifier was trained from the aforementioned EDJ card set, and then the classifier was used to predict the taxonomic affinities of six samples (test set) from von Koenigswald's Chinese Apothecary collection. RESULTS: Results show that EDJ cards in validation set are classified accurately by the CNN learner. More importantly, taxonomic predictions for six specimens in test set match well with the diagnosis results deduced from multiple lines of evidence, implying the great potential of CNN method. DISCUSSION: This workflow paves a way for future studies using CNN to address taxonomic complexity (e.g., distinguishing Pongo and Homo teeth from the Pleistocene of Asia). Further improvements include visual interpretation and extending the applicability to moderately worn teeth.

A robust alternative to assessing three-dimensional relative enamel thickness for the use in taxonomic assessment.

OBJECTIVE: Three-dimensional relative enamel thickness (3DRET) is important for assessing hypotheses about taxonomy, phylogeny, and dietary reconstruction for primates. However, its weaknesses have not been thoroughly investigated. Here, we analyze its weaknesses and propose an index aiming at better taxonomic discrimination. MATERIALS AND METHODS: The dimensionless 3D index, ratio of enamel-thickness to dentine-thickness (3DRED), which is defined as the cubic root of the ratio of 3D average enamel thickness (3DAET) to 3D average dentine thickness (3DADT), is proposed here. To compare 3DRET and 3DRED and their sensitivity to voxel size, a fossil orangutan molar was scanned 14 times with different resolutions ranging from 10 to 50 µm. Enamel thickness analysis was carried out for each resultant digital model. In addition, enamel thickness measurements of 179 mandibular permanent molars (eight genera) were analyzed, followed by investigating the relationship between 3DRET and 3DAET and between 3DRED and 3DAET. RESULTS: Regarding sensitivity, 3DRED is more robust than 3DRET. In addition, 3DRET is correlated with 3DAET by linear curve with regression coefficients approximating or larger than 0.8 in most cases, while 3DRED shows less correlation with 3DAET. Furthermore, there are clear separations between different taxa in the bivariate plot of 3DRED against 3DAET, indicative of the taxonomic value of 3DRED. CONCLUSION: Under certain conditions, 3DRED promises to be a robust and reliable alternative to 3DRET in taxonomic study.

Hominin diversity in East Asia during the Middle Pleistocene: A premolar endostructural perspective.

Following the recent studies of East Asian mid-Middle to early Late Pleistocene hominin material, a large spectrum of morphological diversity has been recognized and the coexistence of archaic ('Homo erectus-like') and derived ('modern-like') dental morphological patterns has been highlighted. In fact, for most of these Chinese fossils, generally categorized as 'archaic Homo sapiens' or 'post-H. erectus Homo', the taxonomic attribution is a matter of contention. With the help of µCT techniques and a deformation-based 3D geometric morphometric approach, we focused on the morphological variation in the enamel-dentine junction (EDJ) of 18 upper and lower premolars from Chinese Middle Pleistocene hominins. We then compared our results with a number of fossil and modern human groups, including Early Pleistocene H. erectus from Sangiran; late Early Pleistocene hominins from Tighenif, Algeria; classic Neanderthals; and modern humans. Our results highlight an evolutionary/chronological trend of crown base reduction, elevation of EDJ topography, and EDJ surface simplification in the hominin groups studied here. Moreover, this study brings insights to the taxonomy/phylogeny of 6 late Middle Pleistocene specimens whose evolutionary placement has been debated for decades. Among these specimens, Changyang premolars show features that can be aligned with the Asian H. erectus hypodigm, whereas Panxian Dadong and Tongzi premolars are more similar to Late Pleistocene Homo. Compared with early to mid-Middle Pleistocene hominins in East Asia, late Middle Pleistocene hominins evince an enlarged morphological variation. A persistence of archaic morphotypes and possible admixture among populations during the late Middle Pleistocene are discussed.

Reassessment of the TM 1517 odonto-postcranial assemblage from Kromdraai B, South Africa, and the maturational pattern of Paranthropus robustus.

OBJECTIVES: The Pleistocene taxon Paranthropus robustus was established in 1938 following the discovery at Kromdraai B, South Africa, of the partial cranium TM 1517a and associated mandible TM 1517b. Shortly thereafter, a distal humerus (TM 1517g), a proximal ulna (TM 1517e), and a distal hallucial phalanx (TM 1517k) were collected nearby at the site, and were considered to be associated with the holotype. TM 1517a-b represents an immature individual; however, no analysis of the potentially associated postcranial elements has investigated the presence of any endostructural remnant of recent epiphyseal closure. This study aims at tentatively detecting such traces in the three postcranial specimens from Kromdraai B. MATERIALS AND METHODS: By using µXCT techniques, we assessed the developmental stage of the TM 1517b's C-M3 roots and investigated the inner structure of TM 1517g, TM 1517e, and TM 1517k. RESULTS: The M2 shows incompletely closed root apices and the M3 a half-completed root formation stage. The distal humerus was likely completely fused, while the proximal ulna and the distal hallucial phalanx preserve endosteal traces of the diaphyseo-epiphyseal fusion process. DISCUSSION: In the hominin fossil record, there are few unambiguously associated craniodental and postcranial remains sampling immature individuals, an essential condition for assessing the taxon-specific maturational patterns. Our findings corroborate the original association of the craniodental and postcranial remains representing the P. robustus type specimen. As with other Plio-Pleistocene hominins, the odonto-postcranial maturational pattern of TM 1517 more closely fits an African great ape rather than the extant human pattern.

Structural analysis of premolar roots in Middle Pleistocene hominins from China.

This study investigates permanent maxillary and mandibular premolar root structural organization in East Asian Middle Pleistocene hominins. In addition to reporting and analyzing the linear and volumetric properties of the roots, we used a landmark-free approach to both qualify and quantify in 3D premolar root shape variation of Middle Pleistocene hominins in East Asia. Moreover, we focus on some mid-to late East Asian Middle Pleistocene hominin specimens whose taxonomic attribution is unclear. We find considerable cementum in this sample of hominins, similar to other fossil groups, but clearly different from modern humans which have a very small amount of cementum. Additionally, a smaller root pulp cavity is found in later Homo (Neanderthals and modern humans). Our analyses on the crown-root surface area ratio show that East Asian Middle Pleistocene Homo erectus as well as one late Middle Pleistocene Homo sp. specimen (PA 81 P4 from Changyang site) are distinguished from other fossil and extant groups by a relatively larger root surface, stout root branches and thick cementum deposits. This may represent a distinct East Asian H. erectus dental pattern. Geometric morphometric analyses on the external root surface reveal a general trend of shape simplification along the Homo lineage examined here, and distinguish Early Pleistocene Homo, Middle Pleistocene H. erectus, Neanderthals and modern human morphologies. The late Middle Pleistocene teeth from Changyang site (PA 76 P3 and PA 81 P4) are close to East Asian H. erectus and Neanderthals, while the mid-Middle Pleistocene P3 from Panxian Dadong falls within the modern human distribution. Combined with dental crown morphology and root number/form reported in previous studies, our results show that the external root shape can be considered a taxonomically relevant indicator. In general, an evolutionary tendency towards modern human morphology is observed in part of the East Asian Middle Pleistocene specimens, while a retention of primitive, H. erectus-like features is expressed in some late Middle Pleistocene specimens, supporting a multi-lineage and discontinuous scenario of human settlements in East Asia.

Dental macrowear and cortical bone distribution of the Neanderthal mandible from Regourdou (Dordogne, Southwestern France).

Tooth wear is an important feature for reconstructing diet, food processing and cultural habits of past human populations. In particular, occlusal wear facets can be extremely useful for detecting information about diet and non-masticatory behaviors. The aim of this study is to reconstruct the diet and cultural behavior of the Neanderthal specimen Regourdou 1 (Dordogne, Southern France) from the analysis of the macrowear pattern, using the occlusal fingerprint analysis method. In addition, we have also examined whether there is any association between the observed dental macrowear and mandibular bone distribution and root dentine thickness. The posterior dentition of Regourdou 1 is characterized by an asymmetric wear pattern, with the right side significantly more worn than the left. In contrast, the left lower P3 shows a more advanced wear than the right premolar, with unusual semicircular enamel wear facets. The results from occlusal fingerprint analysis of this unique pattern suggest tooth-tool uses for daily task activities. Moreover, the left buccal aspect of the mandibular cortical bone is thicker than its right counterpart, and the left P3 has a thicker radicular dentine layer than its antimere. These results show a certain degree of asymmetry in cortical bone topography and dentine tissue that could be associated with the observed dental macrowear pattern. The molar macrowear pattern also suggests that Regourdou 1 had a mixed diet typical of those populations living in temperate deciduous woodlands and Mediterranean habitats, including animal and plant foods. Although this study is limited to one Neanderthal individual, future analyses based on a larger sample may further assist us to better understand the existing relationship between mandibular architecture, occlusal wear and the masticatory apparatus in humans.

Efficacy of diffeomorphic surface matching and 3D geometric morphometrics for taxonomic discrimination of Early Pleistocene hominin mandibular molars.

Morphometric assessments of the dentition have played significant roles in hypotheses relating to taxonomic diversity among extinct hominins. In this regard, emphasis has been placed on the statistical appraisal of intraspecific variation to identify morphological criteria that convey maximum discriminatory power. Three-dimensional geometric morphometric (3D GM) approaches that utilize landmarks and semi-landmarks to quantify shape variation have enjoyed increasingly popular use over the past twenty-five years in assessments of the outer enamel surface (OES) and enamel-dentine junction (EDJ) of fossil molars. Recently developed diffeomorphic surface matching (DSM) methods that model the deformation between shapes have drastically reduced if not altogether eliminated potential methodological inconsistencies associated with the a priori identification of landmarks and delineation of semi-landmarks. As such, DSM has the potential to better capture the geometric details that describe tooth shape by accounting for both homologous and non-homologous (i.e., discrete) features, and permitting the statistical determination of geometric correspondence. We compare the discriminatory power of 3D GM and DSM in the evaluation of the OES and EDJ of mandibular permanent molars attributed to Australopithecus africanus, Paranthropus robustus and early Homo sp. from the sites of Sterkfontein and Swartkrans. For all three molars, classification and clustering scores demonstrate that DSM performs better at separating the A. africanus and P. robustus samples than does 3D GM. The EDJ provided the best results. P. robustus evinces greater morphological variability than A. africanus. The DSM assessment of the early Homo molar from Swartkrans reveals its distinctiveness from either australopith sample, and the "unknown" specimen from Sterkfontein (Stw 151) is notably more similar to Homo than to A. africanus.

A Neanderthal from the Central Western Zagros, Iran. Structural reassessment of the Wezmeh 1 maxillary premolar.

Wezmeh Cave, in the Kermanshah region of Central Western Zagros, Iran, produced a Late Pleistocene faunal assemblage rich in carnivorans along with a human right maxillary premolar, Wezmeh 1, an unerupted tooth from an 8 ± 2 year-old individual. Uranium-series analyses of the fauna by alpha spectrometry provided age estimates between 70 and 11 ka. Crown dimensions place the tooth specimen at the upper limits of Late Pleistocene human ranges of variation. Wezmeh 1 metameric position (most likely a P3) remains uncertain and only its surficial morphology has been described so far. Accordingly, we used microfocus X-ray tomography (12.5 µm isotropic voxel size) to reassess the metameric position and taxonomic attribution of this specimen. We investigated its endostructural features and quantified crown tissue proportions. Topographic maps of enamel thickness (ET) distribution were also generated, and semilandmark-based geometric morphometric analyses of the enamel-dentine junction (EDJ) were performed. We compared Wezmeh 1 with unworn/slightly-moderately worn P3 and P4 of European Neanderthals, Middle Paleolithic modern humans from Qafzeh, an Upper Paleolithic premolar, and Holocene humans. The results confirm that Wezmeh 1 represents a P3. Based on its internal conformation and especially EDJ shape, Wezmeh 1 aligns closely with Neanderthals and is distinct from the fossil and extant modern human pattern of our comparative samples. Wezmeh 1 is thus the first direct evidence of Neanderthal presence on the western margin of the Iranian Plateau.