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The early Middle Pleistocene human material from Boxgrove (West Sussex, UK) consists of a partial left tibia and two lower incisors from a separate adult individual. These remains derive from deposits assigned to the MIS 13 interglacial at about 480 ka and have been referred to as Homo cf. heidelbergensis. The much larger skeletal sample from the Sima de los Huesos (Atapuerca, Spain) is dated to the succeeding MIS 12, at about 430 ka. This fossil material has previously been assigned to Homo heidelbergensis but is now placed within the Neanderthal clade. Because of the scarcity of human remains from the Middle Pleistocene and their morphological variability, this study assessed whether the Boxgrove specimens fit within the morphological variability of the homogeneous Sima de los Huesos population. Based on morphometric analyses performed against 22 lower incisors from Sima de los Huesos and published material, the data from the Boxgrove incisors place them comfortably within the range of Sima de los Huesos. Both assemblages present robust incisors distinct from the overall small recent Homo sapiens incisors, and Boxgrove also aligns closely with Homo neanderthalensis and some other European Middle Pleistocene hominins. Following morphological and cross-sectional analyses of the Boxgrove tibia compared to seven adult Sima de los Huesos specimens and a set of comparative tibiae, Boxgrove is shown to be similar to Sima de los Huesos and Neanderthals in having thick cortices and bone walls, but in contrast resembles modern humans in having a straight anterior tibial crest and a suggestion of a lateral concavity. Based on the patterns observed, there is no justification for assigning the Boxgrove and Sima de los Huesos incisors to distinct paleodemes, but the tibial data show greater contrasts and suggest that all three of these samples are unlikely to represent the same paleodeme., Competing Interests: Declaration of competing interest All authors have no conflict of interest to disclose., (Copyright © 2022 The Authors. Published by Elsevier Ltd.. All rights reserved.)

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., Competing Interests: Declaration of competing interest The authors declare no conflict of interest., (Copyright © 2022 Elsevier Ltd. All rights reserved.)

The two- and three-dimensional assessment of dental tissues has become routine in human taxonomic studies throughout the years. Nonetheless, most of our knowledge of the variability of the enamel and dentine dimensions of the human evolutionary lineage comes from the study of permanent dentition, and particularly from molars. This leads to a biased view of the variability of these features. Due to their early formation and rapid development, the deciduous teeth allow more simplified inferences regarding the processes involved in the dental tissue development of each group. Therefore, their study could be very valuable in dental palaeohistology. In this research, we have explored the dental tissue proportions of the deciduous canines belonging to some human samples of the Early and Middle Pleistocene. The purpose of this was to discuss the meaning of the similarities and differences observed in their histological pattern, as well as to evaluate the degree of covariance with that observed in the permanent dentition of these populations. Our results show that, although there are some similarities in the dental tissue proportions between the deciduous and permanent canines of the study samples, the two dental classes do not provide a similar or comparable pictures of the dental tissue pattern present in the dentition of fossil hominins. Future works on the dental tissue patterns of the anterior and posterior dentition, including deciduous teeth, of fossil samples, may help to shed light on this hypothesis., (© 2021 Anatomical Society.)

The Chinese Middle Pleistocene fossils from Hexian, Xichuan, Yiyuan, and Zhoukoudian have been generally classified as Homo erectus s.s. These hominins share some primitive features with other Homo specimens, but they also display unique cranial and dental traits. Thus, the Chinese Middle Pleistocene hominins share with other European and Asian hominin populations the so-called 'Eurasian dental pattern'. The late Early Pleistocene hominins from Gran Dolina-TD6.2 (Spain), representing the species Homo antecessor, also exhibit the Eurasian dental pattern, which may suggest common roots. To assess phylogenetic affinities of these two taxa, we evaluated and compared nonmetric and metric dental features and interpreted morphological differences within a comparative hominin framework. We determined that the robust roots of the molars, the shelf-like protostylid, the dendrite-like pattern of the enamel-dentine junction surface of the upper fourth premolars and molars, the strongly folded dentine of the labial surface of the upper incisors, and the rare occurrence of a mid-trigonid crest in the lower molars, are all characteristic of Chinese H. erectus. With regard to H. antecessor, we observed the consistent expression of a continuous mid-trigonid crest, the absence of a cingulum in the upper canines, a complex root pattern of the lower premolars, and a rhomboidal occlusal contour and occlusal polygon and protrusion in the external outline of a large a bulging hypocone in the first and second upper molars. Using two-dimensional geometric morphometrics, we further demonstrated that H. antecessor falls outside the range of variation of Chinese H. erectus for occlusal crown outline shape, the orientation of occlusal grooves, and relative locations of anterior and posterior foveae in the P 4 s, P 3 s, M 1 s, M 2 s, and M 2 s. Given their geographic and temporal separation, the differences between these two species suggest their divergence occurred at some point in the Early Pleistocene, and thereafter they followed different evolutionary paths., Competing Interests: Declaration of competing interest The authors declare that there is no conflict of interest., (Copyright © 2021 Elsevier Ltd. All rights reserved.)

The Inhibitory Cascade Model was proposed by Kavanagh and colleagues (Nature, 449, 427-433 [2007]) after their experimental studies on the dental development of murine rodent species. These authors described an activator-inhibitor mechanism that has been employed to predict evolutionary size patterns of mammalian teeth, including hominins. In the present study, we measured the crown area of the three lower permanent molars (M1, M2, and M3) of a large recent modern human sample of male and female individuals from a collection preserved at the Institute of Anthropology of the University of Coimbra (Portugal). The main aim of the present study is to test if the size molar patterns observed in this human sample fits the Inhibitory Cascade Model. For this purpose, we first measured the crown area in those individuals preserving the complete molar series. Measurements were taken in photographs, using a planimeter and following well-tested techniques used in previous works. We then plot the M 3 /M 1 and M 2 /M 1  size ratios. Our results show that the premise of the Inhibitory Cascade Model, according to which the average of the crown area of M2 is approximately one-third of the sum of the crown area of the three molars, is fulfilled. However, our results also show that the individual values of a significant number of males and females are out of the 95% confidence interval predicted by the Inhibitory Cascade Model in rodents. As a result, the present analyses suggest that neither the sample of males, nor that of females, nor the pooled sample fits the Inhibitory Cascade Model. It is important to notice that, although this model has been successfully tested in a large number of current human populations, to the best of our knowledge this is the first study in which individual data have been obtained in a recent human population rather than using the average of the sample. Our results evince that, at the individual level, some factors not yet known could interfere with this model masking the modulation of the size on the molar series in modern humans. We suggest that the considerable delay in the onset of M3 formation in modern humans could be related to a weakening of the possible activation/inhibition process for this tooth. Finally, and in support of our conclusions, we have checked that the absolute and relative size of M1 and M2 is not related to the M3 agenesis in our sample. In line with other studies in primates, our results do not support the Inhibitory Cascade Model in a recent human sample. Further research is needed to better understand the genetic basis of this mechanism and its relationship to the phenotype. In this way, we may be able to find out which evolutionary changes may be responsible for the deviations observed in many species, including Homo sapiens., (© 2021 Anatomical Society.)

The origin and evolution of hominin mortuary practices are topics of intense interest and debate 1-3 . Human burials dated to the Middle Stone Age (MSA) are exceedingly rare in Africa and unknown in East Africa 1-6 . Here we describe the partial skeleton of a roughly 2.5- to 3.0-year-old child dating to 78.3 ± 4.1 thousand years ago, which was recovered in the MSA layers of Panga ya Saidi (PYS), a cave site in the tropical upland coast of Kenya 7,8 . Recent excavations have revealed a pit feature containing a child in a flexed position. Geochemical, granulometric and micromorphological analyses of the burial pit content and encasing archaeological layers indicate that the pit was deliberately excavated. Taphonomical evidence, such as the strict articulation or good anatomical association of the skeletal elements and histological evidence of putrefaction, support the in-place decomposition of the fresh body. The presence of little or no displacement of the unstable joints during decomposition points to an interment in a filled space (grave earth), making the PYS finding the oldest known human burial in Africa. The morphological assessment of the partial skeleton is consistent with its assignment to Homo sapiens, although the preservation of some primitive features in the dentition supports increasing evidence for non-gradual assembly of modern traits during the emergence of our species. The PYS burial sheds light on how MSA populations interacted with the dead.

The Middle Pleistocene Sima de los Huesos (SH) site has yielded more than 7.500 human fossil remains belonging to a minimum of 29 individuals. Most of these individuals preserve either the complete mandibular molar series or at least the first (M 1 ) and second (M 2 ) molars. The inhibitory cascade mathematical model was proposed by Kavanagh et al. (Nature, 449, 427-433 [2007]) after their experimental studies on the dental development of murine rodent species. The activator-inhibitor mechanism of this model has shown its ability for predicting evolutionary size patterns of mammalian teeth, including hominins. The main aim of this study is to test whether the size molar patterns observed in the SH hominins fit the inhibitory cascade model. With this purpose, we have measured the crown area of all SH molars in photographs, using a planimeter and following techniques used and well contrasted in previous works. Following one of the premises of the inhibitory cascade model, we expect that the central tooth (M 2 in our case) of a triplet would have the average size of the two outer teeth. The absolute difference between the observed and the expected values for the M 2 s ranges from 0.23 to 8.46 mm 2 in the SH sample. In terms of percentage, the difference ranges between 0.25% and 10.34%, although in most cases, it is below 5%. The plot of the estimated M 3 /M 1 and M 2 /M 1 size ratios obtained in the SH hominins occupies a small area of the theoretical developmental morphospace obtained for rodent species. In addition, the majority of the values are placed near the theoretical line which defines the relationship predicted by the inhibitory cascade model in these mammals. The values of the slope and intercept of the reduced major regression obtained for the SH individuals do not differ significantly from those obtained for rodent species, thus confirming that the size of the molars of the SH hominins fits the inhibitory cascade model. We discuss these results in terms of dental development. Despite the promising results in the SH sample, we draw the attention to the fact that most Early Pleistocene Homo specimens exhibit a pattern (M 1  < M 2  > M 3 ), which is outside the expected theoretical morphospace predicted by the inhibitory cascade model. The shift from the M1 < M2 < M3 size relationship observed in early hominins (including H. habilis) to the M1 > M2 > M3 size relationship, which is predominant in modern humans, includes sequences that depart from predictions of the inhibitory cascade model. Additional studies are required to understand these deviations., (© 2020 Anatomical Society.)

Competing Interests: Conflict of interest The authors declare no conflict of interest.

Here, we present a metric and morphological study of the molar remains from the Montmaurin-La Niche mandible by means of microcomputed tomography. According to the last analysis, based on the combination of geomorphological and paleontological data, the level bearing this human mandible probably corresponds to the marine isotope stages (MIS) 7. These data place the Montmaurin-La Niche in a chronologically intermediate position between the Neanderthals and the Middle Pleistocene fossils (e.g., Sima de los Huesos, la Caune de l'Arago). A recent study has revealed that while the mandible is more closely related to the Early and Middle Pleistocene African and Eurasian populations, the morphology of the outer enamel surfaces of its molars is typical of the Neanderthal linage. The data presented here are in line with this finding because the morphology of the enamel-dentine junction of the molars is similar to that of Neanderthals, whereas the absolute and relative enamel thickness values (2D and 3D) are closer to those exhibited by some Early Pleistocene hominins. Moreover, the pulp cavity morphology and proportions are in concordance with the Neanderthal populations. Our results strengthen the hypothesis that the settlement of Europe could be the result of several migrations, at different times, originated from a common source population. Thus, the variability in the European Middle Pleistocene populations (e.g., Montmaurin, Sima de los Huesos, Arago, Mala Balanica) could indicate different migrations at different times and/or population fragmentation, without excluding the possible hybridization between residents and new settlers., Competing Interests: Conflict of interest The authors declare that there is no conflict of interest., (Copyright © 2020 Elsevier Ltd. All rights reserved.)

Sexual dimorphism is an important component of the total variation seen in populations and plays a key role in taxonomic debates. In this study, microtomographic (microcomputed tomography) techniques were applied to a sample of hominin teeth from the Sima de los Huesos site (Spain). Dental tissue proportions of the permanent canines were assessed to characterize the pattern and degree of sexual dimorphism within this population. In addition, the possible similarities and differences with the Homo neanderthalensis remains from Krapina (Croatia) and with a recent modern human sample were evaluated. A combination of classical statistical approaches with more novel techniques allowed us not only to ratify the sex allocation of the individuals previously assigned in the literature but also to estimate the sex of the youngest individuals, which were not assessed in previous studies. Likewise, the sexes of certain extensively worn canines and isolated pieces were estimated. As a result, the sex ratio observed in our dental sample from the Sima de los Huesos population is 5:9 (N m :N f ). In general terms, both Sima de los Huesos and Krapina dental samples have a degree of sexual dimorphism in their permanent canine tissue proportions that does not surpass that of modern humans. The marked dimorphic root volume of Sima de los Huesos mandibular canines is the exception, which surpasses the modern human mean, although it falls within the 95% confidence interval. Therefore, our results do not support that dental tissue proportions of the European Middle Pleistocene populations were more dimorphic than in modern humans. However, the differences in canine tissue proportions are great enough to allow sex estimation with a high degree of confidence., Competing Interests: Conflict of interest We declare no conflict of interest., (Copyright © 2020 Elsevier Ltd. All rights reserved.)

Dental enamel thickness, topography, growth and development vary among hominins. In Homo, the thickness of dental enamel in most Pleistocene hominins display variations from thick to hyper-thick, while Neanderthals exhibit proportionally thinner enamel. The origin of the thin trait remains unclear. In this context, the Middle Pleistocene human dental assemblage from Atapuerca-Sima de los Huesos (SH) provides a unique opportunity to trace the evolution of enamel thickness in European hominins. In this study, we aim to test the hypothesis if the SH molar sample approximates the Neanderthal condition for enamel thickness and/or distribution. This study includes 626 molars, both original and comparative data. We analysed the molar inner structural organization of the original collections (n = 124), belonging to SH(n = 72) and modern humans from Spanish origin (n = 52). We compared the SH estimates to those of extinct and extant populations of the genus Homo from African, Asian and European origin (estimates extracted from literature n = 502). The comparative sample included maxillary and mandibular molars belonging to H. erectus, East and North African Homo, European Middle Pleistocene Homo, Neanderthals, and fossil and extant H. sapiens. We used high-resolution images to investigate the endostructural configuration of SH molars (tissue proportions, enamel thickness and distribution). The SH molars exhibit on average thick absolute and relative enamel in 2D and 3D estimates, both in the complete crown and the lateral enamel. This primitive condition is shared with the majority of extinct and extant hominin sample, except for Neanderthals and some isolated specimens. On the contrary, the SH molar enamel distribution maps reveal a distribution pattern similar to the Neanderthal signal (with thicker enamel on the lingual cusps and more peripherally distributed), compared to H. antecessor and modern humans. Due to the phylogenetic position of the SH population, the thick condition in molars could represent the persistence of the plesiomorphic condition in this group. Still, more data is needed on other Early and Middle Pleistocene populations to fully understand the evolutionary meaning of this trait., Competing Interests: The authors have declared that no competing interests exist.

Characterizing dental development in fossil hominins is important for distinguishing between them and for establishing where and when the slow overall growth and development of modern humans appeared. Dental development of australopiths and early Homo was faster than modern humans. The Atapuerca fossils (Spain) fill a barely known gap in human evolution, spanning ~1.2 to ~0.4 million years (Ma), during which H. sapiens and Neandertal dental growth characteristics may have developed. We report here perikymata counts, perikymata distributions and periodicities of all teeth belonging to the TE9 level of Sima del Elefante, level TD6.2 of Gran Dolina (H. antecessor) and Sima de los Huesos. We found some components of dental growth in the Atapuerca fossils resembled more recent H. sapiens. Mosaic evolution of perikymata counts and distribution generate three distinct clusters: H. antecessor, Sima de los Huesos and H. sapiens.

Here we present the descriptive and comparative study of two immature scapulae recovered from the TD6.2 level of the Gran Dolina cave site (Sierra de Atapuerca, Spain) and assigned to Homo antecessor. This is the first time that data on the morphology and dimensions of the scapulae of a European late Early Pleistocene hominin population are provided. Considering the state of development and the linear dimensions, the scapula ATD6-116 could belong to a child of about 2-4 years. The morphology of ATD6-116 clearly departs from that of the Australopithecus afarensis juvenile specimen DIK-1-1, pointing to functional differences in locomotor behavior between Australopithecus and the late Early Pleistocene hominins. The immature scapula ATD6-118 belonged to an immature individual with a development of the scapula equivalent to that of adolescents of recent human populations. The scapulae ATD6-118 and KNM-WT 15000 present a similar state of development. Although the scapula KNM-WT 15000 is clearly larger than ATD6-118, these two specimens share some characteristics such as their relative narrowness and the value of the axilloglenoid and spinoglenoid angles. The glenoid fossa of ATD6-116 show a lateral orientation, whereas in ATD6-118 the glenoid fossa is slightly cranially oriented, but still within the range of variation of modern humans. The glenoid index of both ATD6-116 and ATD6-118 is low in accordance to the values usually observed in other early hominins, thus showing the primitive condition for this feature. Both scapulae show a ventrally placed axillary sulcus. The presence of this primitive feature in ATD-116 confirms that the shape of the axillary border has a genetic basis and it is not related to physical activity., (Published by Elsevier Ltd.)

Here we analyze the unpublished hominin dental remains recovered from the late Early Pleistocene Gran Dolina-TD6.2 level of the Sierra de Atapuerca (northern Spain), as well as provide a reassessment of the whole TD6.2 hominin dental sample. Comparative descriptions of the outer enamel surface (OES) and the enamel-dentine junction (EDJ) are provided. Overall, the data presented here support the taxonomic validity of Homo antecessor, since this species presents a unique mosaic of traits. Homo antecessor displays several primitive features for the genus Homo as well as some traits exclusively shared with Early and Middle Pleistocene Eurasian hominins. Some of these Eurasian traits were retained by the Middle Pleistocene hominins of Europe, and subsequently became the typical condition of the Neanderthal lineage. Although other skeletal parts present resemblances with Homo sapiens, TD6.2 teeth do not show any synapomorphy with modern humans. In addition, TD6.2 teeth can be well differentiated from those of Asian Homo erectus. The dental evidence is compatible with previous hypothesis about H. antecessor belonging to the basal population from which H. sapiens, Homo neanderthalensis, and Denisovans emerged. Future findings and additional research may help to elucidate the precise phylogenetic link among them., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

Several human dental traits typical of modern humans appear to be associated with the prolonged period of development that is a key human attribute. Understanding when, and in which early hominins, these dental traits first appeared is thus of strong interest. Using x-ray multiresolution synchrotron phase-contrast microtomography, we quantify dental growth and development in an archaic Homo juvenile from the Xujiayao site in northern China dating to 161,000-224,000 years or 104,000-125,000 years before present. Despite the archaic morphology of Xujiayao hominins, most aspects of dental development of this juvenile fall within modern human ranges (e.g., prolonged crown formation time and delayed first molar eruption). For its estimated age-at-death (6.5 years), its state of dental development is comparable to that of equivalently aged modern children. These findings suggest that several facets of modern human dental growth and development evolved in East Asia before the appearance of fully modern human morphology.

Tooth crown tissue proportions and enamel thickness distribution are considered reliable characters for inferring taxonomic identity, phylogenetic relationships, dietary and behavioural adaptations in fossil and extant hominids. While most Pleistocene hominins display variations from thick to hyper-thick enamel, Neanderthals exhibit relatively thinner. However, the chronological and geographical origin for the appearance of this typical Neanderthal condition is still unknown. The European late Early Pleistocene species Homo antecessor (Gran Dolina-TD6 site, Sierra de Atapuerca) represents an opportunity to investigate the appearance of the thin condition in the fossil record. In this study, we aim to test the hypothesis if H. antecessor molars approximates the Neanderthal condition for tissue proportions and enamel thickness. To do so, for the first time we characterised the molar inner structural organization in this Early Pleistocene hominin taxon (n = 17) and compared it to extinct and extant populations of the genus Homo from African, Asian and European origin (n = 355). The comparative sample includes maxillary and mandibular molars belonging to H. erectus, East and North African Homo, European Middle Pleistocene Homo, Neanderthals, and fossil and extant H. sapiens. We used high-resolution images to investigate the endostructural configuration of TD6 molars (tissue proportions, enamel thickness and distribution). TD6 permanent molars tend to exhibit on average thick absolute and relative enamel in 2D and 3D estimates, both in the complete crown and the lateral enamel. This condition is shared with the majority of extinct and extant hominin sample, except for Neanderthals and some isolated specimens. However, while the total crown percentage of dentine in TD6 globally resembles the low modern values, the lateral crown percentage of dentine tends to be much higher, closer to the Neanderthal signal. Similarly, the H. antecessor molar enamel distribution maps reveal a relative distribution pattern that is more similar to the Neanderthal condition (with the thickest enamel more spread at the periphery of the occlusal basin) rather than that of other fossil specimens and modern humans (with thicker cuspal enamel). Future studies on European Middle Pleistocene populations will provide more insights into the evolutionary trajectory of the typical Neanderthal dental structural organization., Competing Interests: The authors have declared that no competing interests exist.

The penecontemporaneous Middle Pleistocene sites of Fontana Ranuccio (Latium) and Visogliano (Friuli-Venezia Giulia), set c. 450 km apart in central and northeastern Italy, respectively, have yielded some among the oldest human fossil remains testifying to a peopling phase of the Italian Peninsula broadly during the glacial MIS 12, a stage associated with one among the harshest climatic conditions in the Northern hemisphere during the entire Quaternary period. Together with the large samples from Atapuerca Sima de los Huesos, Spain, and Caune de l'Arago at Tautavel, France, the remains from Fontana Ranuccio and Visogliano are among the few mid-Middle Pleistocene dental assemblages from Western Europe available for investigating the presence of an early Neanderthal signature in their inner structure. We applied two- three-dimensional techniques of virtual imaging and geometric morphometrics to the high-resolution X-ray microtomography record of the dental remains from these two Italian sites and compared the results to the evidence from a selected number of Pleistocene and extant human specimens/samples from Europe and North Africa. Depending on their preservation quality and on the degree of occlusal wear, we comparatively assessed: (i) the crown enamel and radicular dentine thickness topographic variation of a uniquely represented lower incisor; (ii) the lateral crown tissue proportions of premolars and molars; (iii) the enamel-dentine junction, and (iv) the pulp cavity morphology of all available specimens. Our analyses reveal in both samples a Neanderthal-like inner structural signal, for some aspects also resembling the condition shown by the contemporary assemblage from Atapuerca SH, and clearly distinct from the recent human figures. This study provides additional evidence indicating that an overall Neanderthal morphological dental template was preconfigured in Western Europe at least 430 to 450 ka ago., Competing Interests: The authors have declared that no competing interests exist.

The description of a new skull (D4500) from the Dmanisi site (Republic of Georgia) has reopened the debate about the morphological variability within the genus Homo. The new skull fits with a mandible (D2600) often referred as 'big' or 'enigmatic' because of its differences with the other Dmanisi mandibles (D211 and D2735). In this report we present a comparative study of the variability of the Dmanisi mandibles under a different perspective, as we focus in morphological aspects related to growth and development. We have followed the notion of modularity and phenotypic integration in order to understand the architectural differences observed within the sample. Our study reveals remarkable shape differences between D2600 and the other two mandibles, that are established early in the ontogeny (during childhood or even before) and that do not depend on size or sexual dimorphism. In addition, D2600 exhibits a mosaic of primitive and derived features regarding the Homo clade, which is absent in D211 and D2735. This mosaic expression is related to the location of the features and can be explained under the concept of modularity. Our study would support the possibility of two different paleodemes represented at the Dmanisi site. This hypothesis has been previously rejected on the basis that all the individuals were constrained in the same stratigraphic and taphonomic settings. However, our revision of the complex Dmanisi stratigraphy suggests that the accumulation could cover an undetermined period of time. Even if "short" in geological terms, the hominin accumulation was not necessarily synchronic. In the same line we discard that the differences between D2600 and the small mandibles are consequence of wear-related dentoalveolar remodeling. In addition, dental wear pattern of D2600 could suggest an adaptation to a different ecological niche than the other Dmanisi individuals.

Here we present a detailed palaeopathological study of the hominin mandible ATE9-1 found at the Sima del Elefante site (TE), Sierra de Atapuerca, Spain. This fossil represents the earliest hominin remains from Western Europe with an age of ca. 1.3 Ma. The specimen displays several dento-gnathic lesions; the antiquity and geographic location of this fossil justifies a detailed palaeopathological study to determine if the pathologies have significantly altered taxonomically relevant features. Our study reveals severe dental attrition combined with generalized hypercementosis, alveolar root exposure, mild periodontal disease, tooth dislocation, and an anomalous occlusal plane. We have also observed calculus deposits, two cystic lesions and an anomalous wear facet compatible with tooth picking. The majority of these pathological signs can be explained by compensatory eruption. We propose that these lesions are associated as causes, consequences, and amplifiers of one another within the framework of heavy and even traumatic occlusion, masticatory habits, or both traumatic occlusion and masticatory habits. Despite the severity of these lesions, occlusion was at least partially functional so it was unlikely to influence the survival of this individual. In addition, the lesions do not prohibit the taxonomic assessment of the mandible., (Copyright © 2011 Elsevier Ltd. All rights reserved.)

We present a detailed morphological comparative study of the hominin mandible ATE9-1 recovered in 2007 from the Sima del Elefante cave site in Sierra de Atapuerca, Burgos, northern Spain. Paleomagnetic analyses, biostratigraphical studies, and quantitative data obtained through nuclide cosmogenic methods, place this specimen in the Early Pleistocene (1.2-1.3 Ma). This finding, together with archaeological evidence from different European sites, suggests that Western Europe was colonised shortly after the first hominin expansion out of Africa around the Olduvai subchron. Our analysis of the ATE9-1 mandible includes a geometric morphometric analysis of the lower second premolar (LP(4)), a combined and detailed external and internal assessment of ATE9-1 roots through CT and microCT techniques, as well as a comparative study of mandibular and other dental features. This analysis reveals some primitive Homo traits on the external aspect of the symphysis and the dentition shared with early African Homo and the Dmanisi hominins. In contrast, other mandibular traits on the internal aspect of the symphysis are derived with regard to African early Homo, indicating unexpectedly large departures from patterns observed in Africa. Reaching the most occidental part of the Eurasian continent implies that the first African emigrants had to cross narrow corridors and to overcome geographic barriers favouring genetic drift, long isolation periods, and adaptation to new climatic and seasonal conditions. Given these conditions and that we are dealing with a long time period, it is possible that one or more speciation events could have occurred in this extreme part of Eurasia during the Early Pleistocene, originating in the lineages represented by the Sima del Elefante-TE9 hominins and possibly by the Gran Dolina-TD6 hominins. In the absence of any additional evidence, we prefer not include the specimen ATE9-1 in any named taxon and refer to it as Homo sp., (Copyright © 2011 Elsevier Ltd. All rights reserved.)