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The phylogenetic relationships of the small-bodied catarrhine Pliobates cataloniae (∼11.6 Ma, NE Iberian Peninsula) have been controversial since its original description. However, the recent report of additional dentognathic remains has supported its crouzeliid pliopithecoid status. Based on the available hypodigm, the molar enamel-dentine junction (EDJ) shape of P. cataloniae is compared with that of other pliopithecoids from the same basin as well as both extinct and extant hominoids to further evaluate its pliopithecoid affinities. We also quantitatively compare the EDJ shape among these taxa by means of landmark-based three-dimensional geometric morphometrics using principal component analysis (PCA), canonical variate analysis, and between-group PCA. Permutation tests are performed to test whether Pliobates variation exceeds that of extant hominoid genera. Results indicate that Pliobates is similar in molar EDJ shape to other pliopithecoids, particularly crouzeliids. The variation displayed by Pliobates upper molars is less marked at the EDJ level than at the outer enamel surface, probably owing to differential enamel wear and intraspecific differences in enamel thickness. Multivariate analyses of EDJ shape show that all pliopithecoids (including Pliobates) cluster together in the PCAs, canonical variate analyses, and between-group PCAs and occupy a different portion of the morphospaces from extinct and extant hominoids. Posterior and typicality probabilities strongly support the classification of Pliobates as a pliopithecoid, wheras permutation tests fail to reject the single-genus hypothesis for the P. cataloniae hypodigm. We conclude that P. cataloniae is a crouzeliid pliopithecoid, as recently supported by cladistic analyses of craniodental characters, and that previous cladistic results that supported a stem hominoid status are attributable to postcranial convergences with crown hominoids. Our results further highlight the potential of three-dimensional geometric morphometrics analyses of the EDJ shape for better informing fossil primate alpha-taxonomy by means of quantitatively testing hypotheses about tooth shape variation., Competing Interests: Declaration of competing interest There is no conflict of interest., (Copyright © 2024 Elsevier Ltd. All rights reserved.)

Although the first discovery of a human fossil in the Sima de los Huesos took place in 1976, systematic excavations did not begin there until 1984. Since then, this site has been continuously excavated in month-long camps. The site is dated by different radiometric techniques to between 430,000 and 300,000 years ago. Until the 2023 campaign, just over 7000 human fossils have been recovered, constituting the largest collection of fossils prior to Homo sapiens ever discovered. The fossils correspond to a minimum of 29 individuals of both sexes and different ages at death, from preadolescents to a specimen of advanced age. Comparative anatomy and ancient DNA studies both suggest that this is a population closely related to Homo neanderthalensis. The great variety and extraordinary quality of the fossils recovered have allowed us to carry out a series of investigations that have greatly increased our knowledge about the evolution of Homo in the Middle Pleistocene. Among the most important discoveries, it has been possible to establish body size and proportions, the confirmation that the origin of the accumulation of human fossils was of an anthropic nature, that those past humans took care of disabled individuals and who were capable of having an oral language almost as complex and efficient as that of our own species., (© 2024 The Author(s). The Anatomical Record published by Wiley Periodicals LLC on behalf of American Association for Anatomy.)

The Sima de los Huesos site in the Sierra de Atapuerca (Burgos, northern Spain) is a Middle Pleistocene locality with the most important accumulation of humans in the European record for this age. In addition to the hominin collection, the Sima has provided numerous faunal remains. Here, we update the carnivoran mammal fauna from the Sima de los Huesos. The analyzed carnivoran fauna is taxonomically diverse consisting of 11 species from 9 genera and four families. The high taxonomic diversity of carnivorans at the site is interpreted as corresponding to high overall ecosystem productivity. A synthetic description of the carnivoran remains is presented adding new information on the biochronology of the carnivoran species assemblage and its paleoecological association with European Middle Pleistocene hominins. Because of their similarity in taphonomic setting and diets-that the carnivoran fauna and humans were found in the same place and had similar protein-rich diets-clarifying the paleontological and paleoecological context of these carnivorans permits a better understanding of the humans also found at this site., (© 2023 The Authors. The Anatomical Record published by Wiley Periodicals LLC on behalf of American Association for Anatomy.)

Previous studies on the morphology of the inner ear (semicircular canals and cochlea) in the Sima de los Huesos hominin sample have provided important results on the evolution of these structures in the Neandertal lineage. Similarly, studies of the anatomy of the external and middle ear cavities of the Sima de los Huesos hominins have also provided important data on the auditory capacities of this European Middle Pleistocene population. The present contribution provides unpublished data on three new middle ear variables from the Sima de los Huesos fossils and compares these data with values from samples of Pan troglodytes, Homo neanderthalensis and Homo sapiens. The results of this analysis are combined with those obtained in previous studies to characterize the anatomy of the outer, middle and inner ear in the Sima de los Huesos fossils, as well as to establish the order of appearance of the features that characterize Neandertal ears. As in other cranial structures, the ear region in the Sima de los Huesos show a mosaic evolutionary pattern that includes primitive traits, others shared exclusively with Neandertals, and others that are specific to the Sima de los Huesos hominins. Neandertals and Sima de los Huesos hominins share two exclusive features of the middle ear that are among the first characteristics of the Neandertal lineage: a long tympanic cavity and a large entrance and exit of the mastoid antrum. Along with these traits, the Sima de los Huesos hominins present two specialized features: large volumes of the tympanic cavity and the mastoid antrum. Finally, the middle ear of the Neandertals is characterized by the presence of small angles between the tympanic axis and the plane of the oval window., (© 2023 The Authors. The Anatomical Record published by Wiley Periodicals LLC on behalf of American Association for Anatomy.)

The Middle Pleistocene site of the Sima de los Huesos (Sierra de Atapuerca, northern Spain) has yielded a considerable number of human fossils during the period 1984-2020. Among them, up to 253 maxillary teeth have been recovered. In this article, we present the description of the eight dental classes of the maxilla following the Arizona State University Dental Anthropology System classification. In addition, we present the mean mesiodistal and buccolingual diameters of these teeth compared to those of Neanderthals and a modern human sample. The morphology of both the anterior and posterior teeth suggests a close relationship of the Sima de los Huesos hominins with the populations of the second half of the Middle Pleistocene of Europe and the Near East, as well as with the so-called classic Neanderthals of Europe. Features with a recognizable taxonomic signal allow grouping the Sima de los Huesos hominins with different paleodemes into a Neanderthal clade. The dental evidence of the Sima de los Huesos hominins is key to suggest a complex model for the settlement of Europe during the Middle Pleistocene. During this period, different migrations of human groups probably coming from Southwest Asia, replacements, prolonged isolations, as well as hybridization and introgression processes would have contributed to the diversity of hominins in Europe., (© 2021 American Association for Anatomy.)

Some of the Sima de los Huesos (SH) humeri have been previously studied and described elsewhere. Here we present an updated inventory and a review of the specimens recovered to the present day. The morphological key traits of the adult and subadult specimens are described, discussed, and illustrated. The SH humeri share with Neandertals many traits usually considered to be Neandertal specializations, thus, most of this morphological pattern is not exclusive to them. The variation found within fossil samples stresses the frequential nature of all these traits and in the specific case of the SH humeri, most of the traits considered as phylogenetically relevant are retained by their descendants, the Neandertals. Some traits are plesiomorphic for the entire genus Homo or are present in European hominins since the early Pleistocene. Finally, some other traits display high variability within the SH sample or different hominin samples and are of uncertain phylogenetic value. Altogether, this evidence is consistent with the hypothesis based on the overall cranial and postcranial morphology that the SH hominins are a sister group to the later Neandertals., (© 2023 The Authors. The Anatomical Record published by Wiley Periodicals LLC on behalf of American Association for Anatomy.)

The excellent fossil record from Sima de los Huesos (SH) includes three well-known complete adult femora and several partial specimens that have not yet been published in detail. This fossil record provides an opportunity to analyze the morphology of European pre-Neandertal adult femur and its variation with different evolution patterns. Currently, there are a minimum of five adult individuals (males or females). In this study, we compiled previously published basic anatomical and biometric characteristics of SH adult femora, emphasizing the most relevant features compared to other recent and fossil hominins. The SH femora exhibited a primitive morphological pattern common to all non-Homo sapiens femora, as well as most of the Neandertal traits. Therefore, the complete Upper Pleistocene Neandertal pattern was well-established in Middle Pleistocene ancestors long before the proper Neandertals appeared. Additionally, we highlight that the SH and Neandertal femora share some morphological traits and proportions with modern humans that hold sexual significance in our species, regardless of size. Keeping this in mind, we discussed the sex determination of the complete SH specimens and re-evaluated sex allocation in two of them., (© 2023 The Authors. The Anatomical Record published by Wiley Periodicals LLC on behalf of American Association for Anatomy.)

Here, we provide a complete, updated, and illustrated inventory, as well as a comprehensive study, of the tarsals (rearfoot) recovered from the Middle Pleistocene site of Sima de los Huesos (SH, Atapuerca, Spain) in comparison to other Homo comparative samples, both extant and fossil. The minimum number of individuals (MNI) estimated from the tarsals has been established as 15, which represents 51.7% of the 29 dental individuals identified within the SH sample. Within the SH hominin foot sample, an exclusive combination of primitive or plesiomorphic and derived or autapomorphic traits can be observed when compared with other Homo individuals/populations. Other characters are shared among SH hominins and Neandertals that might represent shared derived or autapomorphic traits for this evolutionary line, and most are likely related to robusticity (e.g., rectangular-like trochlea of the talus, broad calcanei, broad naviculars, and short lateral cuneiforms). Additionally, we observed some exclusive autapomorphic traits in the SH tarsal sample (e.g., narrow head of the talus and short intermediate cuneiforms). A few exclusive traits in SH tarsal remains are even more robust than in Neandertals (e.g., broad lateral malleolar facet in talus, more projected sustentaculum tali, and broad medial cuneiform). These traits could suggest a slightly higher level of gracilization in the tarsal bones of Neandertals compared to the SH sample that is also supported by other anatomical postcranial skeleton elements. Additionally, some paleobiological inferences are made in relation to body size (stature and body mass) and some associations are proposed within the SH sample. In conclusion, the morphology of the SH tarsi confirms an evolutionary relationship of sister groups between this population and Neandertals, probably representing a morphotype similar to the Neandertal ancestors., (© 2024 The Authors. The Anatomical Record published by Wiley Periodicals LLC on behalf of American Association for Anatomy.)

The Middle Pleistocene site of the Sima de los Huesos (Sierra de Atapuerca, northern Spain) has yielded a considerable number of human fossils during the period 1984-2020. Among them, up to 314 mandibular teeth have been identified. In this second paper dedicated to the dentition we present the description of the eight dental classes of the mandible following the Arizona State University Dental Anthropology System (ASUDAS) classification. In addition, we show the mean mesiodistal and buccolingual diameters obtained in these teeth compared to those of Neanderthals and a modern human sample. The morphology of both the anterior and posterior teeth suggests a close relationship of the Sima de los Huesos hominins with the populations of the second half of the Middle Pleistocene of Europe and the Near East, as well as with the so-called classic Neanderthals of Europe. The combination of dental traits in these populations is characteristic and diagnostic and suggests grouping the Sima de los Huesos hominins with the other paleodemes in a Neanderthal clade. The dental evidence of the Sima de los Huesos hominins is key to propose a complex model for the settlement of Europe during the Middle Pleistocene. In this period, different migrations of human groups probably coming from Southwest Asia, replacements, prolonged isolations, as well as hybridization and introgression processes would have contributed to the diversity of hominins in Europe., (© 2021 American Association for Anatomy.)

This study provides a complete, updated and illustrated inventory, as well as a comprehensive study, of the metatarsals and foot phalanges (forefoot) recovered from the Middle Pleistocene site of Sima de los Huesos (SH, Atapuerca, Spain) in comparison to other Homo comparative samples, both extant and fossils. This current updated review has established a minimum number of individuals (MNI) of 17, which represent 58.6% of the 29 dental individuals identified within the SH sample. An exclusive or autoapomorphic combination of traits can be recognized within the SH hominin foot sample. A few traits appear primitive or plesiomorphic when compared with earlier Homo individuals and other recent modern humans. There are other metrical and morphological traits that SH hominins and Neandertals have in common that sometimes represent shared derived traits in this evolutionary line, most of which are probably related to robusticity. Furthermore, some exclusive autoapomorphic traits are observed in the SH sample: a very broad first metatarsal, long and broad hallucal proximal foot phalanges and possibly extremely robust lateral distal foot phalanges compared to those of Neandertals and modern humans. In these last traits, the SH metatarsals and pedal phalanges are even more robust than in Neandertals. They are herein named as "hyper-Neandertal" traits, which could suggest a slight gracilization process in this evolutionary line, at least in the hallux toe. Finally, some paleobiological inferences are made in relation to body size (stature and body mass) and some associations are proposed within the SH sample., (© 2024 The Authors. The Anatomical Record published by Wiley Periodicals LLC on behalf of American Association for Anatomy.)

The postcranial skeleton of fossil hominins is crucial for reconstructing the processes that occurred between the time of death and the recovery of the bones. Thousands of postcranial skeletal fragments from at least 29 hominin individuals have been recovered from the Sima de los Huesos Middle Pleistocene site in Spain. This study's primary objective is to address the main taphonomic features of the postcranial remains from the Sima de los Huesos sample, including antemortem, perimortem, and postmortem skeletal disturbances. We present an updated assessment of the bone surface modification analysis, the fracture pattern analysis, and the skeletal part representation to facilitate interpretation of the biostratinomic and fossil-diagenetic processes in this large paleoanthropological collection. We conclude that carnivores (probably bears) had limited access to the hominin bones and complete bodies were probably placed in the site., (© 2023 American Association for Anatomy.)

The basicranium contains multiple synchondroses potentially informative for estimating the developmental stage of individuals. The basilar synchondrosis has been routinely used for this purpose in bioarchaeological, forensic and paleoanthropological research, and studies carried out in modern human populations have shown a close relationship between the fusion of the occipitomastoidal synchondrosis and developmental processes. This synchondrosis articulates the jugular process of the occipital bone with the jugular surface of the temporal bone. As the process of fusion of the synchondrosis progresses, the jugular surface undergoes a series of alterations whose study allows to establish the state of fusion of the synchondrosis when the individual died. The extraordinary preservation of the jugular surface in a large number of individuals represented in the fossil hominin sample from the middle Pleistocene site of the Sima de los Huesos (SH) has made it possible to carry out the first systematic study to assess the usefulness of occipitomastoidal synchondrosis in the establishment of the state of development in fossil hominins. Our results show that the complete closure of the occipitomastoidal synchondrosis occurred toward the end of the growth period in the SH fossils. This result opens up the possibility of using it to determine the developmental stage of fossil hominins for which no other information is available, such as the state of the dentition or the degree of closure of the basilar synchondrosis. This has allowed us to infer a state of development for three SH crania where it could not previously be established with certainty., (© 2023 The Authors. The Anatomical Record published by Wiley Periodicals LLC on behalf of American Association for Anatomy.)

Around 42,000 years ago, anatomically modern humans appeared in Western Europe to the detriment of indigenous Neanderthal groups. It is during this period that new techno-cultural complexes appear, such as the Châtelperronian that extends from northern Spain to the Paris Basin. The Grotte du Renne (Arcy-sur-Cure) is a key site for discussing the biological identity of its makers. This deposit has yielded several Neanderthal human remains in its Châtelperronian levels. However, the last inventory of the paleoanthropological collection attributed to this techno-complex allowed the identification of an ilium belonging to a neonate (AR-63) whose morphology required a thorough analysis to assess its taxonomic attribution. Using geometric morphometrics, we quantified its morphology and compared it to that of 2 Neanderthals and 32 recent individuals deceased during the perinatal period to explore their morphological variation. Our results indicate a morphological distinction between the ilia of Neanderthals and anatomically modern neonates. Although AR-63 is slightly outside recent variability, it clearly differs from the Neanderthals. We propose that this is due to its belonging to an early modern human lineage whose morphology differs slightly from present-day humans. We also explore different hypotheses about the presence of this anatomically modern neonate ilium among Neanderthal remains., (© 2023. Springer Nature Limited.)

Since the discovery of a human mandible in 1887 near the present-day city of Banyoles, northeastern Spain, researchers have generally emphasized its archaic features, including the lack of chin structures, and suggested affinities with the Neandertals or European Middle Pleistocene (Chibanian) specimens. Uranium-series and electron spin resonance dating suggest the mandible dates to the Late Pleistocene (Tarantian), approximately ca. 45-66 ka. In this study, we reassessed the taxonomic affinities of the Banyoles mandible by comparing it to samples of Middle Pleistocene fossils from Africa and Europe, Neandertals, Early and Upper Paleolithic modern humans, and recent modern humans. We evaluated the frequencies and expressions of morphological features and performed a three-dimensional geometric morphometric analysis on a virtual reconstruction of Banyoles to capture overall mandibular shape. Our results revealed no derived Neandertal morphological features in Banyoles. While a principal component analysis based on Euclidean distances from the first two principal components clearly grouped Banyoles with both fossil and recent Homo sapiens individuals, an analysis of the Procrustes residuals demonstrated that Banyoles did not fit into any of the comparative groups. The lack of Neandertal features in Banyoles is surprising considering its Late Pleistocene age. A consideration of the Middle Pleistocene fossil record in Europe and southwest Asia suggests that Banyoles is unlikely to represent a late-surviving Middle Pleistocene population. The lack of chin structures also complicates an assignment to H. sapiens, although early fossil H. sapiens do show somewhat variable development of the chin structures. Thus, Banyoles represents a non-Neandertal Late Pleistocene European individual and highlights the continuing signal of diversity in the hominin fossil record. The present situation makes Banyoles a prime candidate for ancient DNA or proteomic analyses, which may shed additional light on its taxonomic affinities., Competing Interests: Declaration of competing interest The authors declare to have no competing interests., (Copyright © 2022 Elsevier Ltd. All rights reserved.)

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.)

Circum-nasal and nasal cavity morphology add to the picture of the Sima de los Huesos specimens as, at one level, representing a distinct morph and, at another, displaying individual variation. They developed a robust, midline-grooved, three-dimensional spinal ridge lying anteriorly in the nasal cavity floor that was distended posteriorly over the nasal cavity floor, and, typically, an expansive, three-dimensional patch of rugose bone on the nasal cavity wall where a conchal crest would otherwise lie. They vary, for example, in degree of topographic relief of the nasal cavity wall, expression of the spinal ridge, and development of nasal crests and fossae. Lacking an anterior nasal spine, Sima specimens differ from extant and most fossil Homo sapiens, some specimens attributed to H. heidelbergensis, and the Gran Dolina partial face, whose anterior nasal spine is a superoanterior distention of the nasoalveolar clivus, and also from Neanderthals, whose anterior nasal spine projects anteriorly away from the nasoalveolar clivus. Comparison of Neanderthals, the Sima hominin, and specimens regarded as H. heidelbergensis calls for re-evaluating the integrity of "heidelbergensis" and rethinking the phylogenetic relationships of them all. To precisely describe the numerous features and combinations thereof of the nasal region in Sima specimens, and compare them with Neandertals and "H. heidelbergensis", we developed terminology that is applicable not only to hominins, but to mammals in general., (© 2021 American Association for Anatomy.)

After 34 years of research and findings in the Middle Pleistocene site of the Sima de los Huesos (SH) of the Sierra de Atapuerca (Burgos, Spain), we present an update of the estimation of the number of individuals (ENI) identified in the SH hominin assemblage. The last ENI, published in 2004, was 28. Although the number of specimens recovered has almost doubled since then and more complete information is now available, this new analysis suggests that the ENI is 29. Some individuals are still represented by only one or a few teeth and the upper and lower jaws of various individuals have not been completed. We suggest that the amateur cavers, who accessed the SH site for years looking for bear fossils, destroyed a significant number of hominin fossils. Despite this, we have a good picture of the group of hominins represented in the SH assemblage. Because complete corpses were accumulated in SH and it is a closed cavity, it is expected that a significant number of hominin fossils remains to be discovered. According to the previous estimates, the representation of females is greater than that of males. However, the observed sex ratio is not significantly different from the 1:1 ratio. With the exception of the possible presence of a child individual, most of the remaining 28 individuals are teenagers or young adults (24/28 = 0.857). The ages of death of SH hominins appear to conform to a catastrophic profile., (© 2020 American Association for Anatomy.)

Competing Interests: Conflict of interest There is no conflict of interests.

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.

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.)

The cochlea contains taxonomic and phylogenetic information and its morphology is related with hearing abilities among fossil hominins. Data for the genus Homo is presently limited to early Homo and the early Neandertals from Krapina. The present study of the middle Pleistocene hominins from the Sima de los Huesos (SH) provides new evidence on cochlear evolution in the genus Homo. We compared the absolute length, proportional lengths of each turn, number of turns, size and shape of the cross-section of the basal turn, volume, curvature gradient, and thickness of the cochlea between extant Pan troglodytes, extant Homo sapiens, Homo neanderthalensis and the SH hominins. The SH hominins resemble P. troglodytes in the proportionally long basal turn, the small size and round shape of the cross-section of the basal turn, the small cochlear volume and the low cochlear thickness. The SH hominins resemble Neandertals and H. sapiens in their long cochlear length and in the proportionally short third turn. Homo neanderthalensis and H. sapiens share several features, not present in the SH hominins, and that likely represent homoplasies: a larger volume, larger size and oval shape of the cross-section of the basal turn and higher cochlear thickness. Later Neandertals show a derived proportionally shorter apical turn. Changes in cochlear volume in Homo cannot be fully explained by variation in body mass or cochlear length but are more directly related to changes in the cross-sectional area of the basal turn. Based on previous studies of the outer and middle ear in SH hominins, changes in the outer and middle ear preceded changes in the inner ear, and the cochlea and semicircular canals seem to have evolved independently in the Neandertal clade. Finally, the small cochlear volume in the SH hominins suggests a slightly higher upper limit of hearing compared with modern humans., (Copyright © 2019 Elsevier Ltd. All rights reserved.)

Only a few postcranial remains have been assigned to the Miocene great ape Dryopithecus fontani, leading to uncertainties in the reconstruction of its overall body plan and positional behavior. Here we shed light on the locomotor repertoire of this species through the study of the femoral neck cortical bone (FNCB) distribution of IPS41724, a partial proximal femur from the Abocador de Can Mata locality ACM/C3-Az (11.9 Ma, middle Miocene; Vallès-Penedès Basin, Spain) attributed to this taxon. This specimen was scanned through computed tomography to measure the superior (SUP) and inferior (INF) cortical thicknesses at the middle and the base of the femoral neck. Measurements were compared with a sample of extant primates and the femur IPS18800.29 from the younger great ape Hispanopithecus laietanus from Can Llobateres 2 (9.6 Ma, late Miocene; Vallès-Penedès Basin), previously shown to display a homogeneous FNCB distribution at the midneck section coupled with postcranial adaptations to below-branch suspensory behaviors. Our analyses indicate an asymmetric FNCB distribution for IPS41724 (SUP/INF index = ∼0.4 at the midneck and base of the neck sections), comparable with that of quadrupedal primates and bipedal hominins (including early australopiths), but contrasting with the homogeneous FNCB distribution of Hispanopithecus and extant great apes. An asymmetrical FNCB distribution has been associated with stereotyped loads at the hip joint (as in both quadrupedal and bipedal taxa). Our results therefore support a significant quadrupedal component of the positional behavior of Dryopithecus, thus strengthening the argument that plesiomorphic generalized quadrupedalism was still a major locomotor behavior for Miocene great apes. If that were the case, it could have deep implications for the origins of hominin bipedalism., (Copyright © 2019 Elsevier Ltd. All rights reserved.)

Ancient evidence of human presence in Europe is recorded in several Early Pleistocene archaeopalaeontological sites from Spain, France and Italy. This is the case of Barranco León (BL) and Fuente Nueva-3 (FN-3), two localities placed near the town of Orce (depression of Baza and Guadix, SE Spain) and dated to ~1.4 Ma. At these sites, huge assemblages of Oldowan tools and evidence of defleshing, butchering and marrow processing of large mammal bones have been recovered together with a deciduous tooth of Homo sp. in the case of level BL-D. In this study, we: (i) describe in detail the anthropic marks found in the bone assemblages from these sites; (ii) analyse patterns of defleshment, butchery and marrow processing, based on the modifications identified in the cortical surface of the fossils; and (iii) discuss on the subsistence strategies of the first hominins that inhabited the European subcontinent during Early Pleistocene times.

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.)

In the Iberian Peninsula, Miocene apes (Hominoidea) are generally rare and mostly restricted to the Vallès-Penedès Basin. Here we report a new hominoid maxillary fragment with M 2 from this basin. It was surface-collected in March 2017 from the site of Can Pallars i Llobateres (CPL, Sant Quirze del Vallès), where fossil apes had not been previously recorded. The locality of provenance (CPL-M), which has delivered no further fossil remains, is located very close (ca. 50 m) to previously known CPL outcrops, and not very far (ca. 500 m in NW direction) from the classical hominoid-bearing locality of Can Poncic 1. Here we describe the new fossil and, based on the size and proportions of the M 2 , justify its taxonomic attribution to Hispanopithecus cf. laietanus, a species previously recorded from several Vallesian sites of the Vallès-Penedès Basin. Based on the associated mammalian fauna from CPL, we also provide a biochronological dating and a paleoenvironmental reconstruction for the site. The associated fauna enables an unambiguous correlation to the Cricetulodon hartenbergeri - Progonomys hispanicus interval local subzone, with an estimated age of 9.98-9.73 Ma (late Vallesian, MN10). Therefore, CPL-M is roughly coeval with the Hispanopithecus laietanus-bearing localities of Can Llobateres 1 and Can Feu 1, and minimally older than those of La Tarumba 1 and Can Llobateres 2. In contrast, CPL-M is younger than the early Vallesian (MN9) localities of Can Poncic 1 (the type locality of Hispanopithecus crusafonti) as well as Polinyà 2 (Gabarró) and Estació Depuradora d'Aigües Residuals-Riu Ripoll 13, where Hispanopithecus sp. is recorded. The associated fauna from CPL indicates a densely forested and humid paleoenvironment with nearby freshwater. This supports the view that Hispanopithecus might have been restricted to dense wetland forests soon before its extinction during the late Vallesian, due to progressive climatic deterioration. Coupled with the existence of other fossiliferous outcrops in the area, this find is most promising for the prospect of discovering additional fossil hominoid remains in the future., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

The sedimentary record in the Guadix-Baza Basin (southern Spain) has proved to be a great source of information for the Miocene through the Pleistocene periods, due to the abundant faunal remains preserved, in some cases associated with lithic tools. The Solana del Zamborino (SZ) section has been the subject of controversy ever since a magnetostratigraphic analysis resulted in an age of 750-770 Kyr for Acheulean tools, a chronology significantly older than the ~600 Kyr established chronology for the first Acheulean record in Europe. Although recent findings at the "Barranc de la Boella" site (north-east of the Iberian Peninsula) seem to indicate that an earlier introduction of such technique in Europe around 0.96-0.781 Ma is possible, the precise age of the classical site at SZ is still controversial. The aim of this paper is to constrain the chronology of the site by developing a longer magnetostratigraphic record. For this purpose, we carried out an exhaustive sampling in a new succession at SZ. Our results provide a ~65 m magnetostratigraphic record in which 4 magnetozones of normal polarity are found. Our new magnetostratigraphic data suggest an age range between 300-480 Kyr for the lithic tools, closer to the age of traditional Acheulean sites in Europe.

Miocene small-bodied anthropoid primates from Africa and Eurasia are generally considered to precede the divergence between the two groups of extant catarrhines—hominoids (apes and humans) and Old World monkeys—and are thus viewed as more primitive than the stem ape Proconsul. Here we describe Pliobates cataloniae gen. et sp. nov., a small-bodied (4 to 5 kilograms) primate from the Iberian Miocene (11.6 million years ago) that displays a mosaic of primitive characteristics coupled with multiple cranial and postcranial shared derived features of extant hominoids. Our cladistic analyses show that Pliobates is a stem hominoid that is more derived than previously described small catarrhines and Proconsul. This forces us to reevaluate the role played by small-bodied catarrhines in ape evolution and provides key insight into the last common ancestor of hylobatids (gibbons) and hominids (great apes and humans)., (Copyright © 2015, American Association for the Advancement of Science.)

Current knowledge of the evolution of the postcranial skeleton in the genus Homo is hampered by a geographically and chronologically scattered fossil record. Here we present a complete characterization of the postcranium of the middle Pleistocene paleodeme from the Sima de los Huesos (SH) and its paleobiological implications. The SH hominins show the following: (i) wide bodies, a plesiomorphic character in the genus Homo inherited from their early hominin ancestors; (ii) statures that can be found in modern human middle-latitude populations that first appeared 1.6-1.5 Mya; and (iii) large femoral heads in some individuals, a trait that first appeared during the middle Pleistocene in Africa and Europe. The intrapopulational size variation in SH shows that the level of dimorphism was similar to modern humans (MH), but the SH hominins were less encephalized than Neandertals. SH shares many postcranial anatomical features with Neandertals. Although most of these features appear to be either plesiomorphic retentions or are of uncertain phylogenetic polarity, a few represent Neandertal apomorphies. Nevertheless, the full suite of Neandertal-derived features is not yet present in the SH population. The postcranial evidence is consistent with the hypothesis based on the cranial morphology that the SH hominins are a sister group to the later Neandertals. Comparison of the SH postcranial skeleton to other hominins suggests that the evolution of the postcranium occurred in a mosaic mode, both at a general and at a detailed level.

In this study, a new Early Pleistocene proximal hand phalanx (ATE9-2) from the Sima del Elefante cave site (TE - Sierra de Atapuerca, Spain), ascribed to Homo sp., is presented and comparatively described in the context of the evolution of the genus Homo. The ATE9-2 specimen is especially important because of the paucity of hand bones in the human fossil record during the Early Pleistocene. The morphological and metrical analyses of the phalanx ATE9-2 indicate that there are no essential differences between it and comparator fossil specimens for the genus Homo after 1.3 Ma (millions of years ago). Similar to Sima de los Huesos and Neandertal specimens, ATE9-2 is a robust proximal hand phalanx, probably reflecting greater overall body robusticity in these populations or a higher gracility in modern humans. The age of level TE9 from Sima del Elefante and morphological and metrical studies of ATE9-2 suggest that the morphology of the proximal hand phalanges and, thus, the morphology of the hand could have remained stable over the last 1.2-1.3 Ma. Taking into account the evidence recently provided by a metacarpal from Kaitio (Kenya) from around 1.42 Ma, we argue that modern hand morphology is present in the genus Homo subsequent to Homo habilis., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

Correspondence between temporal lobe sulcal pattern and bony impressions on the middle cranial fossae (MCF) was analyzed. MCF bone remains (SD-359, SD-315, and SD-1219) from the El Sidrón (Spain) neandertal site are analyzed in this context. Direct comparison of the soft and hard tissues from the same individual was studied by means of: 1) dissection of two human heads; 2) optic (white light) surface scans; 3) computed tomography and magnetic resonance of the same head. The inferior temporal sulcus and gyrus are the features most strongly influencing MCF bone surface. The Superior temporal sulcus and middle temporal and fusiform gyri also leave imprints. Temporal lobe form differs between Homo sapiens and neandertals. A wider and larger post-arcuate fossa (posterior limit of Brodmann area 20 and the anterior portion of area 37) is present in modern humans as compared to neandertals. However other traits of the MCF surface are similar in these two large-brained human groups. A conspicuous variation is appreciated in the more vertical location of the inferior temporal gyrus in H. sapiens. In parallel, structures of the lower surface of the temporal lobe are more sagittally orientated. Grooves accommodating the fusiform and the lower temporal sulci become grossly parallel to the temporal squama. These differences can be understood within the context of a supero-lateral deployment of the lobe in H. sapiens, a pattern previously identified (Bastir et al., Nat Commun 2 (2011) 588-595). Regarding dural sinus pattern, a higher incidence of petrosquamous sinus is detected in neandertal samples., (© 2014 Wiley Periodicals, Inc.)

The archaeological karstic infill site of Galería Complex, located within the Atapuerca system (Spain), has produced a large faunal and archaeological record (Homo sp. aff. heidelbergensis fossils and Mode II lithic artefacts) belonging to the Middle Pleistocene. Extended-range luminescence dating techniques, namely post-infrared infrared stimulated luminescence (pIR-IR) dating of K-feldspars and thermally transferred optically stimulated luminescence (TT-OSL) dating of individual quartz grains, were applied to fossil-bearing sediments at Galería. The luminescence dating results are in good agreement with published chronologies derived using alternative radiometric dating methods (i.e., ESR and U-series dating of bracketing speleothems and combined ESR/U-series dating of herbivore teeth), as well as biochronology and palaeoenvironmental reconstructions inferred from proxy records (e.g., pollen data). For the majority of samples dated, however, the new luminescence ages are significantly (∼50%) younger than previously published polymineral thermoluminescence (TL) chronologies, suggesting that the latter may have overestimated the true burial age of the Galería deposits. The luminescence ages obtained indicate that the top of the basal sterile sands (GIb) at Galería have an age of up to ∼370 thousand years (ka), while the lowermost sub-unit containing Mode II Acheulean lithics (base of unit GIIa) was deposited during MIS 9 (mean age = 313±14 ka; n = 4). The overlying units GIIb-GIV, which contain the richest archaeopalaeontological remains, were deposited during late MIS 8 or early MIS 7 (∼240 ka). Galería Complex may be correlative with other Middle Pleistocene sites from Atapuerca, such as Gran Dolina level TD10 and unit TE19 from Sima del Elefante, but the lowermost archaeological horizons are ∼100 ka younger than the hominin-bearing clay breccias at the Sima de los Huesos site. Our results suggest that both pIR-IR and single-grain TT-OSL dating are suitable for resolving Middle Pleistocene chronologies for the Sierra de Atapuerca karstic infill sequences.

Here we describe the vertebral fragments from the partial skeleton IPS18800 of the fossil great ape Hispanopithecus laietanus (Hominidae: Dryopithecinae) from the late Miocene (9.6 Ma) of Can Llobateres 2 (Vallès-Penedès Basin, Catalonia, Spain). The eight specimens (IPS18800.5-IPS18800.12) include a fragment of thoracic vertebral body, three partial bodies and four neural arch fragments of lumbar vertebrae. Despite the retention of primitive features (moderately long lumbar vertebral bodies with slightly concave ventrolateral sides), these specimens display a suite of derived, modern hominoid-like features: thoracic vertebrae with dorsally-situated costal foveae; lumbar vertebrae with non-ventrally-oriented transverse processes originating from a robust pedicle, caudally-long laminae with caudally-oriented spinous process, elliptical end-plates, and moderately stout bodies reduced in length and with no ventral keel. These features, functionally related to orthograde behaviors, are indicative of a broad and shallow thorax with a moderately short and stiff lumbar region in Hispanopithecus. Despite its large body mass (ca. 39-40 kg), its vertebral morphology is more comparable to that of hylobatids and Ateles than to extant great apes. This is confirmed by our morphometric analyses, also indicating that Hispanopithecus most closely resembles Pierolapithecus and Morotopithecus among Miocene apes, whereas Proconsul and Nacholapithecus resemble pronograde monkeys. Only in a few features (craniocaudally short and transversely wide pedicles, transverse processes situated on the pedicle, and slight ventral wedging), Hispanopithecus is more derived towards the extant great ape condition than other Miocene apes. Overall, the vertebral morphology of Hispanopithecus supports previous inferences of an orthograde body plan with suspensory and climbing adaptations. However, given similarities with Ateles and the retention of a longer and more flexible spine than in extant great apes, the Hispanopithecus morphology is also consistent with some degree of above-branch quadrupedalism, as previously inferred from other anatomical regions., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

Excavations of a complex of caves in the Sierra de Atapuerca in northern Spain have unearthed hominin fossils that range in age from the early Pleistocene to the Holocene. One of these sites, the 'Sima de los Huesos' ('pit of bones'), has yielded the world's largest assemblage of Middle Pleistocene hominin fossils, consisting of at least 28 individuals dated to over 300,000 years ago. The skeletal remains share a number of morphological features with fossils classified as Homo heidelbergensis and also display distinct Neanderthal-derived traits. Here we determine an almost complete mitochondrial genome sequence of a hominin from Sima de los Huesos and show that it is closely related to the lineage leading to mitochondrial genomes of Denisovans, an eastern Eurasian sister group to Neanderthals. Our results pave the way for DNA research on hominins from the Middle Pleistocene.

New dental remains of the fossil great ape Anoiapithecus brevirostris are described from the Middle Miocene local stratigraphic series of Abocador de Can Mata (ACM) in els Hostalets de Pierola (Vallès-Penedès Basin, NE Iberian Peninsula). These specimens correspond to maxillary fragments with upper teeth from two female individuals from two different localities: left P(3)-M(1) (IPS41712) from ACM/C3-Aj (type locality; 11.9 Ma [millions of years ago]); and right M(1)-M(2) and left P(4)-M(2) (IPS35027) from ACM/C1-E* (12.3-12.2 Ma). Relative enamel thickness is also computed in the latter individual and re-evaluated in other Middle Miocene hominoids from ACM, in order to better assess their taxonomic affinities. With regard to maxillary sinus development, occlusal morphology, molar proportions and enamel thickness, the new specimens show greater resemblances with the (male) holotype specimen of A. brevirostris. They differ from Pierolapithecus catalaunicus in displaying less inflated crests, a more lingually-located hypocone, and relatively lower-crowned molars; from Dryopithecus fontani, in the relatively thicker enamel and lower-crowned molars; from Hispanopithecus spp., in the more inflated crown bases, less peripheral cusps and more restricted maxillary sinus; and from Hispanopithecus laietanus also in the thicker crests, more restricted occlusal foveae, and relatively lower-crowned molars. The new specimens of A. brevirostris show some slight differences compared with the holotype of this species: smaller size (presumably due to sexual size dimorphism), and less distally-tapering M(2) occlusal contour (which is highly variable in both extant and extinct hominoids). The reported remains provide valuable new evidence on dental intraspecific variation and sexual dimorphism in Anoiapithecus. From a taxonomic viewpoint, they support the distinction of this taxon from both Dryopithecus and Pierolapithecus. From a chronostratigraphic perspective, IPS35027 from ACM/C1-E* enlarges the known temporal distribution of Anoiapithecus, further representing the oldest record (first appearance datum) of hominoids in the Iberian Peninsula., (Copyright © 2013 Elsevier Ltd. All rights reserved.)

The fossil ATD6-69 from Atapuerca, Spain, dated to ca. 900 ka (thousands of years ago) has been suggested to mark the earliest appearance of modern human facial features. However, this specimen is a subadult and the interpretation of its morphology remains controversial, because it is unclear how developmental shape changes would affect the features that link ATD6-69 to modern humans. Here we analyze ATD6-69 in an evolutionary and developmental context. Our modern human sample comprises cross-sectional growth series from four populations. The fossil sample covers human specimens from the Pleistocene to the Upper Paleolithic, and includes several subadult Early Pleistocene humans and Neanderthals. We digitized landmarks and semilandmarks on surface and CT scans and analyzed the Procrustes shape coordinates using multivariate statistics. Ontogenetic allometric trajectories and developmental simulations were employed in order to identify growth patterns and to visualize potential adult shapes of ATD6-69. We show that facial differences between modern and archaic humans are not exclusively allometric. We find that while postnatal growth further accentuates the differences in facial features between Neanderthals and modern humans, those features that have been suggested to link ATD6-69's morphology to modern humans would not have been significantly altered in the course of subsequent development. In particular, the infraorbital depression on this specimen would have persisted into adulthood. However, many of the facial features that ATD6-69 shares with modern humans can be considered to be part of a generalized pattern of facial architecture. Our results present a complex picture regarding the polarity of facial features and demonstrate that some modern human-like facial morphology is intermittently present in Middle Pleistocene humans. We suggest that some of the facial features that characterize recent modern humans may have developed multiple times in human evolution., (Copyright © 2013 Elsevier Ltd. All rights reserved.)

Pliopithecinae and Crouzeliinae (Primates: Pliopithecidae) are distinguished dentally by the sharper crests, more compressed cusps, larger foveae, and narrower molars of the latter. Traditionally, such differences were qualitatively related to increased folivory in crouzeliines. This was subsequently disproved by microwear and shearing crest analyses, indicating that all pliopithecids were soft-fruit eaters, except for the more folivorous crouzeliine Barberapithecus. This seems however at odds with the occlusal morphology of the latter, intermediate between those of Pliopithecus and the more derived crouzeliine Anapithecus. To further assess dietary evolution in this group, we report results of dental microwear for two Iberian pliopithecids: Pliopithecus canmatensis, from several Abocador de Can Mata localities (11.8-11.7 Ma, MN8), and Barberapithecus huerzeleri from Castell de Barberà (ca. 11.2-10.5 Ma, MN8, or MN9). Contrary to previously published results, our analyses suggest that all pliopithecids, including Barberapithecus, had a frugivorous diet with a significant sclerocarpic component-apparently more marked in some pliopithecines (such as P. canmatensis) than in the crouzeliine Anapithecus. If our interpretation is correct, it would mean that, within the framework of a frugivorous diet with some hard-object feeding, crouzeliine dental evolution would have been driven by selection pressures towards increased soft-fruit consumption instead of folivory. Dental differences between pliopithecids and hominoids with a significant sclerocarpic component (i.e., orangutans) might be related to phylogenetic constraints, different food-processing methods and/or fracture toughness of hard foods consumed. Although additional research would be required, results suggest that dietary niche partitioning played a significant role in the radiation of European pliopithecids., (Copyright © 2013 Wiley Periodicals, Inc.)

Here we describe the pelvis from the holotype specimen of Pierolapithecus catalaunicus (IPS-21350), a middle Miocene (11.9 Ma [millions of years ago]) stem hominid from Barranc de Can Vila 1 in Abocador de Can Mata (Catalonia, Spain) for which a partial skeleton is preserved. This skeleton includes the oldest known great ape pelvic materials, which have implications for reconstructing the basal hominid body plan from which later hominids evolved. The P. catalaunicus pelvis is composed of a fragment from the ilium (IPS-21350.38) and one from the ischium (IPS-21350.39). Although the P. catalaunicus ischium consists of just a small fragment from the caudal acetabulum, the preserved morphology is consistent with the basal hominoid Proconsul nyanzae. The ilium is similar to Pr. nyanzae in its concave gluteal surface and linea arcuata form, which suggests much of the iliac and pubic form was primitive. However, the ilium was relatively more flaring than Pr. nyanzae and most monkeys, and could be within the range of extant lesser apes. In addition, the iliac tuberosity width was probably intermediate between extant great apes and monkeys, although maximum and minimum estimates could be accommodated within either group. The P. catalaunicus ilium reflects incipient modifications of the basal hominoid torso for the more frequent use of orthograde behaviors described for this taxon on the basis of other preserved anatomical regions, and also supports claims that extant ape pelvic morphology could be homoplastic given the hypothesized phylogenetic positions of Pierolapithecus., (Copyright © 2013 Elsevier Ltd. All rights reserved.)

IPS18800 is a partial skeleton attributed to the fossil great ape Hispanopithecus laietanus, and dated to 9.6 Ma (millions of years ago). Previous studies on the postcranial anatomy of this taxon have shown that it displayed a derived, extant great ape-like orthograde body plan with suspensory adaptations, uniquely coupled with adaptations for above-branch pronograde locomotion. Here, for the first time, we describe and analyze in detail the distal tibia of the IPS18800 skeleton of Hispanopithecus with the aid of three-dimensional geometric morphometrics based on 53 landmarks and semilandmarks collected on a broad sample of extant catarrhines and fossil hominoids. Results of principal components and canonical variate analyses reveal that the distal tibia of Hispanopithecus occupies a unique position in the morphospace, similar in some respects to pronograde monkeys, and in other respects to extant apes. The IPS18800 distal tibia combines adaptations for above branch quadrupedalism, such as a keeled trochlear surface and strong intercollicular groove, with adaptations for vertical climbing, such as an anteroposteriorly flattened shaft, enlarged fibular facet and a tibial stop. These results on the distal tibia agree with those from other anatomical regions, indicating that this taxon displayed a locomotor repertoire unlike any extant ape, combining vertical climbing and clambering with above-branch quadrupedalism., (Copyright © 2013 Elsevier Ltd. All rights reserved.)

Currently restricted to Southeast Asia and Africa, extant hominoids are the remnants of a group that was much more diverse during the Miocene. Apes initially diversified in Africa during the early Miocene, but by the middle Miocene they extended their geographical range into Eurasia, where they experienced an impressive evolutionary radiation. Understanding the role of Eurasian hominoids in the origin and evolution of the great-ape-and-human clade (Hominidae) is partly hampered by phylogenetic uncertainties, the scarcity and incompleteness of fossil remains, the current restricted diversity of the group, and pervasive homoplasy. Nevertheless, scientific knowledge of the Eurasian hominoid radiation has significantly improved during the last decade. In the case of Western Europe, this has been due to the discovery of new remains from the Vallès-Penedès Basin (Catalonia, Spain). Here, I review the fossil record of Vallès-Penedès apes and consider its implications. Although significant disagreements persist among scholars, some important lessons can be learned regarding the evolutionary history of the closest living relatives of humans., (Copyright © 2012 Wiley Periodicals, Inc.)

This paper presents and describes new foot fossils from the species Homo antecessor, found in level TD6 of the site of Gran Dolina (Sierra de Atapuerca, Burgos, Spain). These new fossils consist of an almost complete left talus (ATD6-95) and the proximal three-quarters of a right fourth metatarsal (ATD6-124). The talus ATD6-95 is tentatively assigned to Hominin 10 of the TD6 sample, an adult male specimen with which the second metatarsal ATD6-70+107 (already published) is also tentatively associated. Analysis of these fossils and other postcranial remains has made possible to estimate a stature similar to those of the specimens from the Middle Pleistocene site of Sima de los Huesos (Sierra de Atapuerca, Burgos, Spain). The morphology of the TD6 metatarsals does not differ significantly from that of modern humans, Neanderthals and the specimens from Sima de los Huesos. Talus ATD6-95, however, differs from the rest of the comparative samples in being long and high, having a long and wide trochlea, and displaying a proportionally short neck., (Copyright © 2012 Elsevier Ltd. All rights reserved.)

The internal (nasal and paranasal) cranial anatomy of the Middle Miocene (11.9 Ma [millions of years ago]) great ape Pierolapithecus catalaunicus (Hominidae: Dryopithecini) is described on the basis of computed-tomography scans of the holotype specimen (IPS21350), with particular emphasis on its phylogenetic implications. Pierolapithecus displays the following characters: an anteriorly-restricted maxillary sinus that posteriorly spreads towards the ethmoidal area (thus resembling the pongine condition), although being situated well above the molar roots (as in kenyapithecins, other dryopithecins and pongines); lack of frontal sinus (a synapomorphy of derived pongines, independently acquired by both cercopithecoids and hylobatids); posteriorly-situated turbinals (as in Pongo); anteriorly-projecting nasolacrimal canal (as in Pongo); and probably stepped nasal floor with non-overlapping premaxillary-maxillary contact (as in dryopithecines and stem hominoids, although it cannot be conclusively shown due to bone damage). Overall, Pierolapithecus displays a mosaic of primitive hominid and derived pongine features that are inconsistent with this taxon being a hominine (as previously suggested). Two alternative phylogenetic interpretations are possible: Pierolapithecus may be a stem member of the Hominidae as previously suggested in its original description, or alternatively this taxon may be a stem member of the Ponginae s.l. (with the European dryopithecines being the sister taxon to the Asian pongines)., (Copyright © 2012 Elsevier Ltd. All rights reserved.)

Here we report 12 teeth of the fossil great ape Hispanopithecus (Hominidae: Dryopithecinae: Hispanopithecini), recovered in 2011 from the locality of Can Llobateres 1 (MN9, early Vallesian, Late Miocene, ca. 9.7 Ma [millions of years ago]) in the Vallès-Penedès Basin (Catalonia, Spain). Besides an isolated dP(3) from layer CLL1.1b in the eastern (classical) sector of the site, all of the remaining teeth come from facies CLL1.0 (roughly equivalent to CLL1.2 and CLL1.1b), located in the newly excavated western sector, and representing at least two different individuals. Based on facet congruence and degree of wear, all of the upper cheek teeth, a central incisor and a lateral incisor most likely correspond to a single young adult individual of unknown sex, whereas a very worn I(2) and a female C(1) represent one or two additional individual(s). Morphological and metrical comparisons allow us to attribute these remains to Hispanopithecus laietanus, which is the single hominoid species recognized at CLL1. The newly described teeth represent a significant addition to the hypodigm of this taxon, enabling us to more completely assess the degree of variation displayed by several teeth. In light of the new specimens, the previous tooth position assignment of several upper molars from Can Llobateres and Can Poncic is revised, and the criteria employed to distinguish Hispanopithecus crusafonti from H. laietanus are critically evaluated. On the basis of the available upper cheek teeth from these localities, a distinction at the species level between both samples is tentatively favored, mainly on the basis of P(3), M(1) and M(2) proportions as well as I(1) lingual morphology and proportions. The results of the 2011 field season unambiguously confirm that hominoid-bearing fossiliferous layers from CLL1 are not exhausted. Additional excavations at this site are promising for the discovery of additional remains of H. laietanus in the near future., (Copyright © 2012 Elsevier Ltd. All rights reserved.)

Systematic excavations carried out since 1989 at Bolomor Cave have led to the recovery of four Pleistocene human fossil remains, consisting of a fibular fragment, two isolated teeth, and a nearly complete adult parietal bone. All of these specimens date to the late Middle and early Late Pleistocene (MIS 7-5e). The fibular fragment shows thick cortical bone, an archaic feature found in non-modern (i.e. non-Homo sapiens) members of the genus Homo. Among the dental remains, the lack of a midtrigonid crest in the M(1) represents a departure from the morphology reported for the majority of Neandertal specimens, while the large dimensions and pronounced shoveling of the marginal ridges in the C(1) are similar to other European Middle and late Pleistocene fossils. The parietal bone is very thick, with dimensions that generally fall above Neandertal fossils and resemble more closely the Middle Pleistocene Atapuerca (SH) adult specimens. Based on the presence of archaic features, all the fossils from Bolomor are attributed to the Neandertal evolutionary lineage., (Copyright © 2012 Elsevier Ltd. All rights reserved.)

Atapuerca, in the north of Spain, is the archaeological site where the oldest hominid remains within Europe have been found. In 2008 a jaw fragment, corresponding to the symphyseal area, was discovered in the area called the 'Elephant's pit'. Its age has been estimated at 1.2 million years and it is considered to be the oldest human fossil found in Europe and is from the lower Pleistocene. This work analyses the dental and skeletal damage to the specimen, detected in a macroscopic study of possible horizontal and vertical bone loss at the level of support of the remaining teeth. The limited presence of dental scale, the pattern of destruction and the decreased bone density due to increased marrow spaces suggest the presence of possible periodontal disease.

This paper continues the series of articles initiated in 2006 that analyse hominin dental crown morphology by means of geometric morphometric techniques. The detailed study of both upper premolar occlusal morphologies in a comprehensive sample of hominin fossils, including those coming from the Gran Dolina-TD6 and Sima de los Huesos sites from Atapuerca, Spain, complement previous works on lower first and second premolars and upper first molars. A morphological gradient consisting of the change from asymmetric to symmetric upper premolars and a marked reduction of the lingual cusp in recent Homo species has been observed in both premolars. Although percentages of correct classification based on upper premolar morphologies are not very high, significant morphological differences between Neanderthals (and European middle Pleistocene fossils) and modern humans have been identified, especially in upper second premolars. The study of morphological integration between premolar morphologies reveals significant correlations that are weaker between upper premolars than between lower ones and significant correlations between antagonists. These results have important implications for understanding the genetic and functional factors underlying dental phenotypic variation and covariation., (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.)

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.)

Considerations of Neandertal geographical variation have been hampered by the dearth of remains from Mediterranean Europe and the absence there of sufficiently complete associated postcrania. The 2006 and 2007 excavation of an articulated partial skeleton of a small adult female Neandertal at the Sima de las Palomas, Murcia, southeastern Spain (Sima de las Palomas 96) provides substantial and secure information on body proportions among southern European Neandertals, as well as further documenting the nature of Neandertal biology in southern Iberia. The remains exhibit a suite of cranial, mandibular, dental, and postcranial features, of both Neandertals and archaic Homo generally, that distinguish them from contemporary and subsequent early modern humans. Its lower limbs exhibit the robustness of later Pleistocene Homo generally, and its upper limbs conform to the pattern of elevated robustness of the Neandertals. Its body proportions, including relative clavicular length, distal limb segment lengths, and body mass to stature indicators, conform to the "cold-adapted" pattern of more northern Neandertals. Palomas 96 therefore documents the presence of a suite of "Neandertal" characteristics in southern Iberia and, along with its small body size, the more "Arctic" body proportions of other European Neandertals despite the warmer climate of southern Iberia during marine isotope stage 3.