The Atapuerca human fossils

After 20 years of research, the Atapuerca sites have provided a large amount of archaeological and palaeontological remains. Human fossils have been found in three sites: Gran Dolina, galería and Sima de los Huesos. The Early Pleistocene human fossils from Gran Dolina have been ascribed to a new species,Homo antecessor, that represent the last common ancestor of Neandertals and modern humans. The Sima de los Huesos fossils and all the European Middle Pleistocene human fossils are the ancestors exclusively of the Neandertals, which evolved in Europe in conditions of geographic and genetic isolation.     

The medial pterygoid tubercle in the Atapuerca Early and Middle Pleistocene mandibles: Evolutionary implications

Numerous studies have attempted to identify the presence of uniquely derived (autoapomorphic) Neandertal features. Here, we deal with the medial pterygoid tubercle (MTP), which is usually present on the internal face of the ascending ramus of Neandertal specimens. Our study stems from the identification of a hypertrophied tubercle in ATD6‐96, an Early Pleistocene mandible recovered from the TD6 level of the Atapuerca‐Gran Dolina site and attributed to Homo antecessor. Our review of the literature and study of numerous original fossil specimens and high quality replicas confirm that the MTP occurs at a high frequency in Neandertals (ca. 89%) and is also present in over half (ca. 55%) of the Middle Pleistocene Sima de los Huesos (SH) hominins. In contrast, it is generally absent or minimally developed in other extinct hominins, but can be found in variable frequencies (<ca. 25%) in Pleistocene and recent H. sapiens samples. The presence of this feature in ATD6‐96 joins other traits shared by H. antecessor, the SH hominins and Neandertals. Since the TD6 hominins have been attributed either to MIS 21 or to MIS 25, it seems that a suite of assumed derived Neandertal features appeared in the Early Pleistocene, and they should be interpreted as synapomorphies shared among different taxa. We suggest that H. antecessor, the SH hominins and Neandertals shared a common ancestor in which these features appeared during the Early Pleistocene. The presence of the MTP in taxa other than H. neanderthalensis precludes this feature from being a Neandertal autapomorphy. Am J Phys Anthropol 156:102–109, 2015 © 2014 Wiley Periodicals, Inc.     

The Atapuerca sites and their contribution to the knowledge of human evolution in Europe

Over the last two decades, the Pleistocene sites of the Sierra de Atapuerca (Spain) have provided two extraordinary assemblages of hominin fossils that have helped refine the evolutionary story of the genus Homo in Europe. The TD6 level of the Gran Dolina site has yielded about one hundred remains belonging to a minimum of six individuals of the species Homo antecessor. These fossils, dated to the end of the Lower Pleistocene (800 kyr), provide the earliest evidence of hominin presence in Western Europe. The origin of these hominins is unknown, but they may represent a speciation event from Homo ergaster/Homo erectus. The TD6 fossils are characterized by a significant increase in cranial capacity as well as the appearance of a “sapiens” pattern of craniofacial architecture. At the Sima de los Huesos site, more than 4,000 human fossils belonging to a minimum of 28 individuals of a Middle Pleistocene population (ca. 500–400 kyr) have been recovered. These hominins document some of the oldest evidence of the European roots of Neanderthals deep in the Middle Pleistocene. Their origin would be the dispersal out of Africa of a hominin group carrying Mode 2 technologies to Europe. Comparative study of the TD6 and Sima de la Huesos hominins suggests a replacement model for the European Lower Pleistocene population of Europe or interbreeding between this population and the new African emigrants.     

The costal skeleton of Homo antecessor: preliminary results

The Lower Pleistocene TD6 level at the Gran Dolina site in the Sierra de Atapuerca (Burgos, Spain) has yielded nine ribs that represent a minimum of three individuals of the species, Homo antecessor. We present a detailed morphological and metric study of these costal elements, including the siding and anatomical position of all of the rib remains. The adult or nearly adult ribs are also metrically compared with other fossil hominins and with modern comparative samples. The costal elements recovered to date from the TD6 level at Gran Dolina can neither confirm nor reject the hypothesis that H. antecessor had a large thorax, similar to that of Neandertals. However, the fragmentary evidence of the H. antecessor thoracic skeleton is not inconsistent with this suggestion based on other skeletal elements, such as clavicles.

Resumen

En el nivel TD6 del Pleistoceno inferior del yacimiento de Gran Dolina, en la Sierra de Atapuerca (Burgos, España) se han recuperado nueve costillas que pertenecen a un mínimo de tres individuos de la especie Homo antecessor. Presentamos un detallado estudio métrico y morfológico incluyendo el lado y la determinación anatómica. Las costillas pertenecientes a individuos adultos o casi adultos también son comparadas métricamente a muestras modernas de comparación y otros homininos fósiles. Basándonos en el registro de costillas de Homo antecessor recuperado hasta el momento no podemos probar ni refutar la hipótesis de que esta especie presentaba un tórax grande similar al de los Neandertales. Sin embargo, el registro de costillas no es inconsistente con la hipótesis de un tórax grande como sugiere la gran longitud de sus clavículas.     

New foot remains from the Gran Dolina-TD6 Early Pleistocene site (Sierra de Atapuerca, Burgos, Spain)

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.     

Stature estimation from complete long bones in the Middle Pleistocene humans from the Sima de los Huesos, Sierra de Atapuerca (Spain)

Systematic excavations at the site of the Sima de los Huesos (SH) in the Sierra de Atapuerca (Burgos, Spain) have allowed us to reconstruct 27 complete long bones of the human species Homo heidelbergensis. The SH sample is used here, together with a sample of 39 complete Homo neanderthalensis long bones and 17 complete early Homo sapiens (Skhul/Qafzeh) long bones, to compare the stature of these three different human species. Stature is estimated for each bone using race- and sex-independent regression formulae, yielding an average stature for each bone within each taxon. The mean length of each long bone from SH is significantly greater (p < 0.05) than the corresponding mean values in the Neandertal sample. The stature has been calculated for male and female specimens separately, averaging both means to calculate a general mean. This general mean stature for the entire sample of long bones is 163.6 cm for the SH hominins, 160.6 cm for Neandertals and 177.4 cm for early modern humans. Despite some overlap in the ranges of variation, all mean values in the SH sample (whether considering isolated bones, the upper or lower limb, males or females or more complete individuals) are larger than those of Neandertals. Given the strong relationship between long bone length and stature, we conclude that SH hominins represent a slightly taller population or species than the Neandertals. However, compared with living European Mediterranean populations, neither the Sima de los Huesos hominins nor the Neandertals should be considered ‘short’ people. In fact, the average stature within the genus Homo seems to have changed little over the course of the last two million years, since the appearance of Homo ergaster in East Africa. It is only with the emergence of H. sapiens, whose earliest representatives were ‘very tall’, that a significant increase in stature can be documented.     

Twentieth anniversary of Homo antecessor (1997‐2017): a review

It has been twenty years since diagnosis and publication of the species Homo antecessor.¹ Since then, new human fossils recovered from the TD6 level of the Gran Dolina site (Sierra de Atapuerca, northern Spain) have helped to refine its taxonomic and phylogenetic position. In this paper, we present a synthesis of the most characteristic features of this species, as well as our interpretation derived from the latest investigations. We focus on the phylogenetic interpretation of Homo antecessor, taking into account the most recent paleogenetic analyses and a reassessment of the European Middle Pleistocene hominin record. We try to show that, twenty years after its publication, H. antecessor provides a good opportunity to address the morphology of the last common ancestor of Neandertals and modern humans.     

Brief Communication: Lumbar lordosis in extinct hominins: Implications of the pelvic incidence

Recently, interest has peaked regarding the posture of extinct hominins. Here, we present a new method of reconstructing lordosis angles of extinct hominin specimens based on pelvic morphology, more specifically the orientation of the sacrum in relation to the acetabulum (pelvic incidence). Two regression models based on the correlation between pelvic incidence and lordosis angle in living hominoids have been developed. The mean values of the calculated lordosis angles based on these models are 36°?45° for australopithecines, 45°?47° for Homo erectus, 27°?34° for the Neandertals and the Sima de los Huesos hominins, and 49°?51° for fossil H. sapiens. The newly calculated lordosis values are consistent with previously published values of extinct hominins (Been et al.: Am J Phys Anthropol 147 (2012) 64–77). If the mean values of the present nonhuman hominoids are representative of the pelvic and lumbar morphology of the last common ancestor between humans and nonhuman hominoids, then both pelvic incidence and lordosis angle dramatically increased during hominin evolution from 27° ± 5 to 22° ± 3 (respectively) in nonhuman hominoids to 54° ± 10 and 51° ± 11 in modern humans. This change to a more human‐like configuration appeared early in the hominin evolution as the pelvis and spines of both australopithecines and H. erectus show a higher pelvic incidence and lordosis angle than nonhuman hominoids. The Sima de los Huesos hominins and Neandertals show a derived configuration with a low pelvic incidence and lordosis angle. Am J Phys Anthropol 154:307–314, 2014. © 2014 Wiley Periodicals, Inc.     

Lower limb bone structure of Middle Pleistocene hominins from the Caune de l’Arago (Tautavel,France): Evolutionary and functional comparison with the penecontemporaneous hominins of Sima de los Huesos (Atapuerca,Spain)

The Caune de l’Arago (Arago) and Sima de los Huesos (Sima) human bones from the European Middle Pleistocene are penecontemporaneous, although the Sima hominins are closely related to Neandertal, and Arago hominins present more archaic features. In previous and in press studies, the cross-sectional geometric properties (CSG) of lower limb bones of Arago and Sima have been studied separately without comparative analyses. Here, in order to bridge this gap, we use the same criteria for both samples to highlight evolutionary affinities and to compare their level and pattern of mobility. This study focuses on the femur, fibula and tibia from Arago and Sima with references to fossils from the sites of Trinil, Zhoukoudian and Lazaret, and ancient and recent Homo sapiens (including athletes and non-athletes). We analyze the cross-sectional areas, biomechanical bone “shape” indices (Ix/Iy, Imax/Imin) and pattern of cortical bone distribution. All lower limb bones from Arago have noticeably high to very high relative cortical areas and low to very low medullary areas. The overall femoral pattern in Arago, like Sima, is similar to that of Middle Pleistocene hominins (e.g., low femoral shape indices, Ix/Iy) and Neandertals (e.g., large cross-sectional size). However, the femoral midshaft in Sima presents prominent posteromedial cortical thickening, as a result of a spiral cortical reinforcement along the medial side of the diaphysis. This characteristic is specific to Neandertal and some Middle Pleistocene hominins. In contrast, the midshaft femoral pattern in Arago is close to that of some Homo erectus. We also note that the femoral cross-sectional size and relative cortical area in Arago differ drastically to the small size and low relative cortical area of Lazaret and Trinil hominins. The very high shape index at midshaft (i.e., high Imax/Imin) of the Arago tibia is observed in ancient H. sapiens and runners; the tibial posterior “pilaster” is found in Neandertals and ancient H. sapiens; and the flat or convex tibial faces are similar to Neandertals. Furthermore, the Arago fibulas show marked fibular posterolateral cortical reinforcement with low anteroposterior strengthening. These leg features (tibia, fibula) are also found in some Sima hominins (but not in all individuals). Consequently, this study confirms the presence of archaic features in Arago and the close evolutionary relationship between Sima and Neandertal. This proposition is mainly based on the femoral midshaft pattern influenced by the pelvofemoral complex, considered to be substantially genetically controlled. The leg functional analysis highlights a high level of mobility and travelling in uneven terrains or in mountainous areas in Arago, consistent with known environments and hunting practices. Previously, an analogous hypothesis was proposed put forward for Sima hominins.     

Metric and morphological study of the upper cervical spine from the Sima de los Huesos site (Sierra de Atapuerca, Burgos, Spain)

In this article, the upper cervical spine remains recovered from the Sima de los Huesos (SH) middle Pleistocene site in the Sierra de Atapuerca (Burgos, Spain) are described and analyzed. To date, this site has yielded more than 5000 human fossils belonging to a minimum of 28 individuals of the species Homo heidelbergensis. At least eleven individuals are represented by the upper cervical (C1 and C2) specimens: six adults and five subadults, one of which could represent an adolescent individual. The most complete adult vertebrae (three atlases and three axes) are described, measured, and compared with other fossil hominins and modern humans. These six specimens are associated with one another and represent three individuals. In addition, one of these sets of cervical vertebrae is associated with Cranium 5 (Individual XXI) from the site. The metric analysis demonstrates that the Sima de los Huesos atlases and axes are metrically more similar to Neandertals than to our modern human comparative sample. The SH atlases share with Neandertals a sagittally elongated canal. The most remarkable feature of the SH (and Neandertal) axes is that they are craniocaudally low and mediolaterally wide compared to our modern male sample. Morphologically, the SH sample shares with Neandertals a higher frequency of caudally projected anterior atlas arch, which could reflect greater development of the longus colli muscle. In other features, such as the frequency of weakly developed tubercles for the attachment of the transverse ligament of the atlas, the Sima de los Huesos fossils show intermediate frequencies between our modern comparative samples and the Neandertals, which could represent the primitive condition. Our results are consistent with the previous phylogenetic interpretation of H. heidelbergensis as an exclusively European species, ancestral only to H. neanderthalensis.     

A geometric morphometric analysis of hominin upper second and third molars, with particular emphasis on European Pleistocene populations

The study of dental morphology by means of geometric morphometric methods allows for a detailed and quantitative comparison of hominin species that is useful for taxonomic assignment and phylogenetic reconstruction. Upper second and third molars have been studied in a comprehensive sample of Plio- and Pleistocene hominins from African, Asian and European sites in order to complete our analysis of the upper postcanine dentition. Intraspecific variation in these two molars is high, but some interspecific trends can be identified. Both molars exhibit a strong reduction of the distal cusps in recent hominin species, namely European Homo heidelbergensis, Homo neanderthalensis and Homo sapiens, but this reduction shows specific patterns and proportions in the three groups. Second molars tend to show four well developed cusps in earlier hominin species and their morphology is only marginally affected by allometric effects. Third molars can be incipiently reduced in earlier species and they evince a significant allometric component, identified both inter- and intraspecifically. European Middle Pleistocene fossils from Sima de los Huesos (SH) show a very strong reduction of these two molars, even more marked than the reduction observed in Neanderthals and in modern human populations. The highly derived shape of SH molars points to an early acquisition of typical Neanderthal dental traits by pre-Neanderthal populations and to a deviation of this population from mean morphologies of other European Middle Pleistocene groups.     

Enamel and dentine dimensions of the Pleistocene hominins from Atapuerca (Burgos,Spain): A comparative study of canine teeth

Enamel and dentin patterns have awakened a considerable interest in phylogenetic studies. However, almost nothing is known about the dental tissue proportions of European Pleistocene hominins, apart from Neanderthal populations. This study aims to assess the three-dimensional dental tissue proportions of permanent canines belonging to the extensive sample of hominin teeth at Sierra de Atapuerca (Spain) through the use of microtomographic techniques. Our results show that early and middle Pleistocene populations from Atapuerca exhibit large coronal and root dentine dimensions, as well as a thinly enamelled pattern, which has been traditionally considered an autapomorphic Neanderthal trait. Therefore, these results might support an early enamel thickness decrease which is already observed 800 kyr ago in Homo antecessor and maintained in later groups such as Sima de los Huesos and Neanderthal populations during the middle Pleistocene.     

Right‐handed fossil humans

Fossil hominids often processed material held between their upper and lower teeth. Pulling with one hand and cutting with the other, they occasionally left impact cut marks on the lip (labial) surface of their incisors and canines. From these actions, it possible to determine the dominant hand used. The frequency of these oblique striations in an array of fossil hominins documents the typically modern pattern of 9 right‐ to 1 left‐hander. This ratio among living Homo sapiens differs from that among chimpanzees and bonobos and more distant primate relatives. Together, all studies of living people affirm that dominant right‐handedness is a uniquely modern human trait. The same pattern extends deep into our past. Thus far, the majority of inferred right‐handed fossils come from Europe, but a single maxilla from a Homo habilis, OH‐65, shows a predominance of right oblique scratches, thus extending right‐handedness into the early Pleistocene of Africa. Other studies show right‐handedness in more recent African, Chinese, and Levantine fossils, but the sample compiled for non‐European fossil specimens remains small. Fossil specimens from Sima del los Huesos and a variety of European Neandertal sites are predominately right‐handed. We argue the 9:1 handedness ratio in Neandertals and the earlier inhabitants of Europe constitutes evidence for a modern pattern of handedness well before the appearance of modern Homo sapiens.     

Intrapopulational body size variation and cranial capacity variation in middle pleistocene humans: The Sima de los Huesos sample (Sierra de Atapuerca,Spain)

A sexual dimorphism more marked than in living humans has been claimed for European Middle Pleistocene humans, Neandertals and prehistoric modern humans. In this paper, body size and cranial capacity variation are studied in the Sima de los Huesos Middle Pleistocene sample. This is the largest sample of non-modern humans found to date from one single site, and with all skeletal elements represented. Since the techniques available to estimate the degree of sexual dimorphism in small palaeontological samples are all unsatisfactory, we have used the bootstraping method to asses the magnitude of the variation in the Sima de los Huesos sample compared to modern human intrapopulational variation. We analyze size variation without attempting to sex the specimens a priori. Anatomical regions investigated are scapular glenoid fossa; acetabulum; humeral proximal and distal epiphyses; ulnar proximal epiphysis; radial neck; proximal femur; humeral, femoral, ulnar and tibial shaft; lumbosacral joint; patella; calcaneum; and talar trochlea. In the Sima de los Huesos sample only the humeral midshaft perimeter shows an unusual high variation (only when it is expressed by the maximum ratio, not by the coefficient of variation). In spite of that the cranial capacity range at Sima de los Huesos almost spans the rest of the European and African Middle Pleistocene range. The maximum ratio is in the central part of the distribution of modern human samples. Thus, the hypothesis of a greater sexual dimorphism in Middle Pleistocene populations than in modern populations is not supported by either cranial or postcranial evidence from Sima de los Huesos. Am J Phys Anthropol 106:19–33, 1998. © 1998 Wiley-Liss, Inc.     

The human cranial remains from Gran Dolina Lower Pleistocene site (Sierra de Atapuerca, Spain).

In this article we study the cranial remains of the late Lower Pleistocene human fossils from Gran Dolina (Sierra de Atapuerca, Spain), assigned to the new species Homo antecessor. The cranial remains belong to at least five individuals, both juveniles and adults. The most outstanding feature is the totally modern human morphology of the very complete face ATD6-69, representing the earliest occurrence of the modern face in the fossil record. The Gran Dolina fossils show in the face a suite of modern human apomorphies not found in earlier hominids nor in contemporary or earlier Homo erectus fossils. There are also traits in the Gran Dolina fossils shared with both Neandertals and modern humans, which reinforce the hypothesis that Neandertals and modern humans form a clade, and that the Gran Dolina fossils are a common ancestor to both lineages.     

Talonid crests expression at the enamel–dentine junction of hominin lower permanent and deciduous molars

The application of microtomography (mCT) to dental morphological studies has unveiled a new source of palaeobiological information, particularly in the analysis of the internal structures of teeth. In this study, we assess the expression of talonid crests at the enamel and dentine surfaces in lower permanent and second deciduous molars (M₂ and dm₂) of H. sapiens, H. neanderthalensis and Atapuerca-Sima de los Huesos (SH) hominins. In modern humans, talonid crests are described exclusively in the deciduous teeth (Korenhof, 1982) and interpreted as a primitive mammalian remnant of the talonid attachment to the trigonid. Here we report for the first time the expression of talonid crests of deciduous and permanent molars in H. sapiens, H. neanderthalensis and Middle Pleistocene hominins. We discuss possible evolutionary interpretations and suggest the importance of recording this feature in future studies.     

Pattern of dental development in Hominid XVIII from the Middle Pleistocene Atapuerca-Sima de los Huesos site (Spain)

. We describe the pattern of dental development of Hominid XVIII from the Middle Pleistocene Sima de los Huesos (SH) site of the Sierra de Atapuerca (Burgos, Spain). As expected, this pattern is similar to that of modern humans. A delay of development of the lower and upper canines was observed. In contrast, the relative advanced development of the lower second molars and, especially, the upper and lower third molars is noteworthy. This latter feature seems to be common in Pleistocene hominids, and suggests that the pattern of dental development evolved in the genus Homo during the Pleistocene. In European Middle Pleistocene hominids, this pattern probably was facilitated by the extra space available in the mandible and maxilla for developing teeth.     

Trigonid crests expression in Atapuerca-Sima de los Huesos lower molars: Internal and external morphological expression and evolutionary inferences

Trigonid crest patterning in lower molars is distinctive among Late Pleistocene hominins such as Homo neanderthalensis, fossil Homo sapiens and modern humans. In this paper, we present an examination of trigonid crest patterning in the Middle Pleistocene permanent lower molar sample (n = 62) of Homo heidelbergensis from Sima de los Huesos (SH). Crest expression was assessed from 3D models of the enamel and the dentine surfaces that were produced using micro-computed tomography (microCT). The aims of our analysis are to: 1) characterize the pattern of trigonid crest expression at the outer enamel and enamel-dentine junction surfaces (OES and EDJ) of the SH sample, 2) evaluate the concordance of expression between both surfaces, and 3) place trigonid crest variation in the SH sample into a phylogenetic context. Our results reveal a greater variability in the expression of trigonid crests at the EDJ (14 types) compared to the OES (4 types). Despite this variability, in almost all cases the expression of a continuous mid-trigonid or distal crest at the OES corresponds with the expression of a continuous mesial/mid-trigonid or distal trigonid crest, respectively, at the EDJ. Thus, it is possible to predict the type of trigonid crest pattern that would be at the OES in the case of partially worn teeth. Our study points to increased variability in trigonid crest expression in M₃s compared to M₁s and M₂s. Moreover, our analysis reveals that the SH sample matches broadly the trigonid crest patterns displayed by H. neanderthalensis and differs from those exhibited by H. sapiens, particularly in the almost constant expression of a continuous middle trigonid crest at the EDJ. However, SH hominins also exhibit patterns that have not been reported in H. neanderthalensis and H. sapiens samples. Other aspects of the variability of the trigonid crest expression at the dentine are presented and discussed.     

Early Pleistocene human humeri from the Gran Dolina-TD6 site (Sierra de Atapuerca, Spain)

In this report, we present a morphometric comparative study of two Early Pleistocene humeri recovered from the TD6 level of the Gran Dolina cave site in Sierra de Atapuerca, northern Spain. ATD6-121 belongs to a child between 4 and 6 years old, whereas ATD6-148 corresponds to an adult. ATD6-148 exhibits the typical pattern of the genus Homo, but it also shows a large olecranon fossa and very thin medial and lateral pillars (also present in ATD6-121), sharing these features with European Middle Pleistocene hominins, Neandertals, and the Bodo Middle Pleistocene humerus. The morphology of the distal epiphysis, together with a few dental traits, suggests a phylogenetic relationship between the TD6 hominins and the Neandertal lineage. Given the older geochronological age of these hominins (ca. 900 ka), which is far from the age estimated by palaeogenetic studies for the population divergence of modern humans and Neandertals (ca. 400 ka), we suggest that this suite of derived "Neandertal" features appeared early in the evolution of the genus Homo. Thus, these features are not "Neandertal" apomorphies but traits which appeared in an ancestral and polymorphic population during the Early Pleistocene.     

Human talus bones from the Middle Pleistocene site of Sima de los Huesos (Sierra de Atapuerca,Burgos, Spain)

Here we present and describe comparatively 25 talus bones from the Middle Pleistocene site of the Sima de los Huesos (SH) (Sierra de Atapuerca, Burgos, Spain). These tali belong to 14 individuals (11 adult and three immature). Although variation among Middle and Late Pleistocene tali tends to be subtle, this study has identified unique morphological characteristics of the SH tali. They are vertically shorter than those of Late Pleistocene Homo sapiens, and show a shorter head and a broader lateral malleolar facet than all of the samples. Moreover, a few shared characters with Neanderthals are consistent with the hypothesis that the SH population and Neanderthals are sister groups. These shared characters are a broad lateral malleolar facet, a trochlear height intermediate between modern humans and Late Pleistocene H. sapiens, and a short middle calcaneal facet. It has been possible to propose sex assignment for the SH tali based on their size. Stature estimates based on these fossils give a mean stature of 174.4 cm for males and 161.9 cm for females, similar to that obtained based on the long bones from this same site.