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.     

Human mandibular incisors from the late Middle Pleistocene locality of Hoedjiespunt 1, South Africa.

The Hoedjiespunt 1 locality is an archaeological and palaeontological site located on the Hoedjiespunt Peninsula at Saldanha Bay, South Africa. In 1996 two human teeth, a left central mandibular incisor and a left lateral mandibular incisor, were discovered during excavations in the late Middle Pleistocene palaeontological layers. These teeth are described and are found to belong to a single subadult individual. Despite their developmental stage, these incisors already display early signs of wear. Their crown diameters are larger than modern and archaeological African comparative material and are most closely comparable with crown diameters of an early Middle Pleistocene and late Middle Pleistocene dental sample from Africa, Europe and Asia. In the light of this metrical evidence, data on two previously excavated maxillary molars, most probably belonging to the same individual, were re-examined. It was found that the Hoedjiespunt 1 hominid possessed dental metrical features (large anterior teeth and small molars) comparable with other African and European hominids referred to the Middle Pleistocene.     

Dental size variation in the Atapuerca-SH Middle Pleistocene hominids

The Middle Pleistocene Atapuerca-Sima de los Huesos (SH) site in Spain has yielded the largest sample of fossil hominids so far found from a single site and belonging to the same biological population. The SH dental sample includes a total of 452 permanent and deciduous teeth, representing a minimum of 27 individuals. We present a study of the dental size variation in these hominids, based on the analysis of the mandibular permanent dentition: lateral incisors, n=29; canines, n=27; third premolars, n=30; fourth premolars, n=34; first molars, n=38; second molars, n=38. We have obtained the buccolingual diameter and the crown area (measured on occlusal photographs) of these teeth, and used the bootstrap method to assess the amount of variation in the SH sample compared with the variation of a modern human sample from the Museu Antropologico of the Universidade of Coimbra (Portugal). The SH hominids have, in general terms, a dental size variation higher than that of the modern human sample. The analysis is especially conclusive for the canines. Furthermore, we have estimated the degree of sexual dimorphism of the SH sample by obtaining male and female dental subsamples by means of sexing the large sample of SH mandibular specimens. We obtained the index of sexual dimorphism (ISD=male mean/female mean) and the values were compared with those obtained from the sexed modern human sample from Coimbra, and with data found in the literature concerning several recent human populations. In all tooth classes the ISD of the SH hominids was higher than that of modern humans, but the differences were generally modest, except for the canines, thus suggesting that canine size sexual dimorphism in Homo heidelbergensis was probably greater than that of modern humans. Since the approach of sexing fossil specimens has some obvious limitations, these results should be assessed with caution. Additional data from SH and other European Middle Pleistocene sites would be necessary to test this hypothesis.     

Interproximal grooving in the Atapuerca-SH hominid dentitions

The dental sample recovered from the Sima de los Huesos (SH) Middle Pleistocene cave site of the Sierra de Atapuerca (Spain) includes 296 specimens. Interproximal wear grooves have been observed in 20 maxillary and mandibular posterior teeth belonging to at least five of the 32 individuals identified so far in the SH hypodigm. Interproximal grooving affected only the adults, and at an age between 25 and 40 years. The appearance, morphology, and location pattern of the SH wear grooves are similar to those reported in other fossil hominids and in more recent human populations. Two alternative proposals, the toothpicking and the fiber or sinew processing hypotheses, compete for explaining the formation of this anomalous wear. The characteristics observed in the wear grooves of the SH teeth are compatible only with the habitual probing of interdental spaces by means of hard and inflexible objects. Dietary grit may also have contributed to the abrasion of the root walls during the motion of the dental probes. Am. J. Phys. Anthropol. 102:369–376, 1997. © 1997 Wiley-Liss, Inc.     

Early and Middle Pleistocene hominins from Atapuerca (Spain) show differences in dental developmental patterns

The Bayesian statistical approach considers teeth as forming a developmental module, as opposed to a tooth-by-tooth analysis. This approach has been employed to analyze Upper Pleistocene hominins, including Neandertals and some anatomically modern humans, but never earlier populations. Here, we show its application on five hominins from the TD6.2 level of the Gran Dolina site (Homo antecessor, Early Pleistocene) and the Sima de los Huesos site (Middle Pleistocene) of the Sierra de Atapuerca (Burgos, northern Spain). Our results show an advanced development of the third molars in both populations with respect to modern Homo sapiens. In addition, the Sima de los Huesos hominins differ from H. sapiens and H. antecessor in the relatively advanced development of their second molar. The relative mineralization of I1/M1 in H. antecessor appears to be similar to that of modern humans, as opposed to that of Neandertals, which appear to be unique. These observations, combined with reduced enamel formation times and the advanced development of the third molars, appear to indicate a shorter ontogenetic period in the hominins from Gran Dolina and Sima de los Huesos in comparison to modern human average.     

Right handedness of Homo heidelbergensis from Sima de los Huesos (Atapuerca, Spain) 500,000 years ago

Handedness is a product of brain specialization, which in turn seems to be responsible for the higher cognitive capabilities of humans, such as language and technology. Handedness in living humans is well established and shows the highest degree of manual specialization. Studies on hand laterality in nonhuman primates, particularly in chimpanzees, remain a matter of controversy as results tend to vary depending on factors such as the tasks performed and the environment in which the individuals live. Studies in several disciplines have attempted to determine where in the course of human evolution handedness established itself, with evidence collected from sources such as paleoneurological analyses, stone tool flaking, zooarchaeological studies and dental wear analyses, the last one of which have proven the most reliable source of information. Here we report an experimental and paleoanthropological study on hand laterality of a sample of 28 hominids from Sima de los Huesos (Atapuerca, Spain), dated at about 500,000 years ago, and compare our results with dental microwear analysis in other fossil samples such as that from Krapina (Croatia), as well as modern traditional societies. Our results indicate that European Middle Pleistocene Homo heidelbergensis was already as right-handed as modern populations.     

Earliest humans in Europe: the age of TD6 Gran Dolina, Atapuerca, Spain

Hominid remains found in 1994 from the stratified Gran Dolina karst-filling at the Atapuerca site in NE Spain were dated to somewhat greater than 780 ka based on palaeomagnetic measurements, making these the oldest known hominids in Europe (sensu stricto). We report new ESR and U-series results on teeth from four levels of the Gran Dolina deposit which confirm the palaeomagnetic evidence, and indicate that TD6 (from which the human remains have been recovered) dates to the end of the Early Pleistocene. The results for the other levels are consistent with estimates based mainly on microfaunal evidence, and suggest that TD8, TD10 and TD11 date to the Middle Pleistocene.     

L’approvisionnement en matières premières lithiques au Pléistocène inférieur et moyen dans la Sierra de Atapuerca, Burgos (Espagne)

This paper summarizes the latest studies on lithic raw material raising that are currently undertaken within the Sierra de Atapuerca research project. During the past two years, several works have provided new petrographical and technological data, contributing to the knowledge of the kinds of materials exploited by the Atapuerca hominids, as well as the areas where those materials were raised. The sites yielding this information are Gran Dolina (Lower and Middle Pleistocene) and Galeria (Middle Pleistocene), both located in the Trinchera del Ferrocarril archeopaleontological complex.     

Additional Evidence for Morpho-Dimensional Tooth Crown Variation in a New Indonesian H. erectus Sample from the Sangiran Dome (Central Java)

This contribution reports fifteen human fossil dental remains found during the last two decades in the Sangiran Dome area, in Central Java, Indonesia. Among this sample, only one of the specimens had already been briefly described, with the other fourteen remaining unreported. Seven of the fifteen isolated teeth were found in a secured stratigraphic context in the late Lower-early Middle Pleistocene Kabuh Formation. The remaining elements were surface finds which, based on coincidental sources of information, were inferred as coming from the Kabuh Formation. Mainly constituted of permanent molars, but also including one upper incisor and one upper premolar, this dental sample brings additional evidence for a marked degree of size variation and time-related structural reduction in Javanese H. erectus. This is notably expressed by a significant decrease of the mesiodistal diameter, frequently associated to the reduction or even loss of the lower molar distal cusp (hypoconulid) and to a more square occlusal outline. In addition to the hypoconulid reduction or loss, this new sample also exhibits a low frequency of the occlusal Y-groove pattern, with a dominance of the X and, to a lesser extent, of the+patterns. This combination is rare in the Lower and early Middle Pleistocene paleoanthropological record, including in the early Javanese dental assemblage from the Sangiran Dome. On the other hand, similar dental features are found in Chinese H. erectus and in H. heidelbergensis. As a whole, this new record confirms the complex nature of the intermittent exchanges that occurred between continental and insular Southeast Asia through the Pleistocene.     

东亚中更新世古人类下颔第二臼齿釉质-齿质连接面三维形状和釉质厚度分布

东亚中更新世古人类在头骨、下颌骨、牙齿等解剖部位表现出不同程度的形态多样性,中期成员代表为直立人,而晚期成员的演化地位具有较大争议。为进一步了解东亚中更新世古人类内部的形态变异特点和为东亚中更新世晚期古人类分类提供依据,本文使用微分同胚的表面匹配（Diffeomorphic Surface Matching, DSM）和形态测量图（Morphometric map）对下颌第二臼齿（M₂）釉质-齿质连接面的形状和齿冠侧面釉质厚度分布模式进行了量化分析。结果显示：1）东亚中更新世古人类与晚期人属成员（尼安德特人和现代人）存在较明显的形态差别;2）该时段晚期的东亚古人类相对中期直立人在侧面釉质厚度分布规律上具有独特性,并在釉质-齿质连接面的三维形状上与晚期人属成员更加接近。本文在以往对东亚中更新世古人类牙齿内外结构单个性状研究的基础上,使用三维形态测量方法进一步量化了M₂釉质-齿质连接面三维形状和侧面釉质厚度分布模式两项重要特征的变异特点,这对未来该时段同类型牙齿的形态鉴定以及解决东亚中更新世晚期古人类的分类地位具有一定意义。     

Odontometry of a Spanish neolithic-Bronze age sample. Comparison with other populations of the iberian peninsula. I.-Permanent teeth

The present study concerns the morphometric characteristics of two hundred permanent teeth from the Neolithic-Bronze Age human remains found in «Galería del Sílex» (Atapuerca Cave, Burgos, Spain). The crown index reveals that all the upper maxillary teeth (except incisors) and the mandibular first incisors, canines and premolars have a vestibulo-lingual diameter greater than the mesio-distal. The dental module and the surface value show that the mandibular teeth (except the molars) are smaller than those of the upper maxillary. These results are very similar to other European populations. Finally, the Sílex Gallery sample was compared with five other series of the Iberian Peninsula which have a chronology from the Mesolithic to the Middle Age. The results indicate a general decrease for both diameters (ØMD and ØVL) throughout time.     

A geometric morphometric analysis of hominin upper premolars. Shape variation and morphological integration

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.     

Brief communication: on the optimum number of metacarpals for roentgenogrammetric measurement

Interpretation of dental development of fossil hominids requires understanding of and comparison with the pattern and timing of dental development among living humans and pongids. We report the first study of crown and root calcification in the lower permanent molar teeth among chimpanzees (Pan troglodytes) of known chronological age. A series of 99 lateral head radiographs of 16 captive-born chimpanzees were analyzed. Radiographs were taken at irregular intervals throughout the entire postnatal period of dental development from birth to 13 years of age. Permanent mandibular molars were rated on an eight-point maturation scale from initial radiographic appearance through crown and root calcification and apical closure of the root canals. In addition, we were able to document initial crown calcification and completion, as well as root completion and apical closure in incisors, canines, and premolars. Our results show several differences from the widely cited developmental schedule for pongid dentitions of Dean and Wood (Folia Primatol. 36:111–127, 1981). We found a much greater degree of temporal overlap in calcification of the crowns of adjacent molars, a pattern very unlike that usually seen in human dental development, which is characterized by delays between the onset of crown calcification in the molar series. Also, the ages and durations of crown and root formation in our chimp sample differ from the estimates proposed by Dean and Wood. By more clearly establishing the nature of developmental schedules and the timing of major events in the pongid dentition, these results should aid in the ongoing controversies concerning the human or pongid nature of dental development among Plio-Pleistocene hominids.     

Tissue proportions and enamel thickness distribution in the early Middle Pleistocene human deciduous molars from Tighenif,Algeria

The present study of three human upper deciduous molars from the early Middle Pleistocene site of Tighenif, Algeria, constitutes the first microtomographic-based endostructural exploration of African fossil teeth likely representative of the Homo heidelbergensis morph. Comparative morphological observations and 2-3D measurements describing subtle tooth organization (crown tissue proportions) and enamel thickness topography (site-specific distribution and global patterning) indicate that their virtual extracted structural signature better fits the modern human, rather than the Neanderthal condition. Accordingly, we predict that the inner structural morphology of the deciduous molars from the Middle Pleistocene western European series better fits the primitive, and not the derived Neanderthal figures.     

Later Middle Pleistocene human remains from the Almonda Karstic system, Torres Novas, Portugal

Later Middle Pleistocene archeological deposits of the Galeria Pesada (Gruta da Aroeira), Almonda Karstic System, Torres Novas, Portugal, yielded two archaic human teeth, a mandibular canine and a maxillary third molar. The C(1)presents moderate and asymmetrical shoveling with a stout root. The slightly worn M(3)exhibits at least four cusps with a large hypocone, three roots with large radicular plates, and an absence of taurodontism. They are moderately large for later Middle Pleistocene humans in their buccolingual crown diameters, although the M(3)mesiodistal diameter is modest. The C(1)exhibits labial calculus and multiple linear hypoplastic defects, but the M(3)is lesion free. Both teeth are morphologically similar to those of other Middle Pleistocene European humans and reinforce a pattern of dental hypertrophy among these archaic Homo.     

Discontinuity of Human Presence at Atapuerca during the Early Middle Pleistocene: A Matter of Ecological Competition?

Increasing evidence suggests that the European human settlement is older than 1.2 Ma. However, there is a fierce debate about the continuity or discontinuity of the early human settlement of Europe. In particular, evidence of human presence in the interval 0.7−0.5 Ma is scarce in comparison with evidence for the previous and later periods. Here, we present a case study in which the environmental conditions at Sierra de Atapuerca in the early Middle Pleistocene, a period without evidence of human presence, are compared with the conditions in the previous period, for which a relatively intense human occupation is documented. With this objective in mind, the available resources for a human population and the intensity of competition between secondary consumers during the two periods are compared using a mathematical model. The Gran Dolina site TD8 level, dated to 0.7−0.6 Ma, is taken as representative of the period during which Atapuerca was apparently not occupied by humans. Conditions at TD8 are compared with those of the previous period, represented by the TD6-2 level, which has yielded abundant evidence of intense human occupation. The results show that survival opportunities for a hypothetical human population were lower at TD8 than they were at TD6-2. Increased resource competition between secondary consumers arises as a possible explanation for the absence of human occupation at Atapuerca in the early Middle Pleistocene.     

Updated Atapuerca biostratigraphy: Small-mammal distribution and its implications for the biochronology of the Quaternary in Spain

The Sierra de Atapuerca (Burgos, Spain) contains the most comprehensive archaeological and paleontological evidence of human evolution in Europe during the Quaternary. The time scale based on the microfossil content of the sedimentary infillings of the caves of Atapuerca has been used to attribute relative ages to the various paleontological and archaeological levels and correlate amongst the different sites. Microfossils are particularly significant as they are the dominant constituents of the caves from the Pleistocene to the Holocene of the Sierra de Atapuerca. The microfossils of the Atapuerca sites are mainly composed of isolated bones and teeth from small vertebrates, and the hard tissue of the vertebrate skeleton can be taxonomically classified. In revisiting the distribution and the classification of the small-mammal taxa along five of the main stratigraphic sequences from Atapuerca (Sima del Elefante, Gran Dolina, Galería-Zarpazos, Sima de los Huesos, and Portalón), we apply a new method, the “Unitary Associations” method, in order to refine the existing biozones, adding five new Faunal Units, and interpret their relative age. Finally, we correlate Atapuerca with other Pleistocene sites of Europe.     

Regional variation in tooth size and pathology in fossil hominids.

Tooth size and dental pathology in fossil hominids were studied to test for regional differences in these parameters. The results showed little regional variation in tooth size for the Middle and Upper Pleistocene sites compared (except for Krapina) but considerable differences in the severity of attrition and dental pathology. These differences were considered indicative of regional differences in the functional load borne by the teeth, and in view of the similar technological status of the groups studied, were attributed to environmental differences in the diet. Since, in all regions, reduction in tooth size appeared to continue at the same rate for the periods investigated, no association can be established between presumed selective pressures related to differences in functional demands made on the dentition, and tooth reduction.