Cranial remains of Middle Pleistocene European hominids

Cranial remains of the Middle Pleistocene European hominids are reviewed in a historical context. Swanscombe and Steinheim, the earliest specimens to be discovered, are compared with each other. Following this, the more recently discovered specimens from Vértesszöllös, Petralona and Bilzingsleben are compared. Finally, Riss remains from La Chaise, Biache and Arago are discussed in the context provided by the above specimens. It is suggested that while phyletic evolution may play a role in the observed differences, a marked degree of sexual dimorphism contributes to the variation expressed by these specimens.     

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.     

Senescence in European Middle Pleistocene hominids: The Atapuerca evidence

The mandibular specimens AT-250 and AT-793 (individual IV) and AT-888 (individual XXI) from the Atapuerca Middle Pleistocene sample show a combination of traits interpreted as evidence of senescence. In order to explore age-related mandibular changes relevant to the interpretation of the Atapuerca hominids, the sample of known age and sex from Spitalfields (London) and the Neolithic sample from Abu Hureyra (Syria) were used as a baseline. Results obtained indicate that the occurrence of an enlarged and posteriorly located mental foramen, in association with high alveolar resorption in AT-250 and AT-888, support the hypothesis of senescence in individuals IV and XXI from the Atapuerca sample. The findings suggest that European Middle Pleistocene populations lived long enough to reach senescence. The age at death for Middle Pleistocene hominids and Neanderthals has previously been estimated at around 40–45 years in the oldest specimens. The appearance of senescence processes before this age is indicative of higher rates of morphological aging in the clade which gave rise to Neanderthals. The anatomical significance of ageing features when considered in the context of craniofacial remodelling can help in the understanding of the taxonomic status of these morphological traits.     

Middle Pleistocene hominids from Olduvai Gorge, northern Tanzania

Cranial, dental, and mandibular remains of eight Olduvai hominids are described in detail. Four individuals were recovered in situ in Beds II to IV, while three more are most probably derived from Bed IV, the Masek Beds and the Lower Ndutu Beds. One specimen is of uncertain provenance. Deposits from which the fossils were collected range from late Lower Pleistocene to Middle Pleistocene in age. Of particular interest are three fragmentary lower jaws, which can be compared to mandibles of Homo erectus known from localities in Northwest Africa and China. Olduvai hominid 22, a nearly complete half mandible with crowns of P3-M2 in place, shares many anatomical features with fossils from Ternifine and Choukoutien. This individual is also similar to a jaw from the Kapthurin Formation west of Lake Baringo, Kenya. How best to interpret these comparisons is not clear, but in view of marked similarities between specimens representing geographically diverse populations from different time periods, it may be unwise to rely on mandibular evidence alone to document the presence of regional lineages. Gradual change and continuity within a sequence of Northwest African Homo fossils has been endorsed by many workers, but such hypotheses cannot be tested adequately with the fragmentary jaws available.     

Enamel hypoplasia in the Middle Pleistocene hominids from Atapuerca (Spain)

The prevalence and chronology of enamel hypoplasias were studied in a hominid dental sample from the Sima de los Huesos (SH) Middle Pleistocene site at the Sierra de Atapuerca (Burgos, northern Spain). A total of 89 permanent maxillary teeth, 143 permanent mandibular teeth, and one deciduous lower canine, belonging to a minimum of 29 individuals, were examined. Excluding the antimeres (16 maxillary and 37 mandibular cases) from the sample, the prevalence of hypoplasias in the permanent dentition is 12.8% (23/179), whereas the deciduous tooth also showed an enamel defect. No statistically significant differences were found between both arcades and between the anterior and postcanine teeth for the prevalence of hypoplasias. In both the maxilla and the mandible the highest frequency of enamel hypoplasias was recorded in the canines. Only one tooth (a permanent upper canine) showed two different enamel defects, and most of the hypoplasias were expressed as faint linear horizontal defects. Taking into account the limitations that the incompleteness of virtually all permanent dentitions imposes, we have estimated that the frequency by individual in the SH hominid sample was not greater than 40%. Most of the hypoplasias occurred between birth and 7 years (N = 18, X = 3.5, SD = 1.3). Both the prevalence and severity of the hypoplasias of the SH hominid sample are significantly less than those of a large Neandertal sample. Furthermore, prehistoric hunter-gatherers and historic agricultural and industrial populations exhibit a prevalence of hypoplasias generally higher than that of the SH hominids. Implications for the survival strategies and life quality of the SH hominids are also discussed. © 1995 Wiley-Liss, Inc.     

Probit analysis of Upper Pleistocene hominids

Probit analysis allows the testing of the normality of a sample population where curve fitting and differential weighting occur simultaneously. Probit analysis may be utilized with small samples, and is ideal for use in the study of fossil populations. It is assumed that population which are normally distributed are likely to be in genetic equilibrium, so that graphic and computational probit analysis may be used to discern the effects of evolutionary forces on human populations. Probit analysis of cranial length in a sequence of Athenian population is presented in order to verify its sensitivity to these evolutionary forces. Probit analysis and other tests of homogeneity are applied to similar variables of Upper Pleistocene European Neandertal and modern man.Despite significant differences between variable means of Neandertal and modern man, probit analyses of their combined populations showed no less deviation from normality than either group separately. This suggests that the evolutionary changes in the cranium from the Neandertal stage to the modern stage of man were effected without significantly disrupting the genetic equilibrium of these populations, i.e. that rapid changes in these populations are unlikely to have occurred.     

Ontogenetic approach to variation in Middle Pleistocene hominids. Evidence from the Atapuerca-SH mandibles

The ontogenetic processes underlying the variation detected in the European Middle Pleistocene hominids are explored. Regulation of growth parameters of the ontogenetic trajectory of these hominid is discussed in the context of heterochrony schemes. The sample of mandibles coming from the Atauuerca-SH site is used as a study case. The variability shown by these mandibles can be arranged along a morphological gradient, and study of growth directions has shown that the remodelling patterns are in line with the existence of a single morphogenetic pattern controlling the process. In addition, the spectrum of morphology detected in the Atapuerca sample virtually encompasses the entire range of variability detected in European Middle Pleistocene samples. From a morphological viewpoint, a hypermorphic pattern seems to be the responsible for the individual variation detected in the sample. Yet, when this information is related to mandibular growth and craniofacial interactions, evidence of an interindividual gradient of somatic development for the Atapuerca hominids is provided. Several hypotheses concerning growth parameters, however, may account for this gradient. Time and rate hypermorphosis hypothesis are discussed. Coincidence of the gradient of variation detected in this sample and the general mandibular growth trajectory of living species, support the hypothesis that is “time of maturation” of the organism the key factor.     

The absolute dating of Upper Pleistocene subSaharan fossil hominids and their place in human evolution

In the evolution of anatomically modern man and his subspecies most specialists have concentrated on investigating geographical areas other than Africa as the possible area of origin.In this study 20 fossil hominids and associated faunal remains from South and East Africa were dated by microanalysis, radiocarbon, and amino-acid dating in order to see whether modern man appears later, was sympatric, or even predated Neandertal man.These dates indicate that anatomically modern man occurs sympatrically and possibly even predates the Rhodesian group of Neandertals in Africa. Modern man might also be contemporary to and possibly even predate the occurrence of Neandertal in Europe.This would indicate that modern man did not evolve from but possibly gave rise to the Neandertals as off-shoots.Two possibilities for the evolution of modern man are suggested. First, that Homo sapiens capensis evolved about 90,000 to 100,000 years ago from possibly Homo erectus by way of a “basic” Homo sapiens and later gave rise to Homo sapiens rhodesiensis, Homo sapiens afer, and possibly Homo sapiens palestinus around 50,000 years ago with Homo neanderthalensis and Homo sapiens capensis evolving separately from Homo erectus. In this case Homo neanderthalensis would be a different species from Homo sapiens which includes Homo sapiens capensis, Homo sapiens rhodesiensis, Homo sapiens afer, and possibly Homo sapiens palestinus.Secondly, Homo sapiens capensis evolved by way of a “basic” Homo sapiens with Homo sapiens rhodesiensis and Homo sapiens palestinus branching off from Homo sapiens capensis around 50,000 years ago. Before that, around 90,000 to 100,000 years ago Homo sapiens capensis evolved first and was then followed by Homo sapiens neanderthalensis from a “basic” Homo sapiens stock, but diverged. This means, all Neandertals, Homo sapiens capensis, Homo sapiens sapiens and Homo sapiens afer can be considered as subspecies of Homo sapiens.The author favors the first scheme since on relative dating grounds the existence of Neandertal man in Europe before the earliest date of Homo sapiens capensis and a “basic” Homo sapiens seems to be fairly well documented. Irrespective of either one of these possibilities, modern man evolved in Africa and seems to have migrated into Europe and other parts of the world.New absolute dating techniques are mentioned in detail like the new radiocarbon-collagen method and amino acid dating.     

Dynamic bone remodeling in later Pleistocene fossil hominids

Histomorphometric analysis of femoral and tibial diaphyseal fragments from seven Late Archaic and three Early Modern humans are compared with those of the Pecos, a pre-Columbian Native American population. The ten samples, from Broken Hill (EM-793), Shanidar 2, 3, 4, 5, and 6, Tabun 1, and Skhul 3, 6, and 7, provide age-at-death results consistent with earlier estimates for most individuals. The Pleistocene groups exhibit less bone turnover and smaller osteons than Recent populations. Resorption and formation were both coupled and balanced in these Pleistocene populations, but the overall vigor of individual cells from both the osteoclast and osteoblast cell lines was less than in Recent populations. Thus the greater bone mass in Later Pleistocene members of the genus Homo is not the result of higher levels of bone turnover, at least among adults. © 1996 Wiley-Liss, Inc.     

Natural selection in brain evolution of early hominids

Directional effect of natural selection on the arrangement of brain of anthropoids and man is reviewed. It is demonstrated that the evolution of the human nervous system is an integrated result of several multidirectional processes. At the early stages of the evolution of primates, the general biological principles of survival of the fittest, i.e., natural selection of the most adapted variants of the brain structure prevailed. During the period of hominid specialization, natural selection led to the formation of the neocortical control of voluntary movements, memory, and mental associations. At later stages of human morphological evolution, biological mechanisms of natural selection of the brain arrangement were replaced by social mechanisms. This process initiated hominid migrations and the growth of the brain size and individual variability in human ancestors. A model of cerebral sorting is proposed to explain the mechanisms of multidirectional selection leading to an increase in brain size of early hominids.     

Some comments on the case for Early Pleistocene hominids in South-Eastern Spain

Several sites in the Orce Basin have revealed evidence of the presence of hominids in the Early Pleistocene. These remains are dated to over 1.0 million years, while they may be as old as 1.6 million years. The skeletal remains from Venta Micena in the Orce Basin show a molecular “fossil protein” pattern which aligns them with hominids, but not with equids. This is supported by the anatomical evidence of the two humeral shafts from theEstrato Blanco in the Venta Micena deposits. The biparieto-occipital partial calvaria shows some unusual features if VM-0 is a hominid specimen. the presence of a prominent crest on the internal surface of the occipital fragment adjacent to the point lambda is decidedly unusual for a modern human calvaria. Moreover, theimpressions gyrorum, in the region where the superior parietal lobule of the cerebral hemisphere abutted against the calvaria, point to a bipartite superior parietal lobule with anterior and posterior moieties which, on the endocast, are clearly separated by a depression that represents a sulcus. These morphological traits are rather puzzling if VM-0 is a hominid, and at first they led me to hesitate over the anatomical identification of VM-0. However, the studies of Campillo (1989) and of Campillo and Barcelo (1986) suggest that the features of the fragment VM-0 are compatible with those of a hominid. Because I believe that we do not possess sufficient information on the variability of the endocranial and ectocranial manifestations of the sagittal suture and of its variance with age of the individual, in different hominid species and different equid species, I have not adduced this pattern as evidence in support or rebuttal of the hominid status of VM-0.     

Evolution of the supraorbital region in Upper Pleistocene fossil hominids from South-Central Europe

South-Central European fossil hominids dated to the Upper Pleistocene exhibit a distinct morphological and metric continuum in supraorbital form from early Neandertal (Krapina), through late Neandertals (Vindija), to early Upper Paleolithic hominids. The supraorbital morphologies pertinent to this continuum are documented, and the alterations in size and morphology are discussed ralative to the function of supraorbital superstructures and their relationship to overall craniofacial form. It is concluded that this continuum most likely reflects localized transition between Neandertals and modern man in this region of Europe.     

Relative dental development of Upper Pleistocene hominids compared to human population variation

The relative development of permanent teeth in samples of Neandertal/archaic Homo and Early Modern/Upper Paleolithic hominids is compared to the range of variability found in three recent human samples. Both fossil hominid samples are advanced in relative M2 and M3 development compared to white French-Canadians, but only the Neandertal/archaic Homo M3 sample is advanced when compared to black southern Africans. Both fossil hominid samples are delayed in relative I1 and P3 development compared to the recent human samples. Two hypotheses concerning the significance of the advanced M3 and M2 development found in both hominid groups and southern Africans compared to French-Canadians are discussed. The first postulates that the differences in relative molar development are due simply to variation in tooth/jaw size relationships. The second postulates that the relatively advanced M3 and M2 development found in the fossil hominids and southern Africans is a correlate of their potential for advanced skeletal maturation compared to French-Canadians and other European-derived populations. It appears that dental development patterns have continued to evolve from the Upper Pleistocene to present times, and that Neandertals and Early Moderns shared similar patterns of relative dental development. © 1996 Wiley-Liss, Inc.     

Further analysis of mandibular molar crown and cusp areas in Pliocene and early Pleistocene hominids

Crown and cusp areas of mandibular molars were measured and analyzed on a sample of 249 specimens attributed to Australopithecus afarensis, A. africanus, A. (Paranthropus) robustus, A. (P.) boisei, and early Homo. In addition to intertaxon comparisons, we compared data that had been collected independently by two of the authors using methods that differ slightly in technique of measurement. Interobserver differences were evaluated by the t-test of paired comparisons, method error statistic, percent differences, and principal component analysis. Results suggest that between-technique error of measurement of overall crown area is small. Error estimates for individual cusp area measurements were of larger relative magnitude. However, these were not sufficient to detract from the conclusions derived from comparative analyses. Our results are in general agreement with previous assessments of early hominid dental size. Crown areas of A. africanus, however, exhibit a mosaic pattern, with M1 similar in size to that of A. afarensis and early Homo, and M2 and M3 similar in size to that of A. robustus. Intertaxon comparisons of relative cusp area were undertaken by univariate statistics and principal component analysis. These analyses revealed that while A. (P.) robustus and A. (P.) boisei both possess mandibular molars with cusp proportions significantly different from the ‘non-robust’ taxa, these differences are substantially greater in A. (P.) boisei. © 1994 Wiley-Liss, Inc.     

早-中更新世中国古人类演化及其与气候环境的关系

近一个世纪以来,随着旧石器时代考古与古人类研究的发展,我们已经获得了大量的研究材料和数据。尽管个别遗址的年代、文化性质和化石人类分析及鉴定仍有争议,但如此长时间的积累依然为构建序列性认识和思考提供了基础。目前的研究多集中在单个遗址或者区域的古人类化石、石器打制技术和气候环境背景的个案分析,对早-中更新世中国古人类演化及其与气候环境的关系尚缺乏系统探讨。本文选取过去百万年里地球气候冰期-间冰期旋回经历的两次显著转型事件:中更新世气候转型期和中布容事件作为时间节点,讨论伴随气候转型时期中国北方地区、秦岭及周边地区和中国南方地区出现的早期古人类演化特征的变化。具体从遗址分布、石器技术变化、早期人类化石特征等方面入手,尝试构建人类活动与气候环境变化的耦合关系。     

更新世中期中国古人类演化区域连续性与多样性的化石证据

以往,在东亚大陆发现的更新世中期人类化石被分别归入直立人和古老型智人。这种分类的主要依据是化石形态特征以及年代。魏敦瑞对周口店第一地点人类化石研究描述的一些头骨、下颌骨和牙齿特征通常被作为判定直立人的标准。根据这些化石的年代分布,一般将30万年前的中更新世晚期作为划分直立人与古老型智人的大致年代界限。近20年来,在非洲、欧洲和东亚新发现了一些更新世中期人类化石,目前古人类学界对中国更新世中期人类化石特征及演化有了与以往不同的认识。最近对大荔、许家窑、盘县大洞、许昌、华龙洞等人类化石的研究显示,近30万年以来东亚大陆人类演化呈现复杂的多样性,将这一时期人类全部归入古老型智人难以准确反映更新世中期中国古人类演化模式及规律。本文结合近年中国更新世中期人类演化研究进展,选择部分具有演化及分类价值的形态特征,分析这些特征在更新世中期中国古人类化石的表现特点。在此基础上,对更新世中期中国古人类演化模式做了尝试性探讨。本研究发现,周口店、和县、沂源、南京等中更新世早期人类化石呈现有较多的区域性特征,形态特征表现相对稳定;而大荔、金牛山、许家窑、许昌、华龙洞、马坝、盘县大洞等中更新世晚期人类在化石形态特征表现复杂多样,变异范围大。此外,在这一时期人类化石上发现较多与生存活动、健康、环境适应有关的证据。根据这些发现,作者认为中国中更新世早期组人类演化以形态连续性为主;进入中更新世晚期,中国古人类演化区域性特征减弱,演化模式以多样性为主。一系列新的化石发现和研究证据提示中更新世晚期东亚大陆可能生存有不同的古人类成员。根据目前掌握的化石形态和年代证据,大约30万年前是中国更新世中期演化变化关键时间节点。     

Patterns of variation in Microtus arvalis and Microtus agrestis populations from Middle to Late Pleistocene in southwestern Europe

Fifteen paired fossil populations of Microtus arvalis and Microtus agrestis from southwestern Europe have been analysed from a morphological and morphometric point of view. The sites under consideration are located in the northern Iberian Peninsula and southern France, from the Middle Pleistocene to the end of the Late Pleistocene. The aim of this study is to stress once again the importance of keeping these two species separated in the fossil record in order to recognize specific trends of evolution and divergence and to obtain more precise information on palaeoclimatic and palaeoenvironmental conditions. It was possible to observe remarkable intraspecific differences between Middle and Late Pleistocene populations of both species. Furthermore, in synchronic co-specific populations from the Late Pleistocene, climatic and geographic conditions seem to exert a major influence in shaping intraspecific changes in dental pattern.     

Lethal Interpersonal Violence in the Middle Pleistocene

Evidence of interpersonal violence has been documented previously in Pleistocene members of the genus Homo, but only very rarely has this been posited as the possible manner of death. Here we report the earliest evidence of lethal interpersonal violence in the hominin fossil record. Cranium 17 recovered from the Sima de los Huesos Middle Pleistocene site shows two clear perimortem depression fractures on the frontal bone, interpreted as being produced by two episodes of localized blunt force trauma. The type of injuries, their location, the strong similarity of the fractures in shape and size, and the different orientations and implied trajectories of the two fractures suggest they were produced with the same object in face-to-face interpersonal conflict. Given that either of the two traumatic events was likely lethal, the presence of multiple blows implies an intention to kill. This finding shows that the lethal interpersonal violence is an ancient human behavior and has important implications for the accumulation of bodies at the site, supporting an anthropic origin.