The TD6.3 faunal assemblage of the Gran Dolina site (Atapuerca,Spain): a late Early Pleistocene hyena den

Ethological studies have shown that besides human groups, large-medium carnivores have bone-collecting habits. The research developed since the last half of the twentieth century has attempted to characterise the carnivore’s accumulations and to identify them in the archaeo-paleontological record. At present, we have diagnostic criteria that define the accumulations produced by hyenids (mainly, Crocuta crocuta), thereby allowing us to differentiate them from the other accumulating agents. The faunal assemblage recovered at the Early Pleistocene TD6.3 layer of the Gran Dolina site is characterised by the presence of typical elements described in hyena dens: presence of small carnivores remains, high bone breakage, low epiphysis survival and a high frequency (>30%) of specimens with carnivore induced modifications, including large amounts of digested bones. However, attritional mortality profiles, hyena’s cubs remains, mid-shaft bone cylinders or differential anatomical composition among different weight-sizes, have not been observed or are ambiguous. In addition, anthropic evidences are scarce and concentrated in the uppermost section of the layer. TD6.3 is the result of an accumulation produced by hyenas using the cave as a den, in alternation with sporadic occupations by hominin groups. TD6.3 shows that hyena fossil accumulations may present great anatomical and taxonomic variability.     

Size variation in early human mandibles and molars from Klasies River,South Africa: Comparison with other middle and late Pleistocene assemblages and with modern humans

Previous studies of the Middle Stone Age human remains from Klasies River have concluded that they exhibited more sexual dimorphism than extant populations, but these claims have not been assessed statistically. We evaluate these claims by comparing size variation in the best‐represented elements at the site, namely the mandibular corpora and M₂s, to that in samples from three recent human populations using resampling methods. We also examine size variation in these same elements from seven additional middle and late Pleistocene sites: Skhūl, Dolní Věstonice, Sima de los Huesos, Arago, Krapina, Shanidar, and Vindija. Our results demonstrate that size variation in the Klasies assemblage was greater than in recent humans, consistent with arguments that the Klasies people were more dimorphic than living humans. Variation in the Skhūl, Dolní Věstonice, and Sima de los Huesos mandibular samples is also higher than in the recent human samples, indicating that the Klasies sample was not unusual among middle and late Pleistocene hominins. In contrast, the Neandertal samples (Krapina, Shanidar, and Vindija) do not evince relatively high mandibular and molar variation, which may indicate that the level of dimorphism in Neandertals was similar to that observed in extant humans. These results suggest that the reduced levels of dimorphism in Neandertals and living humans may have developed independently, though larger fossil samples are needed to test this hypothesis. Am J Phys Anthropol, 2009. © 2009 Wiley‐Liss, Inc.     

The endocast from Dana Aoule North (DAN5/P1): A 1.5 million year-old human braincase from Gona,Afar, Ethiopia

The nearly complete cranium DAN5/P1 was found at Gona (Afar, Ethiopia), dated to 1.5–1.6 Ma, and assigned to the species Homo erectus. Its size is, nonetheless, particularly small for the known range of variation of this taxon, and the cranial capacity has been estimated as 598 cc. In this study, we analyzed a reconstruction of its endocranial cast, to investigate its paleoneurological features. The main anatomical traits of the endocast were described, and its morphology was compared with other fossil and modern human samples. The endocast shows most of the traits associated with less encephalized human taxa, like narrow frontal lobes and a simple meningeal vascular network with posterior parietal branches. The parietal region is relatively tall and rounded, although not especially large. Based on our set of measures, the general endocranial proportions are within the range of fossils included in the species Homo habilis or in the genus Australopithecus. Similarities with the genus Homo include a more posterior position of the frontal lobe relative to the cranial bones, and the general endocranial length and width when size is taken into account. This new specimen extends the known brain size variability of Homo ergaster/erectus, while suggesting that differences in gross brain proportions among early human species, or even between early humans and australopiths, were absent or subtle.     

Brain size and encephalization in early to Mid-Pleistocene Homo

Important changes in the brain have occurred during the course of human evolution. Both absolute and relative size increases can be documented for species of Homo, culminating in the appearance of modern humans. One species that is particularly well-represented by fossil crania is Homo erectus. The mean capacity for 30 individuals is 973 cm(3). Within this group there is substantial variation, but brain size increases slightly in specimens from later time periods. Other Middle Pleistocene crania differ from those of Homo erectus. Characters of the facial skeleton, vault, and cranial base suggest that fossils from sites such as Arago Cave in France, the Sima de los Huesos in Spain, Bodo in Ethiopia, Broken Hill in Zambia, and perhaps Dali in China belong to the taxon Homo heidelbergensis. Ten of these mid-Quaternary hominins have brains averaging 1,206 cm(3) in volume, and many fall beyond the limits of size predicted for Homo erectus of equivalent age. When orbit height is used to construct an index of relative brain size, it is apparent that the (significant) increase in volume documented for the Middle Pleistocene individuals is not simply a consequence of larger body mass. Encephalization quotient values confirm this finding. These changes in absolute and relative brain size can be taken as further corroborative evidence for a speciation event, in which Homo erectus produced a daughter lineage. It is probable that Homo heidelbergensis originated in Africa or western Eurasia and then ranged widely across the Old World. Archaeological traces indicate that these populations differed in their technology and behavior from earlier hominins.     

The Atapuerca sites and the ibeas hominids

The Atapuerca railway Trench and Ibeas sites near Burgos, Spain, are cave fillings that include a series of deposits ranging from below the Matuyama/Bruhnes reversal up to the end of Middle Pleistocene. The lowest fossil-bearing bed in the Trench contains an assemblage of large and small Mammals includingMimomys savini, Pitymys gregaloides, Pliomys episcopalis, Crocuta crocuta, Dama sp. and Megacerini; the uppermost assemblage includesCanis lupus, Lynx spelaea, Panthera (Leo) fossilis, Felis sylvestris, Equus caballus steinbeimensis, E.c. germanicus, Pitymys subterraneus, Microtus arvalis agrestis, Pliomys lenki, and alsoPanthera toscana, Dicerorbinus hemitoechus, Bison schoetensacki, which are equally present in the lowest level. The biostratigraphic correlation and dates of the sites are briefly discussed, as are the paleoclimatic interpretation of the Trench sequences. Stone artifacts are found in several layers; the earliest occurrences correspond to the upper beds containingMimomys savini. A set of preserved human occupation floors has been excavated in the top fossil-bearing beds. The stone-tool assemblages of the upper levels are of upper-medial Acheulean to Charentian tradition. The rich bone breccia SH, in the Cueva Mayor-Cueva del Silo, Ibeas de Juarros, is a derived deposit, due to a mud flow that dispersed and carried the skeletons of many carnivores and humans. The taxa represented are:Ursus deningeri (largely dominant),Panthera (Leo) fossilis, Vulpes vulpes, Homo sapiens var. Several traits of both mandibular and cranial remains are summarized. Preliminary attempts at dating suggest that the Ibeas fossil man is older than the Last Interglacial, or oxygen-isotope stage 5.     

On the taxonomic affinities of the Dmanisi mandible (Georgia)

The recent discovery of unexpectedly ancient human remains has fuelled interest about the first dispersion of Homo outside Africa. The Dmanisi mandible is perhaps one of the most interesting findings, as it supposedly represents one of the oldest hominids outside of Africa. Recently, different interpretations have been published about this specimen. Our comparison of the Dmanisi mandible with a large sample of mandibles and teeth has led us to a new interpretation. In our view, the Dmanisi mandible exhibits a unique combination of traits. Some of its features, taken in isolation, may be attributed to morphological extremes within the genus Homo. The architecture of the mandible as well as the morphology and dimensions of incisors, canines, and P3s are clearly primitive. However, dental traits such as the reduction of the talonid in the P4s and a distally decreasing molar series seems to be derived. Some combinations of these traits are found in specimens of Homo ergaster and differ from those generally present in later hominids. Thus, we propose that the Dmanisi mandible might be taxonomically classified as Homo sp. indet. (aff. ergaster). Furthermore, some aspects of the dentition in Dmanisi display close similarities to Asian Homo erectus. If the 1.8–1.6 Myr dating for the Dmanisi mandible is correct, the differentiation of the Asian branch of the genus Homo could be regarded as a very ancient event. Am J Phys Anthropol 107:145–162, 1998. © 1998 Wiley-Liss, Inc.     

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。     

The Omo-Turkana Basin fossil hominins and their contribution to our understanding of human evolution in Africa

The Omo-Turkana Basin, including the hominin fossil sites around Lake Turkana and the sites along the lower reaches of the Omo River, has made and continues to make an important contribution to improving our murky understanding of human evolution. This review highlights the various ways the Omo-Turkana Basin fossil record has contributed to, and continues to challenge, interpretations of human evolution. Despite many diagrams that look suspiciously like comprehensive hypotheses about human evolutionary history, any sensible paleoanthropologist knows that the early hominin fossil record is too meager to do anything other than offer very provisional statements about hominin taxonomy and phylogeny. If history tells us anything, it is that we still have much to learn about the hominin clade. Thus, we summarize the current state of knowledge of the hominin species represented at the Omo-Turkana Basin sites. We then focus on three specific topics for which the fossil evidence is especially relevant: the origin and nature of Paranthropus; the origin and nature of early Homo; and the ongoing debate about whether the pattern of human evolution is more consistent with speciation by cladogenesis, with greater taxonomic diversity or with speciation by anagenetic transformation, resulting in less taxonomic diversity and a more linear interpretation of human evolutionary history.     

The femur in early human evolution.

Uni- and multivariate analyses of 5 fossil and 215 extant hominoid femora show that two morphological patterns of hominid femora existed about two million years ago. Femora classified as Homo sp. indet. (KNMER 1472 and 1481) are more like Homo sapiens although not identical.Those classified as Australopithecus robustus (SK 82 and 97) and A. boisei (KNM-ER 1503) are similar to one another but uniquely different from any living hominoid. The strong mophological constrasts imply biomechanical and possible locomotor differences, although these are as yet unknown.     

Radiocarbon dating the appearance of modern humans and timing of cultural innovations in Europe: new results and new challenges

New radiocarbon dates from the sites of Bockstein-T?rle, Geissenkl?sterle, Hohle Fels, Hohlenstein-Stadel, Sirgenstein, and Vogelherd in the Swabian Jura of southwestern Germany indicate that the Aurignacian of the region spans the period from ca. 40-30ka BP. If the situation at Vogelherd, in which skeletal remains from modern humans underlie an entire Aurignacian sequence, is viewed as representative for the region, the dates from the Swabian Jura support the hypothesis that populations of modern humans entered the region by way of the "Danube Corridor." The lithic technology from the lower Aurignacian of Geissenkl?sterle III is fully developed, and classic Aurignacian forms are well represented. During the course of the Aurignacian, numerous assemblages rich in art works, jewelry, and musical instruments are documented. By no later than 29ka BP the Gravettian was well established in the region. These dates are consistent with the "Kulturpumpe" hypothesis that important cultural innovations of the Aurignacian and Gravettian in Swabia predate similar developments in other regions of Europe. The radiocarbon dates from Geissenkl?sterle corroborate observations from other non-archaeological data sets indicating large global fluctuations in the atmospheric concentrations of radiocarbon between 30 and 50ka calendar years ago. These fluctuations lead to complications in building reliable chronologies during this period and cause the "Middle Paleolithic Dating Anomaly" and the "Coexistence Effect," which tend to exaggerate the temporal overlap between Neanderthals and modern humans.     

Krapina, “Classic” Neanderthals,and the evolution of the European face

Except for the front end of the dental arch, tooth size remained at approximately the same level throughout the Middle Pleistocene. The Krapina Neanderthals at the end of the last interglacial differed from Homo erectus only in having larger front teeth. From that time on, tooth size in populations at the northern edge of the area of human occupation in the Old World has reduced approximately in proportion to the time elapsed. The “Classic” Neanderthals of western Europe, in fact, have teeth that are 15% smaller than those of the earlier Krapina Neanderthals and only 5% larger than the early Upper Palaeolithic. Reduction since the early Upper Palaeolithic has proceeded another full 20%. It is suggested that the development of heated stone cooking in the Mousterian, originally for the purpose of thawing frozen food, reduced the forces of selection that had previously maintained tooth size during the Middle Pleistocene. The operation of the Probable Mutation Effect, then produced the observed reductions.     

Hybridization in human evolution: Insights from other organisms

During the late Pleistocene, isolated lineages of hominins exchanged genes thus influencing genomic variation in humans in both the past and present. However, the dynamics of this genetic exchange and associated phenotypic consequences through time remain poorly understood. Gene exchange across divergent lineages can result in myriad outcomes arising from these dynamics and the environmental conditions under which it occurs. Here we draw from our collective research across various organisms, illustrating some of the ways in which gene exchange can structure genomic/phenotypic diversity within/among species. We present a range of examples relevant to questions about the evolution of hominins. These examples are not meant to be exhaustive, but rather illustrative of the diverse evolutionary causes/consequences of hybridization, highlighting potential drivers of human evolution in the context of hybridization including: influences on adaptive evolution, climate change, developmental systems, sex‐differences in behavior, Haldane's rule and the large X‐effect, and transgressive phenotypic variation.