Pleistocene Chinese cave hyenas and the recent Eurasian history of the spotted hyena,Crocuta crocuta

The living hyena species (spotted, brown, striped and aardwolf) are remnants of a formerly diverse group of more than 80 fossil species, which peaked in diversity in the Late Miocene (about 7–8 Ma). The fossil history indicates an African origin, and morphological and ancient DNA data have confirmed that living spotted hyenas (Crocuta crocuta) of Africa were closely related to extinct Late Pleistocene cave hyenas from Europe and Asia. The current model used to explain the origins of Eurasian cave hyena populations invokes multiple migrations out of Africa between 3.5–0.35 Ma. We used mitochondrial DNA sequences from radiocarbon‐dated Chinese Pleistocene hyena specimens to examine the origin of Asian populations, and temporally calibrate the evolutionary history of spotted hyenas. Our results support a far more recent evolutionary timescale (430–163 kya) and suggest that extinct and living spotted hyena populations originated from a widespread Eurasian population in the Late Pleistocene, which was only subsequently restricted to Africa. We developed statistical tests of the contrasting population models and their fit to the fossil record. Coalescent simulations and Bayes Factor analysis support the new radiocarbon‐calibrated timescale and Eurasian origins model. The new Eurasian biogeographic scenario proposed for the hyena emphasizes the role of the vast steppe grasslands of Eurasia in contrast to models only involving Africa. The new methodology for combining genetic and geological data to test contrasting models of population history will be useful for a wide range of taxa where ancient and historic genetic data are available.     

Homo floresiensis Contextualized: A Geometric Morphometric Comparative Analysis of Fossil and Pathological Human Samples

The origin of hominins found on the remote Indonesian island of Flores remains highly contentious. These specimens may represent a new hominin species, Homo floresiensis, descended from a local population of Homo erectus or from an earlier (pre-H. erectus) migration of a small-bodied and small-brained hominin out of Africa. Alternatively, some workers suggest that some or all of the specimens recovered from Liang Bua are pathological members of a small-bodied modern human population. Pathological conditions proposed to explain their documented anatomical features include microcephaly, myxoedematous endemic hypothyroidism (“cretinism”) and Laron syndrome (primary growth hormone insensitivity). This study evaluates evolutionary and pathological hypotheses through comparative analysis of cranial morphology. Geometric morphometric analyses of landmark data show that the sole Flores cranium (LB1) is clearly distinct from healthy modern humans and from those exhibiting hypothyroidism and Laron syndrome. Modern human microcephalic specimens converge, to some extent, on crania of extinct species of Homo. However in the features that distinguish these two groups, LB1 consistently groups with fossil hominins and is most similar to H. erectus. Our study provides further support for recognizing the Flores hominins as a distinct species, H. floresiensis, whose affinities lie with archaic Homo.     

在中国云南发现狒狒化石

现存狒狒类(Papionin)生活于非洲(如Papio和Theropithoan)、亚洲(如Macaca)和北非(M.sylvanas)。在上新世和更新世,Theropithecus经历了从非洲到亚洲的扩散过程,在印度发现了类似化石。这次在云南中甸金沙江附近发现的下更新世狒狒化石(Papio)证明,如同亚洲猕猴和现代人类祖先一样,非洲狒狒类(Papio和Theropithecus)在同一时期从非洲扩散到亚洲。所不同的是它们没有像猕猴和人类一样生存下来。这次化石的发现对于研究以下生物学问题提供了重要依据1)探讨旧大陆猴类在上新—更新世从非洲到亚洲的扩散过程;2)研究不同旧大陆猴类的进化和环境适应性;3)为现代人类祖先在非洲—亚大陆的扩散研究提供证据;4)由于化石产地包括有人类祖先和其他动物的化石,因此,狒狒在亚洲的生态适应研究将为探讨人类在同一时期的生态适应提供证据。     

Tectonics, orbital forcing, global climate change, and human evolution in Africa: introduction to the African paleoclimate special volume

The late Cenozoic climate of Africa is a critical component for understanding human evolution. African climate is controlled by major tectonic changes, global climate transitions, and local variations in orbital forcing. We introduce the special African Paleoclimate Issue of the Journal of Human Evolution by providing a background for and synthesis of the latest work relating to the environmental context for human evolution. Records presented in this special issue suggest that the regional tectonics, appearance of C(4) plants in East Africa, and late Cenozoic global cooling combined to produce a long-term drying trend in East Africa. Of particular importance is the uplift associated with the East African Rift Valley formation, which altered wind flow patterns from a more zonal to more meridinal direction. Results in this volume suggest a marked difference in the climate history of southern and eastern Africa, though both are clearly influenced by the major global climate thresholds crossed in the last 3 million years. Papers in this volume present lake, speleothem, and marine paleoclimate records showing that the East African long-term drying trend is punctuated by episodes of short, alternating periods of extreme wetness and aridity. These periods of extreme climate variability are characterized by the precession-forced appearance and disappearance of large, deep lakes in the East African Rift Valley and paralleled by low and high wind-driven dust loads reaching the adjacent ocean basins. Dating of these records show that over the last 3 million years such periods only occur at the times of major global climatic transitions, such as the intensification of Northern Hemisphere Glaciation (2.7-2.5 Ma), intensification of the Walker Circulation (1.9-1.7 Ma), and the Mid-Pleistocene Revolution (1-0.7 Ma). Authors in this volume suggest this onset occurs as high latitude forcing in both Hemispheres compresses the Intertropical Convergence Zone so that East Africa becomes locally sensitive to precessional forcing, resulting in rapid shifts from wet to dry conditions. These periods of extreme climate variability may have provided a catalyst for evolutionary change and driven key speciation and dispersal events amongst mammals and hominins in Africa. In particular, hominin species seem to differentially originate and go extinct during periods of extreme climate variability. Results presented in this volume may represent the basis of a new theory of early human evolution in Africa.     

Reconstructing the Diet of an Extinct Hominin Taxon: The Role of Extant Primate Models

Modern humans represent the only surviving species of an otherwise extinct clade of primates, the hominins. As the closest living relatives to extinct hominins, extant primates are an important source of comparative information for the reconstruction of the diets of extinct hominins. Methods such as comparative and functional morphology, finite element analysis, dental wear, dental topographic analysis, and stable isotope biogeochemistry must be validated and tested within extant populations before they can be applied to extinct taxa. Here we review how these methods have and might be used to reconstruct the diet of a particular extinct hominin, Paranthropus boisei, which has no extant analogue for its highly derived masticatory morphology. Our review emphasizes the potential and limitations of using extant primates as models for the reconstruction of extinct hominin diets. We encourage paleoanthropologists and those who study the feeding behaviors of extant primates to work together to investigate and validate methods for interpreting the diets of all extinct primates, including hominins.     

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 hominoid proximal radius: re-interpreting locomotor behaviors in early hominins

Studies of fossil hominins are traditionally taxonomically narrow and often exclude comparisons with hylobatids. Hence, results of functional analyses of postcrania, interpreted as indicating that early hominins are "African-ape-like" in their postcranial skeletons and positional behaviors, may reflect an artifact of inadequate taxonomic and morphological breadth of the comparative sample. To address this problem and better understand early hominin positional behaviors, this study included hylobatids in a comparative analysis, focusing on the hominoid elbow joint. Specifically, morphometric variables of the proximal radius were derived from measurements from a sample of all genera of extant hominoids and casts of extinct hominin species. Univariate and multivariate analyses were performed on these data. Results show that early hominins are morphologically diverse and are not, as a group, similar to any one extant group. Instead, the fossils resemble Pan, Gorilla, and Hylobates, and are not like modern Homo sapiens or Pongo. This suggests that the morphology of Hylobates may reflect a morphotype for all later hominoids, thus complicating the functional interpretations of fossil hominins. The implications of these results are that the proximal radius is not a sensitive indicator of locomotor behavior among hominoids since the morphology in hylobatids and Gorilla and Pan is similar despite widely varying positional repertoires. Furthermore, inferences of function from form in extinct hominins can be drastically affected by the comparative outgroup selection. A re-evaluation of the functional morphology of the proximal radius in early hominins is addressed.     

Locomotor anatomy and biomechanics of the Dmanisi hominins

The Dmanisi hominins inhabited a northern temperate habitat in the southern Caucasus, approximately 1.8 million years ago. This is the oldest population of hominins known outside of Africa. Understanding the set of anatomical and behavioral traits that equipped this population to exploit their seasonal habitat successfully may shed light on the selection pressures shaping early members of the genus Homo and the ecological strategies that permitted the expansion of their range outside of the African subtropics. The abundant stone tools at the site, as well as taphonomic evidence for butchery, suggest that the Dmanisi hominins were active hunters or scavengers. In this study, we examine the locomotor mechanics of the Dmanisi hind limb to test the hypothesis that the inclusion of meat in the diet is associated with an increase in walking and running economy and endurance. Using comparative data from modern humans, chimpanzees, and gorillas, as well as other fossil hominins, we show that the Dmanisi hind limb was functionally similar to modern humans, with a longitudinal plantar arch, increased limb length, and human-like ankle morphology. Other aspects of the foot, specifically metatarsal morphology and tibial torsion, are less derived and similar to earlier hominins. These results are consistent with hypotheses linking hunting and scavenging to improved walking and running performance in early Homo. Primitive retentions in the Dmanisi foot suggest that locomotor evolution continued through the early Pleistocene.     

Postcranial remains and the origin of modern humans

The nature, timing, and location of the origin of modern humans has been the subject of intense controversy for the last 15 years.^1–4 Genetic data and new radiometric dates for key fossils that lie beyond the range of radiocarbon dating have substantially added to the knowledge derived from the fossil evidence documenting the transition from archaic to modern humans. These new data, however, have failed to resolve the problem in its entirety. Most authorities now accept that Africa played an important, and probably central, role in the origin of modern humans.^7–13 The genetic evidence seems to be particularly emphatic that an African population that existed between 200,000 and 100,000 years ago (100 ka) is ancestral to all living humans.^6,7 Controversy still surrounds the question of how much, if at all, archaic humans from outside of Africa, such as Neandertals, late archaic Chinese hominins such as Jinniushan, and the Indonesian Ngandong hominins, may have contributed to the morphological and genetic diversity present in living populations and the morphology of the earliest fossils of modern humans.¹⁰     

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.     

Look in the trees: Hylobatids as evolutionary models for extinct hominins

Studying extant apes is of central importance to paleoanthropology. This approach is informative in inferring how hominin skeletal morphology reflects phylogeny, behavior, development, and ecological context. Traditionally, great apes have dominated the paleoanthropological literature as extant analogs for extinct hominins, to the exclusion of their phylogenetic sister group, the hylobatids. Phylogenetic proximity, large body size, and high encephalization quotients may have contributed to decisions to use great apes as models for hominins. However, if we reexamine hylobatids as extant models for extinct hominins—using modern phylogenetic, behavioral, and ecological data—this clade is uniquely poised to inform future frameworks in paleoanthropology. The following features make hylobatids strong analogs for extinct hominins: taxonomic diversity, the timing of diversification, hybridization between species, small body size, and reduced sexual dimorphism. Based on these shared features, hylobatids offer future opportunities to paleoanthropology, and provide a much richer extant analog than is currently recognized.     

Genomic ancestry of North Africans supports back-to-Africa migrations

North African populations are distinct from sub-Saharan Africans based on cultural, linguistic, and phenotypic attributes; however, the time and the extent of genetic divergence between populations north and south of the Sahara remain poorly understood. Here, we interrogate the multilayered history of North Africa by characterizing the effect of hypothesized migrations from the Near East, Europe, and sub-Saharan Africa on current genetic diversity. We present dense, genome-wide SNP genotyping array data (730,000 sites) from seven North African populations, spanning from Egypt to Morocco, and one Spanish population. We identify a gradient of likely autochthonous Maghrebi ancestry that increases from east to west across northern Africa; this ancestry is likely derived from "back-to-Africa" gene flow more than 12,000 years ago (ya), prior to the Holocene. The indigenous North African ancestry is more frequent in populations with historical Berber ethnicity. In most North African populations we also see substantial shared ancestry with the Near East, and to a lesser extent sub-Saharan Africa and Europe. To estimate the time of migration from sub-Saharan populations into North Africa, we implement a maximum likelihood dating method based on the distribution of migrant tracts. In order to first identify migrant tracts, we assign local ancestry to haplotypes using a novel, principal component-based analysis of three ancestral populations. We estimate that a migration of western African origin into Morocco began about 40 generations ago (approximately 1,200 ya); a migration of individuals with Nilotic ancestry into Egypt occurred about 25 generations ago (approximately 750 ya). Our genomic data reveal an extraordinarily complex history of migrations, involving at least five ancestral populations, into North Africa.     

Early Back-to-Africa Migration into the Horn of Africa

Genetic studies have identified substantial non-African admixture in the Horn of Africa (HOA). In the most recent genomic studies, this non-African ancestry has been attributed to admixture with Middle Eastern populations during the last few thousand years. However, mitochondrial and Y chromosome data are suggestive of earlier episodes of admixture. To investigate this further, we generated new genome-wide SNP data for a Yemeni population sample and merged these new data with published genome-wide genetic data from the HOA and a broad selection of surrounding populations. We used multidimensional scaling and ADMIXTURE methods in an exploratory data analysis to develop hypotheses on admixture and population structure in HOA populations. These analyses suggested that there might be distinct, differentiated African and non-African ancestries in the HOA. After partitioning the SNP data into African and non-African origin chromosome segments, we found support for a distinct African (Ethiopic) ancestry and a distinct non-African (Ethio-Somali) ancestry in HOA populations. The African Ethiopic ancestry is tightly restricted to HOA populations and likely represents an autochthonous HOA population. The non-African ancestry in the HOA, which is primarily attributed to a novel Ethio-Somali inferred ancestry component, is significantly differentiated from all neighboring non-African ancestries in North Africa, the Levant, and Arabia. The Ethio-Somali ancestry is found in all admixed HOA ethnic groups, shows little inter-individual variance within these ethnic groups, is estimated to have diverged from all other non-African ancestries by at least 23 ka, and does not carry the unique Arabian lactase persistence allele that arose about 4 ka. Taking into account published mitochondrial, Y chromosome, paleoclimate, and archaeological data, we find that the time of the Ethio-Somali back-to-Africa migration is most likely pre-agricultural.     

Indian Siddis: African descendants with Indian admixture

The Siddis (Afro-Indians) are a tribal population whose members live in coastal Karnataka, Gujarat, and in some parts of Andhra Pradesh. Historical records indicate that the Portuguese brought the Siddis to India from Africa about 300-500 years ago; however, there is little information about their more precise ancestral origins. Here, we perform a genome-wide survey to understand the population history of the Siddis. Using hundreds of thousands of autosomal markers, we show that they have inherited ancestry from Africans, Indians, and possibly Europeans (Portuguese). Additionally, analyses of the uniparental (Y-chromosomal and mitochondrial DNA) markers indicate that the Siddis trace their ancestry to Bantu speakers from sub-Saharan Africa. We estimate that the admixture between the African ancestors of the Siddis and neighboring South Asian groups probably occurred in the past eight generations (～200 years ago), consistent with historical records.     

Mandibular size and shape variation in the hominins at Dmanisi, Republic of Georgia

The hominin fossils of Dmanisi, Republic of Georgia, present an ideal means of assessing levels of skeletal size and shape variation in a fossil hypodigm belonging to the genus Homo because they have been recovered from a spatially and temporally restricted context. We compare variation in mandible size and shape at Dmanisi to that of extant hominoids and extinct hominins. We use height and breadth measurements of the mandibular corpus at the first molar and the symphysis to assess size, and analyze shape based on size-adjusted (using a geometric mean) versions of these four variables. We compare size and shape variation at Dmanisi relative to all possible pairs of individuals within each comparative taxon using an exact resampling procedure of the ratio of D2600 to D211 and the average Euclidean distance (AED) between D2600 and D211, respectively. Comparisons to extant hominoids were conducted at both the specific and subspecific taxonomic levels and to extinct hominins by adopting both a more, and less speciose, hominin taxonomy. Results indicate that the pattern of variation for the Dmanisi hominins does not resemble that of any living species: they exhibit significantly more size variation when compared to modern humans, and they have significantly more corpus shape variation and size variation in corpus heights and overall mandible size than any extant ape species. When compared to fossil hominins they are also more dimorphic in size (although this result is influenced by the taxonomic hypothesis applied to the hominin fossil record). These results highlight the need to re-examine expectations of levels of sexual dimorphism in members of the genus Homo and to account for marked size and shape variation between D2600 and D211 under the prevailing view of a single hominin species at Dmanisi.     

The effect of selection history on extinction risk during severe environmental change

Environments rarely remain the same over time, and populations are therefore frequently at risk of going extinct when changes are significant enough to reduce fitness. Although many studies have investigated what attributes of the new environments and of the populations experiencing these changes will affect their probability of going extinct, limited work has been directed towards determining the role of population history on the probability of going extinct during severe environmental change. Here, we compare the extinction risk of populations with a history of selection in a benign environment, to populations with a history of selection in one or two stressful environments. We exposed spores and lines of the green alga Chlamydomonas reinhardtii from these three different histories to a range of severe environmental changes. We found that the extinction risk was higher for populations with a history of selection in stressful environments compared to populations with a history of selection in a benign environment. This effect was not due to differences in initial population sizes. Finally, the rates of extinction were highly repeatable within histories, indicating strong historical contingency of extinction risk. Hence, information on the selection history of a population can be used to predict their probability of going extinct during environmental change.     

Chimpanzees (Pan troglodytes verus) and their mammalian sympatriates: Mt. Assirik,Niokolo-Koba National Park,Senegal

In intact, mosaic ecosystems, chimpanzees are sympatric with a wide range of other mammals, which may be predators, prey, or competitors. We delve beyond the nominal data of species lists to interval-level data on 35 medium-bodied and large-bodied mammals encountered at a hot, dry, and open field site in far West Africa. Frequency of encounter, habitat where found, and number of individuals encountered are analysed for species for which enough data were accumulated. Further, we compare findings over three periods (1976–1979, 2000, 2012). Species most often encountered were those normally classed as typical savanna forms. Even a crude classification into forest, woodland, and grassland ecotypes yields differences in species likely to meet apes. Comparison of encounter rates over time was surprisingly congruent, although not all species seen in the 1970s survived to the 2000s. Overall, Assirik’s mammalian fauna is comparable to palaeo-faunal guilds sympatric with various extinct hominins.     

Lumbar lordosis of extinct hominins

The lordotic curvature of the lumbar spine (lumbar lordosis) in humans is a critical component in the ability to achieve upright posture and bipedal gait. Only general estimates of the lordotic angle (LA) of extinct hominins are currently available, most of which are based on the wedging of the vertebral bodies. Recently, a new method for calculating the LA in skeletal material has become available. This method is based on the relationship between the lordotic curvature and the orientation of the inferior articular processes relative to vertebral bodies in the lumbar spines of living primates. Using this relationship, we developed new regression models in order to calculate the LAs in hominins. The new models are based on primate group-means and were used to calculate the LAs in the spines of eight extinct hominins. The results were also compared with the LAs of modern humans and modern nonhuman apes. The lordotic angles of australopithecines (41° ± 4), H. erectus (45°) and fossil H. sapiens (54° ± 14) are similar to those of modern humans (51° ± 11). This analysis confirms the assumption that human-like lordotic curvature was a morphological change that took place during the acquisition of erect posture and bipedalism as the habitual form of locomotion. Neandertals have smaller lordotic angles (LA = 29° ± 4) than modern humans, but higher angles than nonhuman apes (22° ± 3). This suggests possible subtle differences in Neandertal posture and locomotion from that of modern humans.     

Signatures of the preagricultural peopling processes in sub-Saharan Africa as revealed by the phylogeography of early Y chromosome lineages

The study of Y chromosome variation has helped reconstruct demographic events associated with the spread of languages, agriculture, and pastoralism in sub-Saharan Africa, but little attention has been given to the early history of the continent. In order to overcome this lack of knowledge, we carried out a phylogeographic analysis of haplogroups A and B in a broad data set of sub-Saharan populations. These two lineages are particularly suitable for this objective because they are the two most deeply rooted branches of the Y chromosome genealogy. Their distribution is almost exclusively restricted to sub-Saharan Africa where their frequency peaks at 65% in groups of foragers. The combined high-resolution single nucleotide polymorphism analysis with short tandem repeats variation of their subclades reveals strong geographic and population structure for both haplogroups. This has allowed us to identify specific lineages related to regional preagricultural dynamics in different areas of sub-Saharan Africa. In addition, we observed signatures of relatively recent contact, both among Pygmies and between them and Khoisan speaker groups from southern Africa, thus contributing to the understanding of the complex evolutionary relationships among African hunter-gatherers. Finally, by revising the phylogeography of the very early human Y chromosome lineages, we have obtained support for the role of southern Africa as a sink, rather than a source, of the first migrations of modern humans from eastern and central parts of the continent. These results open new perspectives on the early history of Homo sapiens in Africa, with particular attention to areas of the continent where human fossil remains and archaeological data are scant.