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three-dimensional shape variation of talar surface morphology in hominoid primates.the hominoid foot is of particular interest to biological anthropologists, as changes in its anatomy through time reflect the adoption of terrestrial locomotion, particularly in species of australopithecus and homo. understanding the osteological morphology associated with changes in whole foot function and the development of the plantar medial longitudinal foot arch are key to understanding the transition through habitual bipedalism in australopithecines to obligate bipedalism and long-distance ...201424842795
insights into the genetic structure and diversity of 38 south asian indians from deep whole-genome sequencing.south asia possesses a significant amount of genetic diversity due to considerable intergroup differences in culture and language. there have been numerous reports on the genetic structure of asian indians, although these have mostly relied on genotyping microarrays or targeted sequencing of the mitochondria and y chromosomes. asian indians in singapore are primarily descendants of immigrants from dravidian-language-speaking states in south india, and 38 individuals from the general population u ...201424832686
reply to ding et al.: non-neanderthal origin of the hla-dpb1*0401. 201424829968
neandertal demise: an archaeological analysis of the modern human superiority complex.neandertals are the best-studied of all extinct hominins, with a rich fossil record sampling hundreds of individuals, roughly dating from between 350,000 and 40,000 years ago. their distinct fossil remains have been retrieved from portugal in the west to the altai area in central asia in the east and from below the waters of the north sea in the north to a series of caves in israel in the south. having thrived in eurasia for more than 300,000 years, neandertals vanished from the record around 40 ...201424789039
reconstructing the dna methylation maps of the neandertal and the denisovan.ancient dna sequencing has recently provided high-coverage archaic human genomes. however, the evolution of epigenetic regulation along the human lineage remains largely unexplored. we reconstructed the full dna methylation maps of the neandertal and the denisovan by harnessing the natural degradation processes of methylated and unmethylated cytosines. comparing these ancient methylation maps to those of present-day humans, we identified ~2000 differentially methylated regions (dmrs). particular ...201424786081
the shape of the human language-ready brain.our core hypothesis is that the emergence of our species-specific language-ready brain ought to be understood in light of the developmental changes expressed at the levels of brain morphology and neural connectivity that occurred in our species after the split from neanderthals-denisovans and that gave us a more globular braincase configuration. in addition to changes at the cortical level, we hypothesize that the anatomical shift that led to globularity also entailed significant changes at the ...201424772099
reconstructing cultural behavior from dental wear studies: is para-facets analysis approach scientifically valid?it is common knowledge, that in archaic populations teeth were used as tools, this behavior can be studied by evaluating attrition patterns. parafacets were defined as nonmasticatory wear areas that have no antagonist matching wear facets. the presence of the parafacets led to far-reaching conclusions regarding cultural interactions between near eastern neanderthals and anatomically modern humans. this study was aimed to examine the identification of parafacets in some of the qafzeh specimens. b ...201424767822
detecting human presence at the border of the northeastern italian pre-alps. 14c dating at rio secco cave as expression of the first gravettian and the late mousterian in the northern adriatic region.in the northern adriatic regions, which include the venetian region and the dalmatian coast, late neanderthal settlements are recorded in few sites and even more ephemeral are remains of the mid-upper palaeolithic occupations. a contribution to reconstruct the human presence during this time range has been produced from a recently investigated cave, rio secco, located in the northern adriatic region at the foot of the carnic pre-alps. chronometric data make rio secco a key site in the context of ...201424759802
patterns of coding variation in the complete exomes of three neandertals.we present the dna sequence of 17,367 protein-coding genes in two neandertals from spain and croatia and analyze them together with the genome sequence recently determined from a neandertal from southern siberia. comparisons with present-day humans from africa, europe, and asia reveal that genetic diversity among neandertals was remarkably low, and that they carried a higher proportion of amino acid-changing (nonsynonymous) alleles inferred to alter protein structure or function than present-day ...201424753607
a 400,000-year-old mitochondrial genome questions phylogenetic relationships amongst archaic hominins: using the latest advances in ancient genomics, the mitochondrial genome sequence of a 400,000-year-old hominin has been deciphered.by combining state-of-the-art approaches in ancient genomics, meyer and co-workers have reconstructed the mitochondrial sequence of an archaic hominin that lived at sierra de atapuerca, spain about 400,000 years ago. this achievement follows recent advances in molecular anthropology that delivered the genome sequence of younger archaic hominins, such as neanderthals and denisovans. molecular phylogenetic reconstructions placed the atapuercan as a sister group to denisovans, although its morpholo ...201424706482
non-neanderthal origin of the hla-dpb1*0401. 201424706180
taxonomic differences in deciduous upper second molar crown outlines of homo sapiens, homo neanderthalensis and homo erectus.a significant number of middle to late pleistocene sites contain primarily (and sometimes only) deciduous teeth (e.g., grotta del cavallo, mezmaiskaya, blombos). not surprisingly, there has been a recent renewed interest in deciduous dental variation, especially in the context of distinguishing homo neanderthalensis and homo sapiens. most studies of the deciduous dentition of fossil hominins have focused on standard metrical variation but morphological (non-metric and morphometric) variation als ...201424703186
neanderthal ancestry drives evolution of lipid catabolism in contemporary europeans.although neanderthals are extinct, fragments of their genomes persist in contemporary humans. here we show that while the genome-wide frequency of neanderthal-like sites is approximately constant across all contemporary out-of-africa populations, genes involved in lipid catabolism contain more than threefold excess of such sites in contemporary humans of european descent. evolutionally, these genes show significant association with signatures of recent positive selection in the contemporary euro ...201424690587
neanderthal paintings? production of prototypical human (homo sapiens) faces shows systematic distortions.people's sketches of human faces seem to be systematically distorted: the eye position is always higher than in reality. this bias was experimentally analyzed by a series of experiments varying drawing conditions. participants either drew prototypical faces from memory (studies 1 and 2: free reconstruction; study 3: cued reconstruction) or directly copied average faces (study 4). participants consistently showed this positioning bias, which is even in accord with facial depictions published in i ...201424689136
the human condition-a molecular approach.research into when and where modern humans originated and how they differ from, and interacted with, other now-extinct forms of human has so far been the realm of archaeologists and paleoanthropologists. however, over the past decade, molecular geneticists have begun to study genomes of extinct humans. here, i discuss where we stand today with respect to understanding how modern humans came to differ from neandertals and other human forms that existed until about 30,000 years ago.201424679537
human evolution. oldest homo sapiens genome pinpoints neandertal input. 201424675930
genetic variation of the whole icam4 gene in caucasians and african americans.landsteiner-wiener (lw) is the human blood group system number 16, which comprises two antithetical antigens, lw(a) and lw(b) and the high-prevalence antigen lw(ab) . lw is encoded by the intracellular adhesion molecule 4 (icam4) gene. the icam4 protein is part of the rhesus complex in the red cell membrane and is involved in cell-cell adhesion.201424673173
human evolution: the neanderthal in the family. 201424670743
searching for signatures of cold adaptations in modern and archaic humans: hints from the brown adipose tissue genes.adaptation to low temperatures has been reasonably developed in the human species during the colonization of the eurasian landmass subsequent to out of africa migrations of anatomically modern humans. in addition to morphological and cultural changes, also metabolic ones are supposed to have favored human isolation from cold and body heat production and this can be hypothesized also for most neandertal and at least for some denisovan populations, which lived in geographical areas that strongly e ...201424667833
middle paleolithic and uluzzian human remains from fumane cave, italy.the site of fumane cave (western lessini mountains, italy) contains a stratigraphic sequence spanning the middle to early upper paleolithic. during excavations from 1989 to 2011, four human teeth were unearthed from the mousterian (fumane 1, 4, 5) and uluzzian (fumane 6) levels of the cave. in this contribution, we provide the first morphological description and morphometric analysis of the dental remains. all of the human remains, except for fumane 6, are deciduous teeth. based on metric data ( ...201424666601
the chronology of the earliest upper palaeolithic in northern iberia: new insights from l'arbreda, labeko koba and la viña.since the late 1980s, northern iberia has yielded some of the earliest radiocarbon dated aurignacian assemblages in western europe, probably produced by anatomically modern humans (amhs). this is at odds with its location furthest from the likely eastern entry point of amhs, and has also suggested to some that the châtelperronian resulted from cultural transfer from amhs to neanderthals. however, the accuracy of the early chronology has been extensively disputed, primarily because of the poor as ...201424636733
evidence for a (15)n positive excursion in terrestrial foodwebs at the middle to upper palaeolithic transition in south-western france: implications for early modern human palaeodiet and palaeoenvironment.the middle to upper palaeolithic transition around 35,000 years ago coincides with the replacement of neanderthals by anatomically modern humans in europe. several hypotheses have been suggested to explain this replacement, one of them being the ability of anatomically modern humans to broaden their dietary spectrum beyond the large ungulate prey that neanderthals consumed exclusively. this scenario is notably based on higher nitrogen-15 amounts in early upper palaeolithic anatomically modern hu ...201424630359
the doubly conditioned frequency spectrum does not distinguish between ancient population structure and hybridization.distinguishing between hybridization and population structure in the ancestral species is a key challenge in our understanding of how permeable species boundaries are to gene flow. the doubly conditioned frequency spectrum (dcfs) has been argued to be a powerful metric to discriminate between these two explanations, and it was used to argue for hybridization between neandertal and anatomically modern humans. the shape of the observed dcfs for these two species cannot be reproduced by a model tha ...201424627034
in search of our direct ancestor an anthropological and orthodontic summary.this paper reviews the development of human facial anatomy in h erectus, neanderthal and modern man. modern orthodontic measurements are used to compare different jaw size and relationships.201424620439
neandertal clavicle length.the late pleistocene archaic humans from western eurasia (the neandertals) have been described for a century as exhibiting absolutely and relatively long clavicles. this aspect of their body proportions has been used to distinguish them from modern humans, invoked to account for other aspects of their anatomy and genetics, used in assessments of their phylogenetic polarities, and used as evidence for late pleistocene population relationships. however, it has been unclear whether the usual scalin ...201424616525
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 ...201424615397
plant foods and the dietary ecology of neanderthals and early modern humans.one of the most important challenges in anthropology is understanding the disappearance of neanderthals. previous research suggests that neanderthals had a narrower diet than early modern humans, in part because they lacked various social and technological advances that lead to greater dietary variety, such as a sexual division of labor and the use of complex projectile weapons. the wider diet of early modern humans would have provided more calories and nutrients, increasing fertility, decreasin ...201424612646
brief communication: investigation of the semicircular canal variation in the krapina neandertals.previous studies comparing bony labyrinth morphology in geographically-dispersed samples of neandertals and modern homo sapiens (h. sapiens) showed that neandertals generally have smaller semicircular canals than modern h. sapiens (hublin et al., ; spoor et al., ; glantz et al., ). here we analyze the morphology of a single group of neandertal specimens from one locale, the krapina site, to determine the intraspecific variation in neandertal semicircular canal sizes. dimensions of the semicircul ...201424610299
extreme population differences in the human zinc transporter zip4 (slc39a4) are explained by positive selection in sub-saharan africa.extreme differences in allele frequency between west africans and eurasians were observed for a leucine-to-valine substitution (leu372val) in the human intestinal zinc uptake transporter, zip4, yet no further evidence was found for a selective sweep around the zip4 gene (slc39a4). by interrogating allele frequencies in more than 100 diverse human populations and resequencing neanderthal dna, we confirmed the ancestral state of this locus and found a strong geographical gradient for the derived a ...201424586184
neandertal admixture in eurasia confirmed by maximum-likelihood analysis of three genomes.although there has been much interest in estimating histories of divergence and admixture from genomic data, it has proved difficult to distinguish recent admixture from long-term structure in the ancestral population. thus, recent genome-wide analyses based on summary statistics have sparked controversy about the possibility of interbreeding between neandertals and modern humans in eurasia. here we derive the probability of full mutational configurations in nonrecombining sequence blocks under ...201424532731
on the chronology of the uluzzian.the uluzzian, one of europe's 'transitional' technocomplexes, has gained particular significance over the past three years when the only human remains associated with it were attributed to modern humans, instead of neanderthals as previously thought. the position of the uluzzian at stratified sequences, always overlying late mousterian layers and underlying early upper palaeolithic ones, highlights its significance in understanding the passage from the middle to upper palaeolithic, as well as th ...201424513033
hyperostosis frontalis interna in a neandertal from marillac (charente, france).the site of marillac (charente, france) has yielded an important stratigraphic sequence containing numerous neandertal remains (some of them with peri-mortem manipulations) from lithofacies 2 (quina mousterian). this level has been correlated with mis 4 and is associated with a tl date of 57,600 ± 4600 years bp (before present). the study of one of the cranial fragments (marillac 3) revealed a grade 2 or type b hyperostosis frontalis interna (hfi), remodelling and altering the internal table of ...201424491378
influence of lower limb configuration on walking cost in late pleistocene humans.it has been proposed that neandertals had about 30% higher gross cost of transport than anatomically modern humans (amh) and that such difference implies higher daily energy demands and reduced foraging ranges in neandertals. thus, reduced walking economy could be among the factors contributing to the neandertals' loss in competition with their anatomically modern successors. previously, neandertal walking cost had been estimated from just two parameters and based upon a pooled-sex sample. in th ...201424485350
human evolution. neandertals and moderns made imperfect mates. 201424482455
allometry, merism, and tooth shape of the upper deciduous m2 and permanent m1.the aims of this study were to investigate the effect of allometry on the shape of dm(2) and m(1) crown outlines and to examine whether the trajectory and magnitude of scaling are shared between species. the sample included 160 recent homo sapiens, 28 upper paleolithic h. sapiens, 10 early h. sapiens, and 33 h. neanderthalensis (neandertal) individuals. of these, 97 were dm(2) /m(1) pairs from the same individuals. a two-block partial least squares analysis of paired individuals revealed a signi ...201424482249
the genomic landscape of neanderthal ancestry in present-day humans.genomic studies have shown that neanderthals interbred with modern humans, and that non-africans today are the products of this mixture. the antiquity of neanderthal gene flow into modern humans means that genomic regions that derive from neanderthals in any one human today are usually less than a hundred kilobases in size. however, neanderthal haplotypes are also distinctive enough that several studies have been able to detect neanderthal ancestry at specific loci. we systematically infer neand ...201424476815
resurrecting surviving neandertal lineages from modern human genomes.anatomically modern humans overlapped and mated with neandertals such that non-african humans inherit ~1 to 3% of their genomes from neandertal ancestors. we identified neandertal lineages that persist in the dna of modern humans, in whole-genome sequences from 379 european and 286 east asian individuals, recovering more than 15 gigabases of introgressed sequence that spans ~20% of the neandertal genome (false discovery rate = 5%). analyses of surviving archaic lineages suggest that there were f ...201424476670
separating endogenous ancient dna from modern day contamination in a siberian neandertal.one of the main impediments for obtaining dna sequences from ancient human skeletons is the presence of contaminating modern human dna molecules in many fossil samples and laboratory reagents. however, dna fragments isolated from ancient specimens show a characteristic dna damage pattern caused by miscoding lesions that differs from present day dna sequences. here, we develop a framework for evaluating the likelihood of a sequence originating from a model with postmortem degradation-summarized i ...201424469802
study of modern human evolution via comparative analysis with the neanderthal genome.many other human species appeared in evolution in the last 6 million years that have not been able to survive to modern times and are broadly known as archaic humans, as opposed to the extant modern humans. it has always been considered fascinating to compare the modern human genome with that of archaic humans to identify modern human-specific sequence variants and figure out those that made modern humans different from their predecessors or cousin species. neanderthals are the latest humans to ...201324465235
sunlight adaptation region of neanderthal genome found in up to 65% of modern east asian populations. 201424458432
[genes tell of the neanderthal role in human evolution]. 201424432504
loss of olfactory receptor function in hominin evolution.the mammalian sense of smell is governed by the largest gene family, which encodes the olfactory receptors (ors). the gain and loss of or genes is typically correlated with adaptations to various ecological niches. modern humans have 853 or genes but 55% of these have lost their function. here we show evidence of additional or loss of function in the neanderthal and denisovan hominin genomes using comparative genomic methodologies. ten neanderthal and 8 denisovan ors show evidence of loss of fun ...201424392153
sequence variants in slc16a11 are a common risk factor for type 2 diabetes in mexico.performing genetic studies in multiple human populations can identify disease risk alleles that are common in one population but rare in others, with the potential to illuminate pathophysiology, health disparities, and the population genetic origins of disease alleles. here we analysed 9.2 million single nucleotide polymorphisms (snps) in each of 8,214 mexicans and other latin americans: 3,848 with type 2 diabetes and 4,366 non-diabetic controls. in addition to replicating previous findings, we ...201424390345
a penile spine/vibrissa enhancer sequence is missing in modern and extinct humans but is retained in multiple primates with penile spines and sensory vibrissae.previous studies show that humans have a large genomic deletion downstream of the androgen receptor gene that eliminates an ancestral mammalian regulatory enhancer that drives expression in developing penile spines and sensory vibrissae. here we use a combination of large-scale sequence analysis and pcr amplification to demonstrate that the penile spine/vibrissa enhancer is missing in all humans surveyed and in the neandertal and denisovan genomes, but is present in dna samples of chimpanzees an ...201324367647
micro-biomechanics of the kebara 2 hyoid and its implications for speech in neanderthals.the description of a neanderthal hyoid from kebara cave (israel) in 1989 fuelled scientific debate on the evolution of speech and complex language. gross anatomy of the kebara 2 hyoid differs little from that of modern humans. however, whether homo neanderthalensis could use speech or complex language remains controversial. similarity in overall shape does not necessarily demonstrate that the kebara 2 hyoid was used in the same way as that of homo sapiens. the mechanical performance of whole bon ...201324367509
the complete genome sequence of a neanderthal from the altai mountains.we present a high-quality genome sequence of a neanderthal woman from siberia. we show that her parents were related at the level of half-siblings and that mating among close relatives was common among her recent ancestors. we also sequenced the genome of a neanderthal from the caucasus to low coverage. an analysis of the relationships and population history of available archaic genomes and 25 present-day human genomes shows that several gene flow events occurred among neanderthals, denisovans a ...201424352235
identification of putative target genes of the transcription factor runx2.comparisons of the genomes of neandertals and denisovans with present-day human genomes have suggested that the gene runx2, which encodes a transcription factor, may have been positively selected during early human evolution. here, we overexpress runx2 in ten human cell lines and identify genes that are directly or indirectly affected by runx2 expression. we find a number of genes not previously known to be affected by runx2 expression, in particular birc3, genes encoded on the mitochondrial gen ...201324349465
evidence supporting an intentional neandertal burial at la chapelle-aux-saints.the bouffia bonneval at la chapelle-aux-saints is well known for the discovery of the first secure neandertal burial in the early 20th century. however, the intentionality of the burial remains an issue of some debate. here, we present the results of a 12-y fieldwork project, along with a taphonomic analysis of the human remains, designed to assess the funerary context of the la chapelle-aux-saints neandertal. we have established the anthropogenic nature of the burial pit and underlined the taph ...201424344286
early pleistocene third metacarpal from kenya and the evolution of modern human-like hand morphology.despite discoveries of relatively complete hands from two early hominin species (ardipithecus ramidus and australopithecus sediba) and partial hands from another (australopithecus afarensis), fundamental questions remain about the evolution of human-like hand anatomy and function. these questions are driven by the paucity of hand fossils in the hominin fossil record between 800,000 and 1.8 my old, a time interval well documented for the emergence and subsequent proliferation of acheulian technol ...201424344276
human species and mating systems: neandertal-homo sapiens reproductive isolation and the archaeological and fossil records.the present paper examined the assumption of strong reproductive isolation (ri) between homo neanderthalensis and homo sapiens, as well as the question of what form it might have taken, using insights from the parallel case of chimpanzee–bonobo hybridization. ri from hybrid sterility or inviability was thought unlikely based on the short separation-to-introgression timeline. the forms of ri that typically develop in primates have relatively short timelines (especially for partial implementation) ...201324344097
neanderthal introgression at chromosome 3p21.31 was under positive natural selection in east asians.studies of the neanderthal and denisovan genomes demonstrate archaic hominin introgression in eurasians. here, we present evidence of neanderthal introgression within the chromosome 3p21.31 region, occurring with a high frequency in east asians (ranging from 49.4% to 66.5%) and at a low frequency in europeans. we also detected a signal of strong positive selection in this region only in east asians. our data indicate that likely candidate targets of selection include rs12488302-t and its associa ...201424336922
a reassessment of the presumed neandertal remains from san bernardino cave, italy.in 1986-1987, three human remains were unearthed from macro-unit ii of san bernardino cave (berici hills, veneto, italy), a deposit containing a late mousterian lithic assemblage. the human remains (a distal phalanx, a lower right third molar and a lower right second deciduous incisor) do not show diagnostic morphological features that could be used to determine whether they were from homo neanderthalensis or homo sapiens. despite being of small size, and thus more similar to recent h. sapiens, ...201424331083
within a sample from a population, the distribution of the number of descendants of a subsample's most recent common ancestor.sample n individuals uniformly at random from a population, and then sample m individuals uniformly at random from the sample. consider the most recent common ancestor (mrca) of the subsample of m individuals. let the subsample mrca have j descendants in the sample (m ⩽ j ⩽ n). under a moran or coalescent model (and therefore under many other models), the probability that j = n is known. in this case, the subsample mrca is an ancestor of every sampled individual, and the subsample and sample mrc ...201424321308
neanderthal and denisovan retroviruses in modern humans. 201324262833
a novel family of human leukocyte antigen class ii receptors may have its origin in archaic human species.hla class ii α and β chains form receptors for antigen presentation to cd4(+) t cells. numerous pairings of class ii α and β subunits from the wide range of haplotypes and isotypes may form, but most of these combinations, in particular those produced by isotype mixing, yielded mismatched dimers. it is unclear how selection of functional receptors is achieved. at the atomic level, it is not known which interactions of class ii residues regulate selection of matched αβ heterodimers and the evolut ...201424214983
neanderthal axial and appendicular remains from moula-guercy, ardèche, france.excavations carried out during the 1990s at moula-guercy cave ardèche, france, yielded 108 hominid specimens dating to 100-120 ka. in this paper, we describe and compare the 39 axial and appendicular specimens not including hand and foot bones. among these remains are a large adult femur, several clavicles, a likely antimeric pair of radial heads, and a nearly complete superior pubic ramus. analyses of this material indicate a clear affinity with neanderthals by the presence of large and robust ...201324214385
neanderthal hand and foot remains from moula-guercy, ardèche, france.the hand and foot remains from moula-guercy cave (ardèche, france) comprise 24 specimens of eemian age (ca. 120 ka). the specimens include primarily complete elements, which are rare among the moula-guercy postcrania. the hand remains have several characteristic neanderthal traits including a laterally facing (parasagittally oriented) second metacarpal-capitate articulation, a short styloid process, a wide proximal articular surface on the third metacarpal, and absolutely expanded apical tuberos ...201324214384
hapfabia: identification of very short segments of identity by descent characterized by rare variants in large sequencing data.identity by descent (ibd) can be reliably detected for long shared dna segments, which are found in related individuals. however, many studies contain cohorts of unrelated individuals that share only short ibd segments. new sequencing technologies facilitate identification of short ibd segments through rare variants, which convey more information on ibd than common variants. current ibd detection methods, however, are not designed to use rare variants for the detection of short ibd segments. sho ...201324174545
apparent variation in neanderthal admixture among african populations is consistent with gene flow from non-african populations.recent studies have found evidence of introgression from neanderthals into modern humans outside of sub-saharan africa. given the geographic range of neanderthals, the findings have been interpreted as evidence of gene exchange between neanderthals and modern humans descended from the out-of-africa (ooa) migration. here, we examine an alternative interpretation in which the introgression occurred earlier within africa, between ancestors or relatives of neanderthals and a subset of african modern ...201324162011
toothpicking and periodontal disease in a neanderthal specimen from cova foradà site (valencia, spain).we present a neanderthal maxilla (cf-1) from cova foradà site (oliva, valencia, spain) with periodontal disease and evidence of attempts to alleviate pain with the use of a toothpick. two interproximal grooves have been found on the distal surfaces of the upper left pm(3) and m(1) of cf-1 maxilla. the location, morphology and size of the grooves coincide with other interproximal grooves found on the teeth of other fossil specimens. heavy dental wear and periodontal disease would have caused the ...201324146934
san bernardino cave (italy) and the appearance of levallois technology in europe: results of a radiometric and technological reassessment.the introduction of levallois technology in europe marked the transition from the lower to the early middle paleolithic. this new method of flake production was accompanied by significant behavioral changes in hominin populations. the emergence of this technological advance is considered homogeneous in the european archaeological record at the marine isotopic stage (mis) 9/mis 8 boundary. in this paper we report a series of combined electron spin resonance/u-series dates on mammal bones and teet ...201324146836
no known hominin species matches the expected dental morphology of the last common ancestor of neanderthals and modern humans.a central problem in paleoanthropology is the identity of the last common ancestor of neanderthals and modern humans ([n-mh]lca). recently developed analytical techniques now allow this problem to be addressed using a probabilistic morphological framework. this study provides a quantitative reconstruction of the expected dental morphology of the [n-mh]lca and an assessment of whether known fossil species are compatible with this ancestral position. we show that no known fossil species is a suita ...201324145426
elongation as a factor in artefacts of humans and other animals: an acheulean example in comparative context.elongation is a commonly found feature in artefacts made and used by humans and other animals and can be analysed in comparative study. whether made for use in hand or beak, the artefacts have some common properties of length, breadth, thickness and balance point, and elongation can be studied as a factor relating to construction or use of a long axis. in human artefacts, elongation can be traced through the archaeological record, for example in stone blades of the upper palaeolithic (traditiona ...201324101633
the fragmented character of middle palaeolithic stone tool technology.the importance of the transport of stone artefacts in structuring neandertal lithic assemblages has often been addressed, but the degree to which this led to fragmentation of lithic reduction over middle palaeolithic landscapes has not been explicitly studied thus far. large-scale excavations of middle palaeolithic open-air sites and refitting studies of the retrieved assemblages have yielded new, high-resolution data on the mobile aspects of neandertal stone tool technology. in this paper, we i ...201324074611
proteogenomic review of the changes in primate apoc-i during evolution.apolipoprotein c-i has evolved more rapidly than any of the other soluble apolipoproteins. during the course of primate evolution, the gene for this apolipoprotein was duplicated. prompted by our observation that the two resulting genes encode two distinct forms of apoc-i in great apes, we have reviewed both the genomic and proteomic data to examine what changes have occurred during the course of primate evolution. we have found data showing that one of the duplicated genes, known to be a pseudo ...201328757862
neanderthal language? just-so stories take center stage. 201324069017
brief communication: a morphometric analysis of the neandertal upper second molar leuca i.the scarcity of neandertal remains from southern europe hampers our understanding of neandertal variability, and can bias interpretations about neandertal geographic variation. to address this issue, it is often important to reassess human remains that, while discovered decades ago, remain relatively unknown to the scientific community. in this contribution, we provide a complete state-of-the-art comparative morphometric analysis of leuca i, an unworn left second upper molar (lm(2) ) discovered ...201323999832
evaluating developmental shape changes in homo antecessor subadult facial morphology.the fossil atd6-69 from atapuerca, spain, dated to ca. 900 ka (thousands of years ago) has been suggested to mark the earliest appearance of modern human facial features. however, this specimen is a subadult and the interpretation of its morphology remains controversial, because it is unclear how developmental shape changes would affect the features that link atd6-69 to modern humans. here we analyze atd6-69 in an evolutionary and developmental context. our modern human sample comprises cross-se ...201323998458
[mitochondrial dna polymorphisms shared between modern humans and neanderthals: adaptive convergence or evidence for interspecific hybridization?].an analysis of the variability of the nucleotide sequences in the mitochondrial genome of modern humans, neanderthals, denisovans, and other primates has shown that there are shared polymorphisms at positions 2758 and 7146 between modern homo sapiens (in phylogenetic cluster l2'3'4'5'6) and homo neanderthalensis (in the group of european neanderthals younger than 48000 years). it is suggested that the convergence may be due to adaptive changes in the mitochondrial genomes of modern humans and ne ...201325508911
[mitochondrial dna polymorphisms shared between modern humans and neanderthals: adaptive convergence or evidence for interspecific hybridization?].an analysis of the variability of the nucleotide sequences in the mitochondrial genome of modern humans, neanderthals, denisovans, and other primates has shown that there are shared polymorphisms at positions 2758 and 7146 between modern homo sapiens (in phylogenetic cluster l2'3'4'5'6) and homo neanderthalensis (in the group of european neanderthals younger than 48000 years). it is suggested that the convergence may be due to adaptive changes in the mitochondrial genomes of modern humans and ne ...201325486780
hominin evolution and gene flow in the pleistocene africa.africa demonstrates a complex process of the hominin evolution with a series of adaptive radiations during several millions of years that led to diverse morphological forms. recently, hammer et al. (2011) and harvati et al. (2011) provided integrated morphological and genetic evidence of interbreeding between modern humans and unknown archaic hominins in africa as recently as 35,000 years ago. however, a genetic evidence of hybridization between hominin lineages during the lower and middle pleis ...201323980394
using the neanderthal and denisova genetic data to understand the common mapt 17q21 inversion in modern humans.the polymorphic inversion on 17q21, that includes the mapt gene, represents a unique locus in the human genome characterized by a large region with strong linkage disequilibrium. two distinct haplotypes, h1 and h2, exist in modern humans, and h1 has been unequivocally related to several neurodegenerative disorders. recent data indicate that recurrent inversions of this genomic region have occurred through primate evolution, with the h2 haplotype being the ancestral state. neandertals harbored th ...201223959642
neandertals made the first specialized bone tools in europe.modern humans replaced neandertals ∼40,000 y ago. close to the time of replacement, neandertals show behaviors similar to those of the modern humans arriving into europe, including the use of specialized bone tools, body ornaments, and small blades. it is highly debated whether these modern behaviors developed before or as a result of contact with modern humans. here we report the identification of a type of specialized bone tool, lissoir, previously only associated with modern humans. the micro ...201323940333
regional behaviour among late neanderthal groups in western europe: a comparative assessment of late middle palaeolithic bifacial tool variability.population dynamics between and within pleistocene groups are vital to understanding wider behavioural processes like social transmission and cultural variation. the late middle palaeolithic (mis 5d-3, ca. 115,000-35,000 bp [years before present]) permits a novel, data-driven assessment of these concepts through a unique record: bifacial tools made by classic neanderthals. previously, studies of late middle palaeolithic bifacial tools were hampered by a convoluted plethora of competing terms, ty ...201323928352
middle pleistocene ecology and neanderthal subsistence: insights from stable isotope analyses in payre (ardèche, southeastern france).the middle palaeolithic site of payre in southeastern france yields abundant archaeological material associated with fossil hominid remains. with its long sequence of middle pleistocene deposits, payre is a key site to study the middle palaeolithic chronology of this region. this study is the first to investigate carbon and oxygen isotope contents of neanderthal tooth enamel bioapatite, together with a wide range of herbivorous and carnivorous species. the aim is to contribute to the understandi ...201323920410
morphometric analysis of molars in a middle pleistocene population shows a mosaic of 'modern' and neanderthal features.previous studies of upper first molar (m1) crown shape have shown significant differences between homo sapiens and homo neanderthalensis that were already present in the european middle pleistocene populations, including the large dental sample from atapuerca-sima de los huesos (sh). analysis of other m1 features such as the total crown base area, cusp proportions, cusp angles and occlusal polygon have confirmed the differences between both lineages, becoming a useful tool for the taxonomic assi ...201323914934
the cognitive ability of extinct hominins: bringing down the hierarchy using genomic evidences.the availability of the full genomes of homo sapiens, homo neanderthalensis, and denisovans, as well as modern bioinformatic tools, are opening new possibilities for the understanding of the differences and similarities present in these taxa.201323907779
adaptive introgression in animals: examples and comparison to new mutation and standing variation as sources of adaptive variation.adaptive genetic variation has been thought to originate primarily from either new mutation or standing variation. another potential source of adaptive variation is adaptive variants from other (donor) species that are introgressed into the (recipient) species, termed adaptive introgression. here, the various attributes of these three potential sources of adaptive variation are compared. for example, the rate of adaptive change is generally thought to be faster from standing variation, slower fr ...201323906376
dental wear and cultural behavior in middle paleolithic humans from the near east.neanderthals and anatomically modern humans (amhs) may have lived in close proximity in the near east region during middle paleolithic times. although functional morphological analyses suggest a marked behavioral contrast between these two human groups, new dental micro- and macro-wear studies, together with new archaeological data, have revealed some similarities in ecology and dietary habits. in this study, we analyze the tooth wear patterns of neanderthals and amh from middle paleolithic site ...201323904240
an ochered fossil marine shell from the mousterian of fumane cave, italy.a scanty but varied ensemble of finds challenges the idea that neandertal material culture was essentially static and did not include symbolic items. in this study we report on a fragmentary miocene-pliocene fossil marine shell, aspamarginata, discovered in a discoid mousterian layer of the fumane cave, northern italy, dated to at least 47.6-45.0 cal ky bp. the shell was collected by neandertals at a fossil exposure probably located more than 100 kms from the site. microscopic analysis of the sh ...201323874677
neanderthal and denisova genetic affinities with contemporary humans: introgression versus common ancestral polymorphisms.analyses of the genetic relationships among modern humans, neanderthals and denisovans have suggested that 1-4% of the non-sub-saharan african gene pool may be neanderthal derived, while 6-8% of the melanesian gene pool may be the product of admixture between the denisovans and the direct ancestors of melanesians. in the present study, we analyzed single nucleotide polymorphism (snp) diversity among a worldwide collection of contemporary human populations with respect to the genetic constitution ...201323872234
[application of the musculo-skeletal modelling software lhpfusionbox to a paleoanthropological problem: the spyrou neandertal moves!].lhpfusionbox is a program originally designed for biomechanical and clinical studies relating to the musculoskeletal system of anatomically modern humans (amh). the program has recently been adapted for paleontological purposes and used to reconstruct and biomechanically analyse a fossil hominid. there is no complete neandertal skeleton in the fossil record. the aim of the study was to reconstruct a complete three-dimensional (3d) model of a neandertal using the relatively complete spy ii neande ...201323859517
on the antiquity of language: the reinterpretation of neandertal linguistic capacities and its consequences.it is usually assumed that modern language is a recent phenomenon, coinciding with the emergence of modern humans themselves. many assume as well that this is the result of a single, sudden mutation giving rise to the full "modern package." however, we argue here that recognizably modern language is likely an ancient feature of our genus pre-dating at least the common ancestor of modern humans and neandertals about half a million years ago. to this end, we adduce a broad range of evidence from l ...201323847571
continuity of microblade technology in the indian subcontinent since 45 ka: implications for the dispersal of modern humans.we extend the continuity of microblade technology in the indian subcontinent to 45 ka, on the basis of optical dating of microblade assemblages from the site of mehtakheri, (22° 13' 44″ n lat 76° 01' 36″ e long) in madhya pradesh, india. microblade technology in the indian subcontinent is continuously present from its first appearance until the iron age (~3 ka), making its association with modern humans undisputed. it has been suggested that microblade technology in the indian subcontinent was d ...201323840912
earliest floral grave lining from 13,700-11,700-y-old natufian burials at raqefet cave, mt. carmel, israel.flowering plants possess mechanisms that stimulate positive emotional and social responses in humans. it is difficult to establish when people started to use flowers in public and ceremonial events because of the scarcity of relevant evidence in the archaeological record. we report on uniquely preserved 13,700-11,700-y-old grave linings made of flowers, suggesting that such use began much earlier than previously thought. the only potentially older instance is the questionable use of flowers in t ...201323818584
the campanian ignimbrite eruption: new data on volcanic ash dispersal and its potential impact on human evolution.the campanian ignimbrite (ci) volcanic eruption was the most explosive in europe in the last 200,000 years. the event coincided with the onset of an extremely cold climatic phase known as heinrich event 4 (he4) approximately 40,000 years ago. their combined effect may have exacerbated the severity of the climate through positive feedbacks across europe and possibly globally. the ci event is of particular interest not only to investigate the role of volcanism on climate forcing and palaeoenvironm ...201323799050
fibrous dysplasia in a 120,000+ year old neandertal from krapina, croatia.we describe the first definitive case of a fibrous dysplastic neoplasm in a neandertal rib (120.71) from the site of krapina in present-day croatia. the tumor predates other evidence for these kinds of tumor by well over 100,000 years. tumors of any sort are a rare occurrence in recent archaeological periods or in living primates, but especially in the human fossil record. several studies have surveyed bone diseases in past human populations and living primates and fibrous dysplasias occur in a ...201323755126
inferring demographic history from a spectrum of shared haplotype lengths.there has been much recent excitement about the use of genetics to elucidate ancestral history and demography. whole genome data from humans and other species are revealing complex stories of divergence and admixture that were left undiscovered by previous smaller data sets. a central challenge is to estimate the timing of past admixture and divergence events, for example the time at which neanderthals exchanged genetic material with humans and the time at which modern humans left africa. here, ...201323754952
genetic and archaeological perspectives on the initial modern human colonization of southern asia.it has been argued recently that the initial dispersal of anatomically modern humans from africa to southern asia occurred before the volcanic "supereruption" of the mount toba volcano (sumatra) at ∼74,000 y before present (b.p.)-possibly as early as 120,000 y b.p. we show here that this "pre-toba" dispersal model is in serious conflict with both the most recent genetic evidence from both africa and asia and the archaeological evidence from south asian sites. we present an alternative model base ...201323754394
neanderthal teeth from moula-guercy, ardèche, france.here we describe dental remains from a neanderthal fossil assemblage from moula-guercy, france. our report demonstrates that the moula-guercy hominid remains contribute important morphological, developmental, and behavioral data to understanding neanderthal evolutionary history. we include gross comparative morphological descriptions and enamel surface microstructure and microwear data. these teeth reveal numerous characteristics that are diagnostic of neanderthals and provide no evidence for th ...201323737145
population changes across the neanderthal-to-modern-human transition in western france: a reply to dogandžić and mcpherron (2013). 201323726931
a 175 million year history of t cell regulatory molecules reveals widespread selection, with adaptive evolution of disease alleles.t cell activation plays a central role in immune response and in the maintenance of self-tolerance. we analyzed the evolutionary history of t cell regulatory molecules. nine genes involved in triggering t cell activation or in regulating the ensuing response evolved adaptively in mammals. several positively selected sites overlap with positions interacting with the binding partner or with cellular components. population genetic analysis in humans revealed a complex scenario of local (faslg, cd40 ...201323707475
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 later ...201323706407
barium distributions in teeth reveal early-life dietary transitions in primates.early-life dietary transitions reflect fundamental aspects of primate evolution and are important determinants of health in contemporary human populations. weaning is critical to developmental and reproductive rates; early weaning can have detrimental health effects but enables shorter inter-birth intervals, which influences population growth. uncovering early-life dietary history in fossils is hampered by the absence of prospectively validated biomarkers that are not modified during fossilizati ...201323698370
neanderthal culture: old masters. 201323676735
handedness in neandertals from the el sidrón (asturias, spain): evidence from instrumental striations with ontogenetic inferences.the developed cognitive capabilities for homo sapiens seems to be the result of a specialized and lateralized brain, and as a result of this, humans display the highest degree of manual specialization or handedness among the primates. studies regarding its emergence and distribution within the genus homo show that handedness is present very early. the mode in which it was articulated and spread across the different species during the course of human evolution could provide information about our ...201323671635
the genochip: a new tool for genetic anthropology.the genographic project is an international effort aimed at charting human migratory history. the project is nonprofit and nonmedical, and, through its legacy fund, supports locally led efforts to preserve indigenous and traditional cultures. although the first phase of the project was focused on uniparentally inherited markers on the y-chromosome and mitochondrial dna (mtdna), the current phase focuses on markers from across the entire genome to obtain a more complete understanding of human gen ...201323666864
no ancient dna damage in actinobacteria from the neanderthal bone.the neanderthal genome was recently sequenced using dna extracted from a 38,000-year-old fossil. at the start of the project, the fraction of mammalian and bacterial dna in the sample was estimated to be <6% and 9%, respectively. treatment with restriction enzymes prior to sequencing increased the relative proportion of mammalian dna to 15%, but the large majority of sequences remain uncharacterized.201323658776
hiv infection reveals widespread expansion of novel centromeric human endogenous retroviruses.human endogenous retroviruses (hervs) make up 8% of the human genome. the herv-k (hml-2) family is the most recent group of these viruses to have inserted into the genome, and we have detected the activation of herv-k (hml-2) proviruses in the blood of patients with hiv-1 infection. we report that hiv-1 infection activates expression of a novel herv-k (hml-2) provirus, termed k111, present in multiple copies in the centromeres of chromosomes throughout the human genome yet not annotated in the m ...201323657884
dating human cultural capacity using phylogenetic principles.humans have genetically based unique abilities making complex culture possible; an assemblage of traits which we term "cultural capacity". the age of this capacity has for long been subject to controversy. we apply phylogenetic principles to date this capacity, integrating evidence from archaeology, genetics, paleoanthropology, and linguistics. we show that cultural capacity is older than the first split in the modern human lineage, and at least 170,000 years old, based on data on hyoid bone mor ...201323648831
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