Publications

evolution of the base of the brain in highly encephalized human species.	the increase of brain size relative to body size-encephalization-is intimately linked with human evolution. however, two genetically different evolutionary lineages, neanderthals and modern humans, have produced similarly large-brained human species. thus, understanding human brain evolution should include research into specific cerebral reorganization, possibly reflected by brain shape changes. here we exploit developmental integration between the brain and its underlying skeletal base to test ...	2011	22158443
Stature estimation from complete long bones in the Middle Pleistocene humans from the Sima de los Huesos, Sierra de Atapuerca (Spain).	Systematic excavations at the site of the Sima de los Huesos (SH) in the Sierra de Atapuerca (Burgos, Spain) have allowed us to reconstruct 27 complete long bones of the human species Homo heidelbergensis. The SH sample is used here, together with a sample of 39 complete Homo neanderthalensis long bones and 17 complete early Homo sapiens (Skhul/Qafzeh) long bones, to compare the stature of these three different human species. Stature is estimated for each bone using race- and sex-independent reg ...	2011	22196156
genetic and phenotypic consequences of introgression between humans and neanderthals.	strong evidence for introgression of neanderthal genes into parts of the modern human gene pool has recently emerged. the evidence indicates that some populations of modern humans have received infusions of genes from two different groups of neanderthals. one of these neanderthal groups lived in the middle east and central europe and the other group (the denisovans) is known to have lived in central asia and was probably more widespread. this review examines two questions. first, how were these ...	2011	22099691
archaic human ancestry in east asia.	recent studies of ancient genomes have suggested that gene flow from archaic hominin groups to the ancestors of modern humans occurred on two separate occasions during the modern human expansion out of africa. at the same time, decreasing levels of human genetic diversity have been found at increasing distance from africa as a consequence of human expansion out of africa. we analyzed the signal of archaic ancestry in modern human populations, and we investigated how serial founder models of huma ...	2011	22042846
Learning about human population history from ancient and modern genomes.	Genome-wide data, both from SNP arrays and from complete genome sequencing, are becoming increasingly abundant and are now even available from extinct hominins. These data are providing new insights into population history; in particular, when combined with model-based analytical approaches, genome-wide data allow direct testing of hypotheses about population history. For example, genome-wide data from both contemporary populations and extinct hominins strongly support a single dispersal of mode ...	2011	21850041
Neanderthals: fossil evidence and DNA.	Neanderthals inhabited Western Eurasia from approximately 300 to 30 thousand years ago (ka). They are distinguished by a unique combination of anatomical traits, and are commonly associated with Middle Paleolithic lithic industries. Current consensus among paleoanthropologists is that they represented a separate Eurasian human lineage, which evolved in isolation from the rest of the Old World and which shared a common ancestor with modern humans in the Middle Pleistocene. It is thought that some ...	2011	21957644
Estimating the distribution of probable age-at-death from dental remains of immature human fossils.	In two historic longitudinal growth studies, Moorrees et al. (Am J Phys Anthropol 21 (1963) 99-108; J Dent Res 42 (1963) 1490-1502) presented the "mean attainment age" for stages of tooth development for 10 permanent tooth types and three deciduous tooth types. These findings were presented graphically to assess the rate of tooth formation in living children and to age immature skeletal remains. Despite being widely cited, these graphical data are difficult to implement because there are no acco ...	2011	22190291
crown size and cusp proportions in homo antecessor upper first molars. a comment on quam et al. 2009.	a recent evaluation of upper first molar (m¹) crown size and cusp proportions in the genus homo (quam et al. 2009) describes homo antecessor as maintaining a primitive pattern of cusp proportions, similar to that identified in australopithecines and the earliest members of the genus homo. these results contrast with those of gómez-robles et al. (2007), who described the crown shape in these molars as derived and similar to neanderthals and european homo heidelbergensis. the reassessment of these ...	2011	21208207
late pleistocene adult mortality patterns and modern human establishment.	the establishment of modern humans in the late pleistocene, subsequent to their emergence in eastern africa, is likely to have involved substantial population increases, during their initial dispersal across southern asia and their subsequent expansions throughout africa and into more northern eurasia. an assessment of younger (20-40 y) versus older (>40 y) adult mortality distributions for late archaic humans (principally neandertals) and two samples of early modern humans (middle paleolithic a ...	2011	21220336
gorilla and orangutan brains conform to the primate cellular scaling rules: implications for human evolution.	gorillas and orangutans are primates at least as large as humans, but their brains amount to about one third of the size of the human brain. this discrepancy has been used as evidence that the human brain is about 3 times larger than it should be for a primate species of its body size. in contrast to the view that the human brain is special in its size, we have suggested that it is the great apes that might have evolved bodies that are unusually large, on the basis of our recent finding that the ...	2011	21228547
an x-linked haplotype of neandertal origin is present among all non-african populations.	recent work on the neandertal genome has raised the possibility of admixture between neandertals and the expanding population of homo sapiens who left africa between 80 and 50 kya (thousand years ago) to colonize the rest of the world. here, we provide evidence of a notable presence (9% overall) of a neandertal-derived x chromosome segment among all contemporary human populations outside africa. our analysis of 6,092 x-chromosomes from all inhabited continents supports earlier contentions that a ...	2011	21266489
calcaneus length determines running economy: implications for endurance running performance in modern humans and neandertals.	the endurance running (er) hypothesis suggests that distance running played an important role in the evolution of the genus homo. most researchers have focused on er performance in modern humans, or on reconstructing er performance in homo erectus, however, few studies have examined er capabilities in other members of the genus homo. here, we examine skeletal correlates of er performance in modern humans in order to evaluate the energetics of running in neandertals and early homo sapiens. recent ...	2011	21269660
nasopharyngeal carcinoma as a paradigm of cancer genetics.	the unusual incidence patterns for nasopharyngeal carcinoma (npc) in china, northeast india, arctic inuit, peninsular and island southeast asia, polynesian islanders, and north africans indicate a role for npc risk genes in chinese, chinese-related, and not-obviously chinese-related populations. renewed interest in npc genetic risk has been stimulated by a hypothesis that npc population patterns originated in bai-yue / pre-austronesian-speaking aborigines and were dispersed during the last glaci ...	2011	21272439
what lies beneath? an evaluation of lower molar trigonid crest patterns based on both dentine and enamel expression.	the nearly ubiquitous presence of a continuous crest connecting the protoconid and metaconid of the lower molars (often referred to as the middle trigonid crest), is one of several dental traits that distinguish homo neanderthalensis from homo sapiens. this study examined variation in trigonid crest patterns on the enamel and dentine surfaces to (1) evaluate the concordance between the morphology of trigonid crests at the inner dentine and the outer enamel surfaces; (2) examine their development ...	2011	21312178
a comprehensive functional analysis of ancestral human signal peptides.	with the sequencing of the neandertal genome, it has become possible to identify amino acid substitutions that occurred on the human lineage since its separation from the neandertal lineage. conceptually, it will therefore be possible to functionally analyze all such amino acid substitutions in the future. here, we analyze the function of substitutions that occurred during recent human evolution in n-terminal signal peptides. we develop a high-throughput flow cytometry-based assay to analyze sig ...	2011	20798139
linking dopamine neurotransmission and neurogenesis: the evolutionary history of the ntad (ncam1-ttc12-ankk1-drd2) gene cluster.	genetic studies have long suggested the important role of the drd2 gene in psychiatric disorders and behavior. further research has shown a conjoined effect of genes in the chr11q22-23 region, which includes the ncam1, ttc12, ankk1 and drd2 genes, or ntad cluster. despite a growing need to unravel the role of this cluster, few studies have taken into account interspecies and evolutionary approaches. this study shows that behaviorally relevant snps from the ntad cluster, such as rs1800497 (taq1a) ...	2012	23412349
comment: low dental caries rate in neandertals: the result of diet or the oral flora composition?	dental caries is an infectious disease caused by oral acidophilic bacteria feeding on fermentable sugars, e.g. streptococcus mutans. the frequency of dental caries in neandertals was very low. this was usually explained as the result of a low-sugar diet. recent research, however, revealed some regional differences between european and near eastern neandertals, with the latter consuming considerable amounts of plants including highly cariogenic dates. this discovery, compared with the results of ...	2012	22409830
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 th ...	2012	22052243
Evolutionary origin of bipolar disorder-revised: EOBD-R.	The hypothesis of the evolutionary origin of bipolar disorder (EOBD) synthesized ideas about the biological clock and seasonal shifts in mood (Rosenthal, Wehr) with theorizing that bipolar disorder descends from a pyknic (compact, cold-adapted) group (Kretchmer). The hypothesis suggested that bipolar behaviors evolved in the northern temperate zone as highly derived adaptations to the selective pressures of severe climatic conditions during the Pleistocene. Given evidence of Neandertal contribut ...	2012	22036090
la ferrassie 6 and the development of neandertal pubic morphology.	functional interpretations of the mediolateral elongation and superoinferior flattening of the superior pubic ramus of neandertals require knowledge of its ontogeny. metric comparisons between the la ferrassie 6 neandertal infant, aged 3-5 years, and a sample of modern infants reveal that the acetabulosymphyseal length of la ferrassie 6, relative to femoral length and iliac breadth, falls at the limits of the range of variation of the modern infants, while the relative height of its superior pub ...	2012	3303957
Articulatory capacity of Neanderthals, a very recent and human-like fossil hominin.	Scientists seek to use fossil and archaeological evidence to constrain models of the coevolution of human language and tool use. We focus on Neanderthals, for whom indirect evidence from tool use and ancient DNA appears consistent with an adaptation to complex vocal-auditory communication. We summarize existing arguments that the articulatory apparatus for speech had not yet come under intense positive selection pressure in Neanderthals, and we outline some recent evidence and analyses that chal ...	2012	22106429
Morphological description and comparison of the dental remains from Atapuerca-Sima de los Huesos site (Spain).	The systematic excavation of the Sima de los Huesos (SH) site in Sierra de Atapuerca (Burgos, Spain) has yielded the largest hominin collection worldwide for the Middle Pleistocene. The dental sample now consists of more than 500 teeth that provide exceptional opportunities to define the dental morphological pattern of a Middle Pleistocene population as well as develop hypotheses about the origins of the Neanderthals. The dental collection has now increased to over 533 specimens (525 permanent a ...	2012	22118969
paleoclimate during neandertal and anatomically modern human occupation at amud and qafzeh, israel: the stable isotope data.	the δ(13)c(en) and δ(18)o(en) values of goat and gazelle enamel carbonate indicate that neandertals at amud cave, israel (53-70 ka) lived under different ecological conditions than did anatomically modern humans at qafzeh cave, israel (approximately 92 ka). during the last glacial period, neandertals at amud cave lived under wetter conditions than those in the region today. neither faunal species ate arid-adapted c(4) plants or drought-stressed c(3) plants. the variation in gazelle δ(18)o(en) va ...	2012	22115545
palaeogenetic research at the el sidrón neanderthal site.	el sidrón (asturias, north of spain) is a subterranean karstic system, where the remains of a contemporaneous social neanderthal group dated to about 49,000 years ago have been being excavated since their accidental discovery in 1994. due to the particular preservation conditions of this site, all individuals identified so far have preserved dna, and the anticontamination measures implemented during the excavation have made palaeogenetic studies possible on all individuals. the el sidrón samples ...	2012	21482084
caveman couture. neandertals may have worn dark feathers.		2012	23230791
homo sapiens, homo neanderthalensis and the denisova specimen: new insights on their evolutionary histories using whole-genome comparisons.	after a brief review of the most recent findings in the study of human evolution, an extensive comparison of the complete genomes of our nearest relative, the chimpanzee (pan troglodytes), of extant homo sapiens, archaic homo neanderthalensis and the denisova specimen were made. the focus was on non-synonymous mutations, which consequently had an impact on protein levels and these changes were classified according to degree of effect. a total of 10,447 non-synonymous substitutions were found in ...	2012	23413113
periodontal disease and dental caries from krapina neanderthal to contemporary man - skeletal studies.	the aim of this study was the quantification of alveolar bone resorption as well as the number and percentage of teeth with dental caries.	2012	23331387
[neandertal and homo sapiens: to meet, or not to meet?].		2012	23290415
dynamics of genetic and morphological variability within neandertals.	paleogenomics may suggest changes to the way anthropologists have discussed the dynamics and morphological diversity among neandertals. genetic comparisons show that later neandertals had relatively low autosomal genetic variation compared to recent humans. the known mitochondrial sample from neandertals covers a broader geographic and temporal range, and shows greater diversity. this review addresses how genetic data compare to morphological and archaeological evidence about neandertal variatio ...	2012	23274747
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 ...	2012	23959642
hominin dispersal into the nefud desert and middle palaeolithic settlement along the jubbah palaeolake, northern arabia.	the arabian peninsula is a key region for understanding hominin dispersals and the effect of climate change on prehistoric demography, although little information on these topics is presently available owing to the poor preservation of archaeological sites in this desert environment. here, we describe the discovery of three stratified and buried archaeological sites in the nefud desert, which includes the oldest dated occupation for the region. the stone tool assemblages are identified as a midd ...	2012	23185454
nanogp8: evolution of a human-specific retro-oncogene.	nanogp8 is a human (homo sapiens) retrogene, expressed predominantly in cancer cells where its protein product is tumorigenic. it arose through retrotransposition from its parent gene, nanog, which is expressed predominantly in embryonic stem cells. based on identification of fixed and polymorphic variants in a genetically diverse set of human nanog and nanogp8 sequences, we estimated the evolutionary origin of nanogp8 at approximately 0.9 to 2.5 million years ago, more recent than previously es ...	2012	23173096
intrapopulation variation in macro tooth wear patterns--a case study from igloolik, canada.	the pattern of human tooth wear-the way it varies between teeth in the mouth-is crucial to our understanding of important questions in archeology and paleoanthropology, such as the contrasts in diet and behavior between neanderthals and early modern humans in europe and asia, or with the adoption of agriculture in the americas. little is known, however, about the way in which wear patterns develop with increasing age or the way in which they differ between males and females. one explanation is t ...	2012	23125036
archaic human genomics.	for much of the 20th century, the predominant view of human evolutionary history was derived from the fossil record. homo erectus was seen arising in africa from an earlier member of the genus and then spreading throughout the old world and into the oceania. a regional continuity model of anagenetic change from h. erectus via various intermediate archaic species into the modern humans in each of the regions inhabited by h. erectus was labeled the multiregional model of human evolution (mre). a c ...	2012	23124308
extremely rare interbreeding events can explain neanderthal dna in living humans.	considering the recent experimental discovery of green et al that present-day non-africans have 1 to [formula: see text] of their nuclear dna of neanderthal origin, we propose here a model which is able to quantify the genetic interbreeding between two subpopulations with equal fitness, living in the same geographic region. the model consists of a solvable system of deterministic ordinary differential equations containing as a stochastic ingredient a realization of the neutral wright-fisher proc ...	2012	23112810
radiocarbon dates from the grotte du renne and saint-césaire support a neandertal origin for the châtelperronian.	the transition from the middle paleolithic (mp) to upper paleolithic (up) is marked by the replacement of late neandertals by modern humans in europe between 50,000 and 40,000 y ago. châtelperronian (cp) artifact assemblages found in central france and northern spain date to this time period. so far, it is the only such assemblage type that has yielded neandertal remains directly associated with up style artifacts. cp assemblages also include body ornaments, otherwise virtually unknown in the ne ...	2012	23112183
neandertal diet, voracious bacteria and the hunger for knowledge: reply to tomczyk's comments.		2012	23103051
modeling neanderthal clothing using ethnographic analogues.	although direct evidence for neanderthal clothing is essentially nonexistent, information about paleolithic clothing could provide insights into the biological, technological, and behavioral capabilities of neanderthals. this paper takes a new approach to understanding neanderthal clothing through the collection and analysis of clothing data for 245 recent hunter-gatherer groups. these data are tested against environmental factors to infer what clothing humans tend to wear under different condit ...	2012	23084621
new chronology for the middle palaeolithic of the southern caucasus suggests early demise of neanderthals in this region.	neanderthal populations of the southern and northern caucasus became locally extinct during the late pleistocene. the timing of their extinction is key to our understanding of the relationship between neanderthals and anatomically modern humans (amh) in eurasia. recent re-dating of the end of the middle palaeolithic (mp) at mezmaiskaya cave, northern caucasus, and ortvale klde, southern caucasus, suggests that neanderthals did not survive after 39 ka cal bp (thousands of years ago, calibrated be ...	2012	23084367
north african populations carry the signature of admixture with neandertals.	one of the main findings derived from the analysis of the neandertal genome was the evidence for admixture between neandertals and non-african modern humans. an alternative scenario is that the ancestral population of non-africans was closer to neandertals than to africans because of ancient population substructure. thus, the study of north african populations is crucial for testing both hypotheses. we analyzed a total of 780,000 snps in 125 individuals representing seven different north african ...	2012	23082212
cervical and crown outline analysis of worn neanderthal and modern human lower second deciduous molars.	despite the general increase in digital techniques for dental morphometric analyses, only a few methods are available to study worn teeth. moreover, permanent dentitions are studied much more frequently than deciduous teeth. in this study, we address both issues by providing a taxonomic classification of neanderthal and modern human (mh) lower second deciduous molars (dm(2) s) through the analysis of crown and cervical outlines. crown and cervical outlines were obtained from a three-dimensional ...	2012	23077004
the date of interbreeding between neandertals and modern humans.	comparisons of dna sequences between neandertals and present-day humans have shown that neandertals share more genetic variants with non-africans than with africans. this could be due to interbreeding between neandertals and modern humans when the two groups met subsequent to the emergence of modern humans outside africa. however, it could also be due to population structure that antedates the origin of neandertal ancestors in africa. we measure the extent of linkage disequilibrium (ld) in the g ...	2012	23055938
towards the middle palaeolithic in western europe: the case of orgnac 3 (southeastern france).	the sequence of orgnac 3 in southern europe is dated to mis 9 and the beginning of mis 8. the site contains records of upper acheulian occupations with evidence of middle palaeolithic technological strategies at the top of the sequence. in order to address the question of gradual versus punctuated changes in the onset of the middle palaeolithic, nine criteria on subsistence strategies and technological behaviour were selected throughout the whole stratigraphic sequence to describe behavioural pa ...	2012	23040107
long anterior mandibular tooth roots in neanderthals are not the result of their large jaws.	tooth root length has been shown to taxonomically distinguish neanderthals from modern humans. however, this may result from differences in jaw size between both taxa, although most previous studies have revealed a very low or non-existent correlation between tooth size and jaw size in recent modern humans. we therefore investigated, within a broader taxonomical frame, to what extent measurements on the anterior tooth roots and the symphyseal region covary. our samples comprise permanent mandibu ...	2012	23000085
archaeology. did neandertals truly bury their dead?		2012	22997298
middle pleistocene human facial morphology in an evolutionary and developmental context.	neanderthals and modern humans exhibit distinct facial architectures. the patterning of facial morphology of their predecessors, the middle pleistocene humans, is more mosaic showing a mix of archaic and modern morphologies. significant changes in facial size and robusticity occurred throughout pleistocene human evolution, resulting in temporal trends in both facial reduction and enlargement. however, the allometric patterning in facial morphology in archaic humans is not well understood. this s ...	2012	22981042
the mousterian child from teshik-tash is a neanderthal: a geometric morphometric study of the frontal bone.	in the 1930s subadult hominin remains and mousterian artifacts were discovered in the teshik-tash cave in south uzbekistan. since then, the majority of the scientific community has interpreted teshik-tash as a neanderthal. however, some have considered aspects of the morphology of the teshik-tash skull to be more similar to fossil modern humans such as those represented at skhūl and qafzeh, or to subadult upper paleolithic modern humans. here we present a 3d geometric morphometric analysis of th ...	2012	22976750
neandertal mobility and large-game hunting: the exploitation of reindeer during the quina mousterian at chez-pinaud jonzac (charente-maritime, france).	neandertals were effective hunters of large ungulates throughout their geographic and temporal ranges. equipped with this knowledge, researchers in paleoanthropology continue to seek insight on the relationships between hunting and subsistence strategies with other components of the neandertals' niche, such as mobility, site use, and lithic technology. the quina mousterian deposits from the rockshelter site of chez pinaud jonzac (charente-maritime, france; hereafter jonzac) offer an excellent op ...	2012	22951376
unexpectedly many extinct hominins.	recent studies indicate that neanderthal and denisova hominins may have been separate species, while debate continues on the status of homo floresiensis. the decade-long debate between "splitters," who recognize over 20 hominin species, and "lumpers," who maintain that all these fossils belong to just a few lineages, illustrates that we do not know how many extinct hominin species to expect. here, we present probability distributions for the number of speciation events and the number of contempo ...	2012	22946817
hand to mouth in a neandertal: right-handedness in regourdou 1.	we describe and analyze a neandertal postcranial skeleton and dentition, which together show unambiguous signs of right-handedness. asymmetries between the left and right upper arm in regourdou 1 were identified nearly 20 years ago, then confirmed by more detailed analyses of the inner bone structure for the clavicle, humerus, radius and ulna. the total pattern of all bones in the shoulder and arm reveals that regourdou 1 was a right-hander. confirmatory evidence comes from the mandibular inciso ...	2012	22937134
a high-coverage genome sequence from an archaic denisovan individual.	we present a dna library preparation method that has allowed us to reconstruct a high-coverage (30×) genome sequence of a denisovan, an extinct relative of neandertals. the quality of this genome allows a direct estimation of denisovan heterozygosity indicating that genetic diversity in these archaic hominins was extremely low. it also allows tentative dating of the specimen on the basis of "missing evolution" in its genome, detailed measurements of denisovan and neandertal admixture into presen ...	2012	22936568
new foot remains from the gran dolina-td6 early pleistocene site (sierra de atapuerca, burgos, spain).	this paper presents and describes new foot fossils from the species homo antecessor, found in level td6 of the site of gran dolina (sierra de atapuerca, burgos, spain). these new fossils consist of an almost complete left talus (atd6-95) and the proximal three-quarters of a right fourth metatarsal (atd6-124). the talus atd6-95 is tentatively assigned to hominin 10 of the td6 sample, an adult male specimen with which the second metatarsal atd6-70+107 (already published) is also tentatively associ ...	2012	22921478
effect of ancient population structure on the degree of polymorphism shared between modern human populations and ancient hominins.	recent comparisons between anatomically modern humans and ancient genomes of other hominins have raised the tantalizing, and hotly debated, possibility of hybridization. although several tests of hybridization have been devised, they all rely on the degree to which different modern populations share genetic polymorphisms with the ancient genomes of other hominins. however, spatial population structure is expected to generate genetic patterns similar to those that might be attributed to hybridiza ...	2012	22893688
a haplotype at stat2 introgressed from neanderthals and serves as a candidate of positive selection in papua new guinea.	signals of archaic admixture have been identified through comparisons of the draft neanderthal and denisova genomes with those of living humans. studies of individual loci contributing to these genome-wide average signals are required for characterization of the introgression process and investigation of whether archaic variants conferred an adaptive advantage to the ancestors of contemporary human populations. however, no definitive case of adaptive introgression has yet been described. here we ...	2012	22883142
profile: joão zilhão. neandertal champion defends the reputation of our closest cousins.		2012	22879480
comments on soltysiak's paper: "comment: low dental caries rate in neandertals: the result of diet or the oral flora compositions?".	a low frequency of dental caries in neandertal population is still puzzling. many authors stress that the lower frequency of dental caries was related to a meat diet. however, a recent publication in homo - journal comparative human biology presented a new interpretation of dental caries in neandertals. in this article, soltysiak supports the thesis that the lower frequency of caries in the neandertal population from the near east could not be related to the low-sugar diet, but rather to the abs ...	2012	22858155
border cave and the beginning of the later stone age in south africa.	the transition from the middle stone age (msa) to the later stone age (lsa) in south africa was not associated with the appearance of anatomically modern humans and the extinction of neandertals, as in the middle to upper paleolithic transition in western europe. it has therefore attracted less attention, yet it provides insights into patterns of technological evolution not associated with a new hominin. data from border cave (kwazulu-natal) show a strong pattern of technological change at appro ...	2012	22847432
volcanic ash layers illuminate the resilience of neanderthals and early modern humans to natural hazards.	marked changes in human dispersal and development during the middle to upper paleolithic transition have been attributed to massive volcanic eruption and/or severe climatic deterioration. we test this concept using records of volcanic ash layers of the campanian ignimbrite eruption dated to ca. 40,000 y ago (40 ka b.p.). the distribution of the campanian ignimbrite has been enhanced by the discovery of cryptotephra deposits (volcanic ash layers that are not visible to the naked eye) in archaeolo ...	2012	22826222
neandertal humeri may reflect adaptation to scraping tasks, but not spear thrusting.	unique compared with recent and prehistoric homo sapiens, neandertal humeri are characterised by a pronounced right-dominant bilateral strength asymmetry and an anteroposteriorly strengthened diaphyseal shape. remodeling in response to asymmetric forces imposed during regular underhanded spear thrusting is the most influential explanatory hypothesis. the core tenet of the "spear thrusting hypothesis", that underhand thrusting requires greater muscle activity on the right side of the body compare ...	2012	22815742
neanderthal medics? evidence for food, cooking, and medicinal plants entrapped in dental calculus.	neanderthals disappeared sometime between 30,000 and 24,000 years ago. until recently, neanderthals were understood to have been predominantly meat-eaters; however, a growing body of evidence suggests their diet also included plants. we present the results of a study, in which sequential thermal desorption-gas chromatography-mass spectrometry (td-gc-ms) and pyrolysis-gas chromatography-mass spectrometry (py-gc-ms) were combined with morphological analysis of plant microfossils, to identify mater ...	2012	22806252
revisiting dental fluctuating asymmetry in neandertals and modern humans.	previous studies have suggested that neandertals experienced greater physiological stress and/or were less capable of mitigating stress than most prehistoric modern human populations. the current study compares estimates of dental fluctuating asymmetry (dfa) for prehistoric inupiat from point hope alaska, the late archaic, and protohistoric periods from ohio and west virginia, and a modern sample from ohio to neandertals from europe and southwest asia. dfa results from developmental perturbation ...	2012	22791408
the first modern europeans.	the discovery of new human fossil remains is one of the most obvious ways to improve our understanding of the dynamics of human evolution. the reanalysis of existing fossils using newer methods is also crucial, and may lead to a reconsideration of the biological and taxonomical status of some specimens, and improve our understanding of highly debated periods in human prehistory. this is particularly true for those remains that have previously been studied using traditional approaches, with only ...	2012	22781582
hominoid dispersal patterns and human evolution.	recent advances in dna and isotope analyses have allowed tentative reconstructions of dispersal strategies of plio-pleistocene hominins.(1,2) comparing their findings to dispersal patterns of some extant apes and humans suggested groups of related males and unrelated females in neandertals indicating patrilocality(2) and pan-like male philopatry in australopiths.(1) here we review the demographic, ethnographic, and genetic evidence of dispersal patterns in extant apes and humans and compare the ...	2012	22718478
u-series dating of paleolithic art in 11 caves in spain.	paleolithic cave art is an exceptional archive of early human symbolic behavior, but because obtaining reliable dates has been difficult, its chronology is still poorly understood after more than a century of study. we present uranium-series disequilibrium dates of calcite deposits overlying or underlying art found in 11 caves, including the united nations educational, scientific, and cultural organization (unesco) world heritage sites of altamira, el castillo, and tito bustillo, spain. the resu ...	2012	22700921
[progresses on neandertal genomics].	neandertal is our closest known relative and also an archaic hominid reserving the richest fossils. whether the neandertals exchanged their dna with modern human or not is a matter of debate on the modern human origin. the progresses on the mitochondrial and nuclear genomes of neandertals in recent years were reviewed in this paper. recent study has revealed possible genetic contribution of neandertals to the modern human to some extent, which arose the rethinking of modern human origin. the exp ...	2012	22698735
neandertal and denisovan retroviruses.		2012	22677281
specific inactivation of two immunomodulatory siglec genes during human evolution.	sialic acid-recognizing ig-like lectins (siglecs) are signaling receptors that modulate immune responses, and are targeted for interactions by certain pathogens. we describe two primate siglecs that were rendered nonfunctional by single genetic events during hominin evolution after our common ancestor with the chimpanzee. siglec13 was deleted by an alu-mediated recombination event, and a single base pair deletion disrupted the orf of siglec17. siglec-13 is expressed on chimpanzee monocytes, inna ...	2012	22665810
differences between neandertal and modern human infant and child growth models.	studying the emergence of distinctive human growth patterns is essential to understanding the evolution of our species. the large number of neandertal fossils makes this species the best candidate for a comparative study of growth patterns in archaic and modern humans. here, neandertal height growth during infancy and early childhood is described using a mathematical model. height growth velocities for individuals five years old or younger are modelled as age functions based on different estimat ...	2012	22658332
did a discrete event 200,000-100,000 years ago produce modern humans?	scenarios for modern human origins are often predicated on the assumption that modern humans arose 200,000-100,000 years ago in africa. this assumption implies that something 'special' happened at this point in time in africa, such as the speciation that produced homo sapiens, a severe bottleneck in human population size, or a combination of the two. the common thread is that after the divergence of the modern human and neandertal evolutionary lineages ∼400,000 years ago, there was another discr ...	2012	22658331
middle paleolithic human remains from the gruta da oliveira (torres novas), portugal.	additional middle paleolithic human remains from layers 17, 18, and 22 of the gruta da oliveira, portugal consist of a proximal manual phalanx 2 (oliveira 5), a partial postcanine tooth (oliveira 6), a humeral diaphysis (oliveira 7), a distal mandibular molar (oliveira 8), and a mandibular premolar (p(3) ) (oliveira 9). oliveira 5, 6, and 8 are unremarkable for late pleistocene humans. the oliveira 7 right humerus is moderately robust or the individual had the stocky body proportions of other eu ...	2012	22610966
different cranial ontogeny in europeans and southern africans.	modern human populations differ in developmental processes and in several phenotypic traits. however, the link between ontogenetic variation and human diversification has not been frequently addressed. here, we analysed craniofacial ontogenies by means of geometric-morphometrics of europeans and southern africans, according to dental and chronological ages. results suggest that different adult cranial morphologies between southern africans and europeans arise by a combination of processes that i ...	2012	22558270
variations and asymmetries in regional brain surface in the genus homo.	paleoneurology is an important field of research within human evolution studies. variations in size and shape of an endocast help to differentiate among fossil hominin species whereas endocranial asymmetries are related to behavior and cognitive function. here we analyse variations of the surface of the frontal, parieto-temporal and occipital lobes among different species of homo, including 39 fossil hominins, ten fossil anatomically modern homo sapiens and 100 endocasts of extant modern humans. ...	2012	22542169
new human fossil to the last neanderthals in central spain (jarama vi, valdesotos, guadalajara, spain).		2012	22516305
ancient structure in africa unlikely to explain neanderthal and non-african genetic similarity.	neanderthals have been shown to share more genetic variants with present-day non-africans than africans. recent admixture between neanderthals and modern humans outside of africa was proposed as the most parsimonious explanation for this observation. however, the hypothesis of ancient population structure within africa could not be ruled out as an alternative explanation. we use simulations to test whether the site frequency spectrum, conditioned on a derived neanderthal and an ancestral yoruba ...	2012	22513287
genetic adaptation of fatty-acid metabolism: a human-specific haplotype increasing the biosynthesis of long-chain omega-3 and omega-6 fatty acids.	omega-3 and omega-6 long-chain polyunsaturated fatty acids (lc-pufas) are essential for the development and function of the human brain. they can be obtained directly from food, e.g., fish, or synthesized from precursor molecules found in vegetable oils. to determine the importance of genetic variability to fatty-acid biosynthesis, we studied fads1 and fads2, which encode rate-limiting enzymes for fatty-acid conversion. we performed genome-wide genotyping (n = 5,652 individuals) and targeted res ...	2012	22503634
a "copernican" reassessment of the human mitochondrial dna tree from its root.	mutational events along the human mtdna phylogeny are traditionally identified relative to the revised cambridge reference sequence, a contemporary european sequence published in 1981. this historical choice is a continuous source of inconsistencies, misinterpretations, and errors in medical, forensic, and population genetic studies. here, after having refined the human mtdna phylogeny to an unprecedented level by adding information from 8,216 modern mitogenomes, we propose switching the referen ...	2012	22482806
fossil human remains from bolomor cave (valencia, spain).	systematic excavations carried out since 1989 at bolomor cave have led to the recovery of four pleistocene human fossil remains, consisting of a fibular fragment, two isolated teeth, and a nearly complete adult parietal bone. all of these specimens date to the late middle and early late pleistocene (mis 7-5e). the fibular fragment shows thick cortical bone, an archaic feature found in non-modern (i.e. non-homo sapiens) members of the genus homo. among the dental remains, the lack of a midtrigoni ...	2012	22440746
human evolution out of africa: the role of refugia and climate change.	although an african origin of the modern human species is generally accepted, the evolutionary processes involved in the speciation, geographical spread, and eventual extinction of archaic humans outside of africa are much debated. an additional complexity has been the recent evidence of limited interbreeding between modern humans and the neandertals and denisovans. modern human migrations and interactions began during the buildup to the last glacial maximum, starting about 100,000 years ago. by ...	2012	22422974
analysis of human accelerated dna regions using archaic hominin genomes.	several previous comparisons of the human genome with other primate and vertebrate genomes identified genomic regions that are highly conserved in vertebrate evolution but fast-evolving on the human lineage. these human accelerated regions (hars) may be regions of past adaptive evolution in humans. alternatively, they may be the result of non-adaptive processes, such as biased gene conversion. we captured and sequenced dna from a collection of previously published hars using dna from an iberian ...	2012	22412940
predicting homo pigmentation phenotype through genomic data: from neanderthal to james watson.	human pigmentation is regulated by several genes acting at different stages of melanin formation. functional and association studies have elucidated the role of several of these genes in pigmentation phenotypes. forensic and evolutionary studies can benefit from this knowledge.	2012	22411106
right-handedness, lateralization and language in neanderthals: a comment on frayer et al. (2010).		2012	22408126
presumed symbolic use of diurnal raptors by neanderthals.	in africa and western eurasia, occurrences of burials and utilized ocher fragments during the late middle and early late pleistocene are often considered evidence for the emergence of symbolically-mediated behavior. perhaps less controversial for the study of human cognitive evolution are finds of marine shell beads and complex designs on organic and mineral artifacts in early modern human (emh) assemblages conservatively dated to ≈ 100-60 kilo-years (ka) ago. here we show that, in france, neand ...	2012	22403717
the mesosternum of the regourdou 1 neandertal revisited.	fossil hominin mesosterna, while scarce, can provide useful morphological data in addition to rib remains regarding aspects of thoracic size and shape. these data, in turn, can address hypotheses related to respiratory dynamics, climatic adaptation, and ecogeographical patterning. in this study, we re-evaluate the anatomical representation of the mesosternum of the regourdou 1 neandertal individual that alters key aspects of the original description of the fossil remains. we compare this specime ...	2012	22382084
partial genetic turnover in neandertals: continuity in the east and population replacement in the west.	remarkably little is known about the population-level processes leading up to the extinction of the neandertal. to examine this, we use mitochondrial dna sequences from 13 neandertal individuals, including a novel sequence from northern spain, to examine neandertal demographic history. our analyses indicate that recent western european neandertals (<48 kyr) constitute a tightly defined group with low mitochondrial genetic variation in comparison with both eastern and older (>48 kyr) european nea ...	2012	22362080
variation in enamel thickness within the genus homo.	recent humans and their fossil relatives are classified as having thick molar enamel, one of very few dental traits that distinguish hominins from living african apes. however, little is known about enamel thickness in the earliest members of the genus homo, and recent studies of later homo report considerable intra- and inter-specific variation. in order to assess taxonomic, geographic, and temporal trends in enamel thickness, we applied micro-computed tomographic imaging to 150 fossil homo tee ...	2012	22361504
tooth wear, neanderthal facial morphology and the anterior dental loading hypothesis.	the anterior dental loading hypothesis states that the unique neanderthal facial and dental anatomy was an adaptive response to the regular application of heavy forces resulting from both the masticatory and cultural use of the anterior teeth. heavy anterior tooth wear frequently observed in neanderthal specimens is cited as a main source of evidence for heavy forces being applied to these teeth. from this, it might be predicted that the wear shown on the anterior teeth of neanderthals would gre ...	2012	22341317
early pleistocene human humeri from the gran dolina-td6 site (sierra de atapuerca, spain).	in this report, we present a morphometric comparative study of two early pleistocene humeri recovered from the td6 level of the gran dolina cave site in sierra de atapuerca, northern spain. atd6-121 belongs to a child between 4 and 6 years old, whereas atd6-148 corresponds to an adult. atd6-148 exhibits the typical pattern of the genus homo, but it also shows a large olecranon fossa and very thin medial and lateral pillars (also present in atd6-121), sharing these features with european middle p ...	2012	22328492
an ancestral mir-1304 allele present in neanderthals regulates genes involved in enamel formation and could explain dental differences with modern humans.	genetic changes in regulatory elements are likely to result in phenotypic effects that might explain population-specific as well as species-specific traits. micrornas (mirnas) are posttranscriptional repressors involved in the control of almost every biological process. these small noncoding rnas are present in various phylogenetic groups, and a large number of them remain highly conserved at the sequence level. microrna-mediated regulation depends on perfect matching between the seven nucleotid ...	2012	22319171
use of red ochre by early neandertals.	the use of manganese and iron oxides by late neandertals is well documented in europe, especially for the period 60-40 kya. such finds often have been interpreted as pigments even though their exact function is largely unknown. here we report significantly older iron oxide finds that constitute the earliest documented use of red ochre by neandertals. these finds were small concentrates of red material retrieved during excavations at maastricht-belvédère, the netherlands. the excavations exposed ...	2012	22308348
extension of cortical synaptic development distinguishes humans from chimpanzees and macaques.	over the course of ontogenesis, the human brain and human cognitive abilities develop in parallel, resulting in a phenotype strikingly distinct from that of other primates. here, we used microarrays and rna-sequencing to examine human-specific gene expression changes taking place during postnatal brain development in the prefrontal cortex and cerebellum of humans, chimpanzees, and rhesus macaques. we show that the most prominent human-specific expression change affects genes associated with syna ...	2012	22300767
why levallois? a morphometric comparison of experimental 'preferential' levallois flakes versus debitage flakes.	middle palaeolithic stone artefacts referred to as 'levallois' have caused considerable debate regarding issues of technological predetermination, cognition and linguistic capacities in extinct hominins. their association with both neanderthals and early modern humans has, in particular, fuelled such debate. yet, controversy exists regarding the extent of 'predetermination' and 'standardization' in so-called 'preferential levallois flakes' (plfs).	2012	22291888
comment on "late mousterian persistence near the arctic circle".	slimak et al. (reports, 13 may 2011, p. 841) reanalyzed the lithic assemblage from the northern site of byzovaya (russia) and concluded that it was mousterian and produced by neandertals. the previous interpretation of this assemblage as falling within early upper paleolithic variability remains the most parsimonious explanation; pending additional fossil discoveries, there is no evidence supporting the occurrence of neandertals at these high latitudes.	2012	22246757
a uniquely modern human pattern of endocranial development. insights from a new cranial reconstruction of the neandertal newborn from mezmaiskaya.	the globular braincase of modern humans is distinct from all fossil human species, including our closest extinct relatives, the neandertals. such adult shape differences must ultimately be rooted in different developmental patterns, but it is unclear at which point during ontogeny these group characteristics emerge. here we compared internal shape changes of the braincase from birth to adulthood in neandertals (n = 10), modern humans (n = 62), and chimpanzees (n = 62). incomplete fossil specimen ...	2012	22221766
can we still hear the cro-magnon man?	this paper is an essay to connect with the stone age coastal hunters who sheltered 28,000 years ago in the caves of today's italian mediterranean sea shore cliffs. we have focused on the archaeological gravettian layer in the cavillon cave from grimaldi occupied by cro magnons (informal name for anatomically modern humans of the european upper paleolithic), which demonstrates the technical skills of the phase. cro magnons, like neanderthals, were seasonally nomadic; however, the diversity of the ...	2012	23560760
birds of a feather: neanderthal exploitation of raptors and corvids.	the hypothesis that neanderthals exploited birds for the use of their feathers or claws as personal ornaments in symbolic behaviour is revolutionary as it assigns unprecedented cognitive abilities to these hominins. this inference, however, is based on modest faunal samples and thus may not represent a regular or systematic behaviour. here we address this issue by looking for evidence of such behaviour across a large temporal and geographical framework. our analyses try to answer four main quest ...	2012	23029321
[adaptive evolution of the homo mitochondrial genome].	adaptive evolution of 12 protein-coding mitochondrial genes in members of genus homo (denisova hominin (h. sp. altai), neandertals (h. neanderthalensis) and modern humans (h. sapiens)) has been evaluated by assessing the pattern of changes in the physicochemical properties of amino acid replacements during the primate evolution. it has been found that in the homo molecular adaptation (positive destabilizing selection) become apparent in the form of 12 radical amino acid replacements accompanied ...	2012	22393781
generation times in wild chimpanzees and gorillas suggest earlier divergence times in great ape and human evolution.	fossils and molecular data are two independent sources of information that should in principle provide consistent inferences of when evolutionary lineages diverged. here we use an alternative approach to genetic inference of species split times in recent human and ape evolution that is independent of the fossil record. we first use genetic parentage information on a large number of wild chimpanzees and mountain gorillas to directly infer their average generation times. we then compare these gene ...	2012	22891323
a geometric morphometric analysis of hominin upper second and third molars, with particular emphasis on european pleistocene populations.	the study of dental morphology by means of geometric morphometric methods allows for a detailed and quantitative comparison of hominin species that is useful for taxonomic assignment and phylogenetic reconstruction. upper second and third molars have been studied in a comprehensive sample of plio- and pleistocene hominins from african, asian and european sites in order to complete our analysis of the upper postcanine dentition. intraspecific variation in these two molars is high, but some inters ...	2012	22840714
the status of homo heidelbergensis (schoetensack 1908).	the species homo heidelbergensis is central to many discussions about recent human evolution. for some workers, it was the last common ancestor for the subsequent species homo sapiens and homo neanderthalensis; others regard it as only a european form, giving rise to the neanderthals. following the impact of recent genomic studies indicating hybridization between modern humans and both neanderthals and "denisovans", the status of these as separate taxa is now under discussion. accordingly, clari ...	2012	22718477
fission-fusion and the evolution of hominin social systems.	the course of hominin evolution has involved successive migrations towards higher absolute latitudes over the past three million years. poorer habitat quality further from the equator has led to the necessity for groups occupying higher latitudes to live at lower population densities. coupled with a trend towards increasing group size over this time period, this tendency towards expansion has led to exponential increases in the area requirements of hominin groups, and a concomitant need to adjus ...	2012	22197359