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The ∼25 m-thick Manyara Beds section near Makuyuni, northern Tanzania, contain abundant Acheulean lithics and vertebrate fossils, including possible Homo erectus remains. However, the age of the unit (and its most productive fossil and archaeological localities) has been provisional. To address this we measured 20 new stratigraphic sections and collected approximately 30 tephra and volcanic rock samples for 40Ar/39Ar geochronology and 12 mudstone/siltstone samples for a trial paleomagnetic analysis. Most sampled tephra deposits (from the greenish-gray, lacustrine lower member) proved too fine-grained or lithologically heterogeneous for 40Ar/39Ar analysis, but crystals from a pumice-rich unit (the ‘Hollywood Tuff;’ at the base of the reddish-brown, fluvial upper member), analyzed in three batches, yielded a middle Pleistocene 40Ar/39Ar age of 0.633 ± 0.039 Ma. Two-step analysis of 10 feldspars from a nephelinite underlying the Manyara Beds produced a weighted mean 40Ar/39Ar age of 6.00 ± 0.03 Ma, linking it to the nearby stratovolcano Essimingor. Paleomagnetic analysis showed that the Manyara Beds carry a stable ancient remanence, and characteristic directions obtained by AF demagnetization define a preliminary magnetostratigraphy for the unit. Four magnetozones over 21 m of section include two intervals of normal polarity tentatively correlated with the Jaramillo Subchron and Brunhes Chron. The Matuyama-Brunhes boundary is located within the upper half of the lower member, at least 6 m below the lower-upper member contact. Using undecompacted sedimentation rates of 24–46 m/Ma (calculated using geomagnetic polarity time scale correlations and the stratigraphic position of the ∼0.633 Ma tephra) we estimate that the base of the Manyara Beds was deposited between 0.98 and 1.3 Ma and the top between 0.27 and 0.44 Ma. Beds III–IV and the Masek Beds at Olduvai were probably also deposited in the same early–middle Pleistocene interval, during which the Mid-Pleistocene Revolution climatic event and active rifting and volcanism in the southernmost Gregory Rift jointly contributed to landscape and ecosystem instability.

The nasal cavity is essential for humidifying and warming the air before it reaches the sensitive lungs. Because humans inhabit environments that can be seen as extreme from the perspective of respiratory function, nasal cavity shape is expected to show climatic adaptation. This study examines the relationship between modern human variation in the morphology of the nasal cavity and the climatic factors of temperature and vapor pressure, and tests the hypothesis that within increasingly demanding environments (colder and drier), nasal cavities will show features that enhance turbulence and air-wall contact to improve conditioning of the air. We use three-dimensional geometric morphometrics methods and multivariate statistics to model and analyze the shape of the bony nasal cavity of 10 modern human population samples from five climatic groups. We report significant correlations between nasal cavity shape and climatic variables of both temperature and humidity. Variation in nasal cavity shape is correlated with a cline from cold-dry climates to hot-humid climates, with a separate temperature and vapor pressure effect. The bony nasal cavity appears mostly associated with temperature, and the nasopharynx with humidity. The observed climate-related shape changes are functionally consistent with an increase in contact between air and mucosal tissue in cold-dry climates through greater turbulence during inspiration and a higher surface-to-volume ratio in the upper nasal cavity.

In1978,twowell-preservedfossilhumancraniawerediscoveredat the Apidima site, a karstic cave complex in the Mani peninsula(southernPeloponnese;Fig.1),duringexcavationsbytheUniversityof Athens Medical School (Pitsios, 1995; Harvati and Delson, 1999;Harvati et al., 2009). Together with the Petralona cranium, the Api-dima specimens represent the most important paleoanthropolog-ical finds in Greece, and some of the most significant human fossildiscoveries in south-eastern Europe, a region whose paleoanthro-pological record remains relatively unexplored. However, despitetheir importance, these specimens remain obscure; little has beenpublished about them in the international literature, and detaileddescription and documentation are still pending.Ofthetwospecimens,Apidima2isthebetterpreserved,thoughsomewhat distorted and showing multiple cracks (Fig. 2A). Theoccipital bone, parts of the temporals, as well as all of the teeth andpartofthepalate,areallabsent.Afragmentof therightmandibularramus ispreserved, attachedtotheencasingmatrixand cranium inroughly anatomical position. Apidima 2 has been described asshowing an elongated, low vault with a pronounced supra-orbitaltorus, a wide interorbital breadth, large rounded orbits, no caninefossa, a large nasal aperture, and a prognathic face (Harvati andDelson, 1999; Geanacos, 2001; Pitsios, 2002; Harvati et al., 2009).It is relatively gracile, and has been considered a female (Harvatiand Delson, 1999; Pitsios, 2002; Harvati et al., 2009). Neanderthalaffinities have been noted and the specimen has been proposed torepresent a pre-Neanderthal. Of particular interest is its possiblerelationship with the Petralona cranium from Northern Greece(Fig.2B),andthesuggestionthatApidima2mayrepresentafemaleHomo heidelbergensis comparable to Petralona in geological age.Apidima 1 is less complete and preserves only the posterior part ofthe neuro- and basicranium.Attempts at identifying Apidima 2 to taxon have been stymiedby the lack of both detailed description and extensive metric data.On the basis of its general morphology, Pitsios (1999) suggestedthat Apidima 2 lies on the Neanderthal lineage and proposed thename Homo (sapiens) teanarius (Pitsios, 1995). Harvati and Delson(1999) suggested affinities with H. heidelbergensis (s.l.). Nineosteometric measurements published in a forensic report byKoutselinis et al. (1995) have been used as variables in previousmetric analyses by Manolis (1996) and, more recently, by Harvatiet al. (2009). Both studies were inconclusive in their results,placing Apidima 2 either in an area of overlap between H. hei-delbergensis (s.l.) and early anatomically modern humans (Manolis,1996), or between Homo neanderthalensis and H. heidelbergensis(s.l.) (Harvati et al., 2009).Given the importance of the Apidima specimens and the smallamount of information on them that currently exists, we re-examine the metric data published for Apidima 2. We 1) test theoriginal nine reported measurements for possible error associatedwith distortion; 2) correct the measurements for size, so that wecan examine shape differences; and 3) take into account as manymeasurements and Middle-Late Pleistocene specimens as possible,inordertoachievethebestresolutionforclassifyingApidima2.Wereview the geology and chronology of the site in light of recent re-evaluation of regional geomorphologyand discuss the implicationsof our findings for the interpretation of the Apidima remains.Materials and methodsOurcomparativesamplecomprisesnineteenfossilhumancrania(Table 1) on which all of the variables known for Apidima 2 can bemeasured (Table 2). Most data were collected by one of us (CS),although some were taken from the literature (Trinkaus, pers.comm.,forShanidar1;Arsuaga etal.,1997).Forthepurposesofouranalyses, we grouped the European Middle Pleistocene specimens

The classification and phylogenetic relationships of the middle Pleistocene human fossil record remains one of the most intractable problems in paleoanthropology. Several authors have noted broad resemblances between European and African fossils from this period, suggesting a single taxon ancestral to both modern humans and Neanderthals. Others point out 'incipient' Neanderthal features in the morphology of the European sample and have argued for their inclusion in the Neanderthal lineage exclusively, following a model of accretionary evolution of Neanderthals. We approach these questions using geometric morphometric methods which allow the intuitive visualization and quantification of features previously described qualitatively. We apply these techniques to evaluate proposed cranio-facial 'incipient' facial, vault, and basicranial traits in a middle-late Pleistocene European hominin sample when compared to a sample of the same time depth from Africa. Some of the features examined followed the predictions of the accretion model and relate the middle Pleistocene European material to the later Neanderthals. However, although our analysis showed a clear separation between Neanderthals and early/recent modern humans and morphological proximity between European specimens from OIS 7 to 3, it also shows that the European hominins from the first half of the middle Pleistocene still shared most of their cranio-facial architecture with their African contemporaries.

In the originally published version of this Letter, the x axis in Fig. 3a should have been: ‘PC1: 26%’ rather than ‘PC1: 46%’, and the y axis should have been: ‘PC2: 16%’ rather than ‘PC2: 29%’. We also noticed an error in the numbering of the fossils from Qafzeh: Qafzeh 27 should be removed, and Qafzeh 26 is actually Qafzeh 25, following Tillier (2014)1 and Schuh et al. (2017)2 and personal communication with B. Vandermeersch and M. D. Garralda. The correct enumeration of Qafzeh samples in the ‘Mandibular metric data’ section of the Methods is therefore: ‘Qafzeh (9, 25)’ rather than ‘Qafzeh (9, 26, 27)’. Owing to the removal of Qafzeh 27, the convex hull of early modern humans changes slightly in Extended Data Fig. 1c. The sample sizes in Extended Data Fig. 1c should have read: Middle Pleistocene archaic Homo n = 19 (instead of 11), Neanderthals n = 40 (instead of 41), early modern humans n = 12 (instead of 7), and recent modern humans n = 46 (instead of 48). In Extended Data Table 2, the mean and standard deviation of corpus height and breadth at mental foramen for early modern humans should have been: x = 33.15, σ = 3.26 for height (rather than x = 34.23, σ = 4.57); and x = 16.25, σ = 1.28 for breadth (rather than x = 16.04, σ = 1.75). Accordingly, n = 12 (rather than n = 13) for both breadth and height. These errors have been corrected in the Letter online (the original Extended Data Fig. 1 is shown in Supplementary Information to this Amendment). These changes do not alter any inferences drawn from the data.

Since the discovery of the human remains from the Upper Cave of Zhoukoudian in the 1930s there has been speculation over their affinities. In particular, the degree to which the three adult crania exhibit recent East Asian morphology, as well as their degree of within-group variability, has long been debated. Several early researchers described a resemblance to East Asian populations, but these findings have been for the most part rejected by more recent metric and non-metric analyses. Nevertheless, the Upper Cave specimens have not been classified conclusively into any recent modern human population to which they have been compared, and classification results differ for each cranium. Here, the question of the affinities of Upper Cave 101 and 103, the two better-preserved crania, is examined from the perspective of the Late Pleistocene human fossil record using the methodology of 3-D geometric morphometrics. The degree of morphological variation between the two specimens is also evaluated within the context of recent population variability. Neurocranial and facial morphology are analyzed separately so as to maximize comparative samples. Results show a morphological resemblance of the Upper Cave material to Upper Paleolithic Europeans. It is proposed that the Upper Cave specimens retain important aspects of modern human ancestral morphology, and possibly share a recent common ancestral population with Upper Paleolithic Europeans, in accordance with the Single Origin model of modern human origins.

The ability of cranial morphology to reflect population/phylogenetic history, and the degree to which it might be influenced by environmental factors and selection pressures have been widely discussed. Recent consensus views cranial morphology as largely indicative of population history in humans, with some anatomical cranial regions/measurements being more informative on population history, while others being under selection pressure. We test earlier findings using the largest and most diverse cranial dataset available as yet: 7,423 male specimens from 135 geographic human population samples represented by 33 standard craniometric linear measurements. We calculated Mahalanobis D2 for three datasets: complete cranial dataset; facial measurement dataset; and neurocranial measurement dataset; these morphological distance matrices were then compared to matrices of geographic distances as well as of several climatic variables. Additionally, we calculated Fst values for our cranial measurements and compared the results to the expected Fst values for neutral genetic loci. Our findings support the hypothesis that cranial, and especially neurocranial morphology, is phylogenetically informative, and that aspects of the face and cranium are subject to selection related to climatic factors. The Fst analysis suggest that selection to climate is largely restricted to groups living in extremely cold environments, including Northeast Asia, North America, and Northern Europe, though each of these regions appears to have arrived at their morphology through distinct adaptive pathways. Anat Rec, 2009. © 2009 Wiley-Liss, Inc.

We report here the first results of a method to extract and sequence mature enamel proteins from modern human and Neanderthal tooth enamel. Using MALDI-TOF/TOF mass spectrometry and a combination of direct sequencing and peptide mass mapping we have sequenced a peptide from the tyrosine-rich amelogenin peptide (TRAP) of the X isoform of the amelogenin protein for modern and recent human samples. We also report our results from two Neanderthal enamel samples where we were also able to recover fragments of the TRAP protein, which had a similar sequence to the modern human samples.

European paleoanthropology and paleolithic archeology were already well-established by the early twentieth century. The human fossil record from this continent is the longest known and perhaps most intensively studied. Nonetheless, important gaps remain to this day in the map of Pleistocene Europe; perhaps the most glaring of these is located in the southeastern corner of the continent. This region's record is critical for addressing questions about the course of human evolution in Europe because its geographic position lends it a dual role: on one hand, it encompasses a frequently hypothesized dispersal corridor from Africa into Europe for both archaic and early modern humans; on the other, as one of the three Mediterranean peninsulas, it acted as a refugium for plant, animal, and, most likely, human populations during glacial conditions. This article is a review of the paleoanthopological record of Greece, one of the least known in Europe.

Developmental and structural affinities between modern human and Neanderthal dental remains continue to be a subject of debate as well as their utility for informing assessments of life history and taxonomy. Excavation of the Middle Paleolithic cave site Lakonis in southern Greece has yielded a lower third molar (LKH 1). Here, we detail the crown development and enamel thickness of the distal cusps of the LKH 1 specimen, which has been classified as a Neanderthal based on the presence of an anterior fovea and mid-trigonid crest. Crown formation was determined using standard histological techniques, and enamel thickness was measured from a virtual plane of section. Developmental differences include thinner cuspal enamel and a lower periodicity than modern humans. Crown formation in the LKH 1 hypoconid is estimated to be 2.6-2.7 years, which is shorter than modern human times. The LKH 1 hypoconid also shows a more rapid overall crown extension rate than modern humans. Relative enamel thickness was approximately half that of a modern human sample mean; enamel on the distal cusps of modern human third molars is extremely thick in absolute and relative terms. These findings are consistent with recent studies that demonstrate differences in crown development, tissue proportions, and enamel thickness between Neanderthals and modern humans. Although overlap in some developmental variables may be found, the results of this and other studies suggest that Neanderthal molars formed in shorter periods of time than modern humans, due in part to thinner enamel and faster crown extension rates.

We report here direct evidence for Neanderthal mobility through the measurement of strontium isotope ratios in tooth enamel using laser-ablation, which allows us to use much smaller samples than traditional methods. There has been a long-standing debate over the extent of Neanderthal mobility, with some arguing for Neanderthals having a very limited geographic range and others for more substantial, and even seasonal, lifetime movements. We sampled across the enamel of a Neanderthal third molar from the site of Lakonis, Greece, dating to ca. 40,000 years ago. The tooth was found in a coastal limestone cave, yet the strontium isotope values indicate the enamel was formed while the individual resided in a region with bedrock consisting of older (more radiogenic) volcanic bedrock. Therefore, this individual must have lived in a different (more radiogenic) location during this period of third molar crown formation (likely to be between the ages of 7 and 9 years) than where the tooth was found. This strontium isotope evidence therefore indicates that this Neanderthal moved over a relatively wide (i.e. at least 20 km) geographical range in their lifetime.

Enamel thickness and dental tissue proportions have been recognized as effective taxonomic discriminators between Neanderthal and modern humans teeth. However, most of the research on this topic focused on permanent teeth, and little information is available for the deciduous dentition. Moreover, although worn teeth are more frequently found than unworn teeth, published data for worn teeth are scarce and methods for the assessment of their enamel thickness need to be developed. Here, we addressed this issue by studying the 2D average enamel thickness (AET) and 2D relative enamel thickness (RET) of Neanderthal and modern humans unworn to moderately worn upper first deciduous molars (dm(1)s) and upper second deciduous molars (dm(2)s). In particular, we used 3D μCT data to investigate the mesial section for dm(1)s and both mesial and buccal sections for dm(2)s. Our results confirmed previous findings of an Neanderthal derived condition of thin enamel, and thinner enamel in dm(1)s than dm(2)s in both Neanderthal and modern humans. We demonstrated that the Neanderthal 2D RET indices are significantly lower than those of modern humans at similar wear stages in both dm(1)s and dm(2)s (p < 0.05). The discriminant analysis showed that using 2D RET from dm(1) and dm(2) sections at different wear stages up to 93% of the individuals are correctly classified. Moreover, we showed that the dm(2) buccal sections, although non-conventionally used, might have an advantage on mesial sections since they distinguish as well as mesial sections but tend to be less worn. Therefore, the 2D analysis of enamel thickness is suggested as a means for taxonomic discrimination between modern humans and Neanderthal unworn to moderately worn upper deciduous molars.

The hominin fossil record of Romania comprises some of the earliest and best preserved early modern humans in Europe. As such, these fossils play an important role in our understanding of the timing of the modern human arrival in Europe, their local evolution, and their potential interactions with Neanderthal populations. In this chapter, we briefly review the Romanian human fossil record and present new 3D geometric morphometric analyses of the mandibular remains from Oase and Muierii. Our findings are discussed in the context of admixture between Neanderthals and modern humans and in light of paleogenetic results (Fu et al. 2015) indicating recent Neanderthal ancestry for Oase 1.

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The timing and route of early human dispersals out of the African continent are among the most important issues currently discussed in paleoanthropology. Several questions arise concerning both early and later dispersals: When did migration events happen? From which populations did these dispersing hominins stem? Which routes did they use? One of the likely dispersal corridors passes through Turkey, which is situated between three continents and therefore can be seen as an important bridge between them. Despite its geographic position, paleoanthropological research in Turkey has been limited, and the known fossil human record from this region is small. Although most of the known fossil human remains were found during early investigations, in the last decade new finds have further highlighted the region’s potential for paleoanthropological research. This chapter reviews the human fossil record from Turkey, and presents the results of a preliminary geometric morphometric study of the Kocabas hominin, the oldest and most important fossil human specimen known from the country.

Greece lies at the crossroads between Europe, Asia, and Africa, and represents a logical gateway through which early human populations might have repeatedly passed on the way to and from Europe. It also represents one of the three European Mediterranean peninsulas which acted as a refugium for fauna, flora and, very likely, human populations during glacial times. Evidence from this region is therefore essential in order to test hypotheses about the course of human evolution in Europe. Despite the importance of the region, paleoanthropological research has until recently been relatively neglected. In recent years, however, renewed research efforts have produced new human fossils from Greece, recovered from excavated contexts. This chapter reviews the Greek human fossil evidence in the context of broader questions in European paleoanthropology.

Modern humans arrived in Europe ∼45,000 years ago, but little is known about their genetic composition before the start of farming ∼8,500 years ago. Here we analyse genome-wide data from 51 Eurasians from ∼45,000-7,000 years ago. Over this time, the proportion of Neanderthal DNA decreased from 3-6% to around 2%, consistent with natural selection against Neanderthal variants in modern humans. Whereas there is no evidence of the earliest modern humans in Europe contributing to the genetic composition of present-day Europeans, all individuals between ∼37,000 and ∼14,000 years ago descended from a single founder population which forms part of the ancestry of present-day Europeans. An ∼35,000-year-old individual from northwest Europe represents an early branch of this founder population which was then displaced across a broad region, before reappearing in southwest Europe at the height of the last Ice Age ∼19,000 years ago. During the major warming period after ∼14,000 years ago, a genetic component related to present-day Near Easterners became widespread in Europe. These results document how population turnover and migration have been recurring themes of European prehistory.

Sub-fossilised remains may still contain highly degraded ancient DNA (aDNA) useful for palaeogenetic investigations. Whether X-ray computed [micro-] tomography ([μ]CT) imaging of these fossils may further damage aDNA remains debated. Although the effect of X-ray on DNA in living organisms is well documented, its impact on aDNA molecules is unexplored.Here we investigate the effects of synchrotron X-ray irradiation on aDNA from Pleistocene bones. A clear correlation appears between decreasing aDNA quantities and accumulating X-ray dose-levels above 2000 Gray (Gy). We further find that strong X-ray irradiation reduces the amount of nucleotide misincorporations at the aDNA molecule ends. No representative effect can be detected for doses below 200 Gy. Dosimetry shows that conventional μCT usually does not reach the risky dose level, while classical synchrotron imaging can degrade aDNA significantly. Optimised synchrotron protocols and simple rules introduced here are sufficient to ensure that fossils can be scanned without impairing future aDNA studies.

Unlike other catarrhines, colobines show early molar eruption relative to that of the anterior dentition. The pattern is variable, with Asian genera (Presbytina) showing a greater variability than the African genera (Colobina). The polarity of early relative molar eruption, as well as the degree to which it is related to phylogeny, are unclear. Schultz (1935) suggested that the trend reflects phylogeny and is primitive for catarrhines. More recently, however, researchers have proposed that life history and dietary hypotheses account for early relative molar eruption. If the colobine eruption pattern is primitive for catarrhines, it implies that cercopithecines and hominoids converged on delayed relative molar eruption. Alternatively, if the colobine condition is derived, factors such as diet and mortality patterns probably shaped colobine eruption patterns. Here we update our knowledge on eruption sequences of living colobines, and explore the evolutionary history of the colobine dental eruption pattern by examining fossil colobine taxa from Eurasia (Mesopithecus) and Africa (Kuseracolobus aramisi and Colobus sp.) and the basal cercopithecoid Victoriapithecus macinnesi. We scored specimens per Harvati (2000). The Late Miocene-Early Pliocene Mesopithecus erupts the second molar early relative to the incisors, while the Early Pliocene Kuseracolobus aramisi does not. These results demonstrate that the common colobine tendency for early molar eruption relative to the anterior dentition had appeared by the Late Miocene, and that some of the diversity observed among living colobines was already established in the Late Miocene/Early Pliocene. We discuss the implications of these results for phylogenetic, life history, and dietary hypotheses of dental development.

The occipital bun (‘‘chignon’’) is cited widely as a Neanderthal derived trait. It encompasses the posterior projection/convexity of the occipital squama and is associated with lambdoid flattening on the parietal. A ‘hemibun’ in some Upper Paleolithic Europeans is thought by some authors to indicate interbreeding between Neanderthals and early modern Europeans. However, ‘bunning’ is difficult to measure, and the term has been applied to a range of morphological patterns. Furthermore, its usefulness in phylogenetic reconstruction and its homologous status across modern and fossil humans have been disputed. We present a geometric morphometric study that quantitatively evaluates the chignon, assesses its usefulness in separating Neanderthals from modern humans, and its degree of similarity to Upper Paleolithic ‘hemibuns.’ We measured the three-dimensional coordinates of closely spaced points along the midsagittal plane from bregma to inion and of anatomical landmarks in a large series of recent human crania and several Middle and Late Pleistocene European and African fossils. These coordinate data were processed using the techniques of geometric morphometrics and analyzed with relative warps, canonical variates, and singular warps. Our results show no separation between Neanderthals and modern humans, including early modern Europeans, when the shape of the occipital plane midsagittal-profile is considered alone. On the other hand, Neanderthals are well separated from both recent and fossil modern humans when information about the occipital’s relative position and relative size are also included. Furthermore, the occurrence of a highly convex and posteriorly projecting midline occipital profile (interpreted as the occipital bun) is highly correlated (>0.8) with a flat parietal midsagittal profile and with antero-superiorly positioned temporal bones across both our recent and our fossil human samples. We conclude that the shape of the occipital profile alone should not be considered an independent trait, as it is very tightly integrated with braincase shape. Our analysis does not support differences in integration of the posterior midsagittal profile and the cranial base in Pleistocene and recent humans. 2006 Elsevier Ltd. All rights reserved.

The morphology of the human bony labyrinth is thought to preserve a strong phylogenetic signal and to be minimally, if at all, affected by postnatal processes. The form of the semicircular canals is considered a derived feature of Neanderthals and different from the modern human anatomy. Among other hominins, European Middle Pleistocene humans have been found to be most similar to Neanderthals. Early modern humans have been proposed to show a pattern that is distinct, but most similar to that of Holocene people. Here we examine the inner ear structures of the Cioclovina calvaria, one of the earliest reliably dated and relatively complete modern human crania from Europe, in the context of recent and fossil human variation.Bony labyrinths were virtually extracted from CT scans of recent Europeans and Cioclovina. Using univariate and multivariate methods, measurements of the semicircular canals were compared with published measurements of other fossil specimens.Our results show that Cioclovina's inner ear morphology falls within the range of modern variation, with affinities to both Late Pleistocene modern humans and recent Europeans. Using discriminant functions, the sex of the Cioclovina specimen is estimated as male.Results agree with previous work showing that Cioclovina exhibits fully modern cranial morphology.

The appearance of anatomically modern humans in Europe and the nature of the transition from the Middle to Upper Palaeolithic are matters of intense debate. Most researchers accept that before the arrival of anatomically modern humans, Neanderthals had adopted several ‘transitional’ technocomplexes. Two of these, the Uluzzian of southern Europe and the Châtelperronian of western Europe, are key to current interpretations regarding the timing of arrival of anatomically modern humans in the region and their potential interaction with Neanderthal populations. They are also central to current debates regarding the cognitive abilities of Neanderthals and the reasons behind their extinction1, 2, 3, 4, 5, 6. However, the actual fossil evidence associated with these assemblages is scant and fragmentary7, 8, 9, 10, and recent work has questioned the attribution of the Châtelperronian to Neanderthals on the basis of taphonomic mixing and lithic analysis11, 12. Here we reanalyse the deciduous molars from the Grotta del Cavallo (southern Italy), associated with the Uluzzian and originally classified as Neanderthal13, 14. Using two independent morphometric methods based on microtomographic data, we show that the Cavallo specimens can be attributed to anatomically modern humans. The secure context of the teeth provides crucial evidence that the makers of the Uluzzian technocomplex were therefore not Neanderthals. In addition, new chronometric data for the Uluzzian layers of Grotta del Cavallo obtained from associated shell beads and included within a Bayesian age model show that the teeth must date to ~45,000–43,000 calendar years before present. The Cavallo human remains are therefore the oldest known European anatomically modern humans, confirming a rapid dispersal of modern humans across the continent before the Aurignacian and the disappearance of Neanderthals., This work was supported by the NSF 01-120 Hominid Grant 2007, A.E.R.S. Dental Medicine Organisations GmbH FA547013, the Fondation Fyssen, the DFG INST 37/706-1 FUGG and the NERC Grant (NE/D014077/1).

The temporal bone is the location of sev- eral traits thought to differentiate Neanderthals from modern humans, including some proposed Neanderthal- derived traits. Most of these, however, are difficult to measure and are usually described qualitatively. This study applied the techniques of geometric morphometrics to the complex morphology of the temporal bone, in order to quantify the differences observed between Neanderthal and modern human anatomy. Two hundred and seventy modern human crania were measured, representing 9 populations of 30 individuals each, and spanning the ex- tremes of the modern human geographical range. Twelve Neanderthal specimens, as well as Reilingen, Kabwe, Skhul 5, Qafzeh 9, and 4 Late Paleolithic European spec- imens, were included in the fossil sample. The data were collected in the form of three-dimensional (3-D) landmark coordinates, and specimen configurations were superim- posed using generalized Procrustes analysis. The fitted coordinates were then analyzed by an array of multivari- ate statistical methods, including principal components analysis, canonical variates analysis, and Mahalanobis D 2 . The temporal bone landmark analysis was very suc- cessful in separating Neanderthals from modern humans. Neanderthals were separated from modern humans in both the principal components and canonical variates analyses. They were much further in Mahalanobis dis- tances from all modern human populations than any two modern human groups were from each other. Most of the previously described temporal bone traits contributed to this separation. Am J Phys Anthropol 120:323-338, 2003. © 2003 Wiley-Liss, Inc.

The Neanderthal taxonomic position is a matter of wide disagreement among paleoanthropologists. Some workers consider this fossil human group to represent a different species, Homo neanderthalensis, while others see it as a subspecies of Homo sapiens. This study developed two models of morphological variation to be applied to a comparison between Neanderthals and modern humans: modern human populations provided a measure of intra-specific variation, while the species and subspecies of Pan provided measures of both intra- and inter-specific morphological differences. Although such an approach has been advocated strongly, it has not been systematically undertaken until recently. The techniques of geometric morphometrics were used to collect data in the form of three-dimensional coordinates of craniofacial landmarks. The data were processed using generalized procrustes analysis, and analyzed by an array of multivariate statistical methods, including principal components analysis, canonical variates analysis and Mahalanobis D 2 . The morphological distances between Neanderthals and modern humans, and between Neanderthals and Late Paleolithic/early anatomically modern specimens, are consistently greater than the distances among recent human populations, and greater than the distances between the two chimpanzee species. Furthermore, no strong morphological similarities were found between Neanderthals and Late Paleolithic Europeans. This study does not find evidence for Neanderthal contribution to the evolution of modern Europeans. Results are consistent with the recognition of Neanderthals as a distinct species. 2003 Elsevier Science Ltd. All rights reserved.

Early South American groups have been described as possessing a different cranial morphology from the one typically observed among recent Native American populations. This peculiar morphology has been recurrently included in discussions about the initial human dispersion into the Americas, with different authors favoring contrasting scenarios to explain the origin of the morphological and biological diversity inside the continent. In this chapter, we summarize our latest results on the morphological affinities of Early Holocene South American groups and their implications for our understanding of the settlement of the Americas. Our analyses focus on early series from Lagoa Santa, East-Central Brazil, and Sabana de Bogota, Colombia, and are based on multivariate analyses of linear craniometric measurements of these series and reference worldwide populations. Our results indicate strong morphological affinities between early Americans and Late Pleistocene specimens from Europe and East Asia, suggesting that the first American groups shared the same morphological pattern seen in other parts of the planet by the end of the Pleistocene. These analyses suggest that the best scenario to explain the morphological diversity seen in the continent across time is one that assumes an extra influx of morphological diversity into the continent during the Holocene, given that in situ differentiation cannot explain the diversity observed across time. However, it is unclear with the present data the nature of this extra influx, if due to two discrete dispersion waves into the continent or if due to continuous gene-flow from Asia during the Holocene.

The annual turnover rate of trabecular bone by far exceeds that of cortical bone and, therefore, is very sensitive to its daily loading regime. Here we test the hypothesis that the study of the trabecular bone architecture of the human humerus is able to differentiate between different habitual manual activities.For this purpose, we compared the trabecular architecture of the humeral head in a Neolithic population to that of a sample of contemporary Europeans using micro-computed tomography (microCT). We defined in each specimen a spherical volume of interest with a diameter of 57.5 ± 2.5% of the maximal diameter of the humeral head to metrically analyze the bulk of humeral head trabecular architecture. We subsequently quantified the trabecular architectures in the VOIs, measuring seven standard 3D-morphometric parameters, and used univariate and multivariate statistical analyses for comparisons within and between populations.Univariate statistical analysis showed significant differences in a combination of 3D-morphometric parameters. A principal components analysis of the 3D-morphometrics of the trabecular architectures separated the Neolithic from the contemporary samples on the basis of differences in their gross trabecular architecture, including differences in the bone volume fraction (BV/TV), the number of trabeculae per unit length (Tb N), and the distance between trabeculae (Tb Sp).We interpret the significant differences found in the humeral trabecular bone of the Neolithic and the contemporary group as likely reflecting the distinct manual working routines. The trabecular bone configuration in the Neolithic sample shows presumably functional signatures of prehistoric subsistence techniques and activity levels.

Many studies have described shape variation of the modern human cranium in relation to subsistence; however, patterns of covariation within the masticatory apparatus (MA) remain largely unexplored. The patterns and intensity of shape covariation, and how this is related to diet, are essential for understanding the evolution of functional masticatory adaptations of the human cranium. Within a worldwide sample (n = 255) of 15 populations with different modes of subsistence, we use partial least squares analysis to study the relationships between three components of the MA: upper dental arch, masseter muscle, and temporalis muscle attachments. We show that the shape of the masseter muscle and the shape of the temporalis muscle clearly covary with one another, but that the shape of the dental arch seems to be rather independent of the masticatory muscles. On the contrary, when relative positioning, orientation, and size of the masticatory components is included in the analysis, the dental arch shows the highest covariation with the other cranial parts, indicating that these additional factors are more important than just shape with regard to covariation within the MA. Covariation patterns among these cranial regions differ mainly between hunting-fishing and gathering-agriculture groups, possibly relating to greater masticatory strains resulting from a large meat component in the diet. High-strain groups show stronger covariation between upper dental arch and masticatory muscle shape when compared with low-strain groups. These results help to provide a clearer understanding of constraints and interlinkage of shape variation within the human MA and allow for more realistic modeling and predictions in future biomechanical studies.

a Paleoanthropology, Senckenberg Center for Human Evolution and Paleoenvironment, Eberhard Karls Universit€ at Tubingen, Rumelinstrasse 23, Tubingen 72070, Germany b Department of Anthropology, Stony Brook University, Stony Brook, 11794-4364 New York, USA c Department of Anatomical Sciences, Stony Brook University, Stony Brook, 11794-4364 New York, USA d Department of Cultural Heritage, University of Bologna, Via degli Ariani 1, 48121 Ravenna, Italy e Department of Human Evolution, Max Planck Institute for Evolutionary Anthropology, Deutscher Platz 6, D-04103 Leipzig, Germany f Department of Archaeology, University of Cape Town, Rondebosch 7701, South Africa g Institute for Archaeology, History, Culture and Religious Studies, University of Bergen, Oysteinsgate 3, N-5007 Bergen, Norway h Evolutionary Studies Institute, University of the Witwatersrand, 1 Jan Smuts Avenue, Braamfontein 2000, Johannesburg, South Africa

Geometric morphometric methods have been increasingly used in paleoanthropology in the last two decades, lending greater power to the analysis and interpretation of the human fossil record. More recently the advent of the wide use of computed tomography and surface scanning, implemented in combination with geometric morphometrics (GM), characterizes a new approach, termed Virtual Anthropology (VA). These methodological advances have led to a number of developments in human evolutionary studies. We present some recent examples of GM and VA related research in human evolution with an emphasis on work conducted at the University of Tubingen and other German research institutions.

Despite broad consensus on Africa as the main place of origin for anatomically modern humans, their dispersal pattern out of the continent continues to be intensely debated. In extant human populations, the observation of decreasing genetic and phenotypic diversity at increasing distances from sub-Saharan Africa has been interpreted as evidence for a single dispersal, accompanied by a series of founder effects. In such a scenario, modern human genetic and phenotypic variation was primarily generated through successive population bottlenecks and drift during a rapid worldwide expansion out of Africa in the Late Pleistocene. However, recent genetic studies, as well as accumulating archaeological and paleoanthropological evidence, challenge this parsimonious model. They suggest instead a “southern route” dispersal into Asia as early as the late Middle Pleistocene, followed by a separate dispersal into northern Eurasia. Here we test these competing out-of-Africa scenarios by modeling hypothetical geographical migration routes and assessing their correlation with neutral population differentiation, as measured by genetic polymorphisms and cranial shape variables of modern human populations from Africa and Asia. We show that both lines of evidence support a multiple-dispersals model in which Australo-Melanesian populations are relatively isolated descendants of an early dispersal, whereas other Asian populations are descended from, or highly admixed with, members of a subsequent migration event.

Dental development is one aspect of growth that is linked to diet and to life history but has not been investigated among colobines since the work of Schultz [1935]. This study establishes the dental eruption sequence for several colobine species and compares it to that of other catarrhines. The mandibles and maxillae of two hundred and four juvenile colobine specimens were scored for presence or absence of permanent teeth and for stages of partial eruption. Eruption was defined as ranging between tooth emergence (any part of a tooth crown above the alveolar margin) and full occlusion, with three intermediate levels manifest between these boundaries. In African colobines, represented by C. guereza, C. angolensis and P. badius, M2 erupts before I2, and in C. angolensis it also erupts before I1. The canine is delayed, erupting after the premolars in females and after M3 in males. Asian colobines show greater diversity in eruption sequences. Nasalis shows no early eruption of the molars and is very similar to Macaca. In Trachypithecus and Pygathrix M(2) erupts before I(2). The canine in Trachypithecus is delayed, erupting after the premolars and, in some males, after M3. In Presbytis M2 erupts before both incisors; M3 erupts before C in both sexes, and often before both premolars. Although the actual timing of eruption is unknown, all colobine species examined except N. larvatus showed some degree of relatively early eruption of M2 and M3. The lack of this tendency in Nasalis sets this genus apart from all other colobines represented in this study. Dental eruption sequence is thought to reflect life history patterns. Early molar eruption in colobines was thought by Schultz (1935) to be a primitive character reflecting shorter life history. Faster growth rates found in folivorous primates have been interpreted as being related to an adaptation to folivory (Leigh 1994), and early eruption of molars may be part of this dietary specialization. The relationships between dental development and both diet and life history are investigated.

Cet article relate la conference qui s'est tenue a Athenes et Areopolis en 1998 et dont le sujet etait la paleoanthropologie de la peninsule de Mani, en Grece. Des ossements humains ont ete mis au jour dans l'une des quatres grottes du site d'Apidima. La decouverte d'un crâne retient, ici, l'attention des AA.

Attempts to place Palaeolithic finds within a precise climatic framework are complicated by both uncertainty over the radiocarbon calibration beyond about 21,500 14C years bp and the absence of a master calendar chronology for climate events from reference archives such as Greenland ice cores or speleothems. Here we present an alternative approach, in which 14C dates of interest are mapped directly onto the palaeoclimate record of the Cariaco Basin by means of its 14C series, circumventing calendar age model and correlation uncertainties, and placing dated events in the millennial-scale climate context of the last glacial period. This is applied to different sets of dates from levels with Mousterian artefacts, presumably produced by late Neanderthals, from Gorham's Cave in Gibraltar: first, generally accepted estimates of about 32,000 14C years bp for the uppermost Mousterian levels; second, a possible extended Middle Palaeolithic occupation until about 28,000 14C years bp; and third, more contentious evidence for persistence until about 24,000 14C years bp. This study shows that the three sets translate to different scenarios on the role of climate in Neanderthal extinction. The first two correspond to intervals of general climatic instability between stadials and interstadials that characterized most of the Middle Pleniglacial and are not coeval with Heinrich Events. In contrast, if accepted, the youngest date indicates that late Neanderthals may have persisted up to the onset of a major environmental shift, which included an expansion in global ice volume and an increased latitudinal temperature gradient. More generally, our radiocarbon climatostratigraphic approach can be applied to any 'snapshot' date from discontinuous records in a variety of deposits and can become a powerful tool in evaluating the climatic signature of critical intervals in Late Pleistocene human evolution.

Uncovering links between skeletal morphology and locomotor behavior is an essential component of paleobiology because it allows researchers to infer the locomotor repertoire of extinct species based on preserved fossils. In this study, we explored ulnar shape in anthropoid primates using 3D geometric morphometrics to discover novel aspects of shape variation that correspond to observed differences in the relative amount of forelimb suspensory locomotion performed by species. The ultimate goal of this research was to construct an accurate predictive model that can be applied to infer the significance of these behaviors. We studied ulnar shape variation in extant species using principal component analysis. Species mainly clustered into phylogenetic groups along the first two principal components. Upon closer examination, the results showed that the position of species within each major clade corresponded closely with the proportion of forelimb suspensory locomotion that they have been observed to perform in nature. We used principal component regression to construct a predictive model for the proportion of these behaviors that would be expected to occur in the locomotor repertoire of anthropoid primates. We then applied this regression analysis to Pliopithecus vindobonensis, a stem catarrhine from the Miocene of central Europe, and found strong evidence that this species was adapted to perform a proportion of forelimb suspensory locomotion similar to that observed in the extant woolly monkey, Lagothrix lagothricha.

On a basis of a method for muscle cross-sectional area estimation from cortical bone area that was previously developed (Slizewski et al. Anat Rec 2013; 296:1695-1707), we reconstructed muscle cross-sectional area at 65% of radius length for a sample of Neolithic human remains from the Linear Pottery Culture (ca. 5,700-4,900 years BC). Muscle cross-sectional area estimations for the Neolithic sample were compared to in vivo measurements from a recent human sample. Results demonstrate that the Neolithic individuals had larger muscle cross-sectional area relative to radius length than the contemporary humans and that their forearms were more muscular and robust. We also found significant differences in relative muscle cross-sectional area between Neolithic and recent children that indicate different levels of physical stress and isometric activities. Our results fit into the framework of studies previously published about the sample and the Linear Pottery Culture. Therefore, the new approach was successfully applied to an archaeological sample for the first time here. Results of our pilot study indicate that muscle cross-sectional area estimation could in the future supplement other anthropological methods currently in use for the analysis of postcranial remains.

Neanderthals are the group of fossil humans that inhabited Western Eurasia from the mid-Middle Pleistocene until ca. 40 Ka ago, when they disappeared from the fossil record, only a few millennia after the first modern humans appear in Europe. They are characterized by a suite of morphological features that in combination produce a uniquemorphotype. They are commonly associated with theMousterian lithic industry, although toward the end of their tenure they are sometimes found with assemblages resembling those produced by early modern humans. Although there is still discussion over their taxonomic status and relationship with modern humans, it is now commonly recognized that they represent a distinct, Eurasian evolutionary lineage sharing a common ancestor with modern humans in the Middle Pleistocene. This lineage is thought to have been isolated from the rest of the Old World, probably due to the climatic conditions of the glacial cycles. Glacial climate conditions are often thought to have been at least in part responsible for the evolution of some of the distinctive Neanderthal morphology, although genetic drift was probably also very important. The causes of the Neanderthal extinction are not well understood. Worsening climate and competition with modern humans are implicated. The Discovery of Neanderthals: Historical Background Although the first Neanderthal remains were discovered in the early nineteenth century (Engis child in 1830, Forbes Quarry adult in 1848), it was not until the discovery of the skeleton from the Neander Valley in 1856, roughly coinciding with the publication of Darwin’s “The Origin of Species” in 1859, that the existence of an extinct kind of archaic humans was recognized. It is this locality that lends its name to the group, and it is there that the debate surrounding the relationship of Neanderthals with modern humans began. The antiquity of the Neanderthal skeleton and its status as an archaic human was not immediately accepted. Instead, its peculiar anatomical attributes were considered the result of various pathologies, including rickets. Its antiquity was only firmly established with the eventual discovery of additional skeletons of similar morphology associated with lithic artifacts and extinct fauna. Neanderthals were assigned to the species H. neanderthalensis as early as 1864 (King 1864). However, once their status as archaic predecessors of modern humans was accepted, their relationship with modern humans, and particularly modern Europeans, began to be intensely debated. The predominant view in the 1910s and 1920s was represented by scientists like Marcellin Boule and Sir Arthur Keith, who were among the most influential scholars of their day. They placed Neanderthals in their own species and rejected any ancestral role for them in the evolution of modern people, pointing out their “primitiveness” and presumed inferiority (Boule 1911/1913; Boule and Vallois 1957). 2244 K. Harvati

During a recent examination of our morphological 3-D coordinate dataset, it was ascertained that it included a subadult specimen not listed in the publication. This specimen was not shown in the plots but mistakenly remained in the analysis as part of the early modern human sample (EAM). We thank Dr. Waddell for bringing this issue to our attention. We have re-analyzed our data excluding this specimen. Although some of the numerical results change slightly, the main findings, relationships among samples and our conclusions do not change. The PCA and CVA plots, as well as the minimum spanning tree, are nearly identical to those originally reported (Corrected Figure 3). The new Mahalanobis D2 results are similar in the values obtained and pattern of relationships among samples to the ones originally reported (Corrected SI Table 2). The main difference is the distance between Iwo Eleru and the early modern human Qafzeh-Skhul sample from the Levant (EAM), which dates to between 90-130 ka, in the distances corrected for unequal sample sizes. This group, and not the much later Zhoukoudian Upper Cave sample as originally reported, is the closest neighbor to Iwo Eleru in the corrected Mahalanobis D2. This result, although different from that originally reported, strengthens our conclusion that Iwo Eleru bears greater morphological similarity to early rather than late modern human samples from Africa and Eurasia. The revised dataset and the updated key for the labels used in the dataset are also provided with this Correction. Figure 3: SI Table 2: Click here for additional data file.(29K, xls) Dataset: http://www.plosone.org/corrections/pone.0024024.001.cn.txt Label Key: http://www.plosone.org/corrections/pone.0024024.002.cn.doc

Diet-related cranial variation in modern humans is well documented on a regional scale, with ample examples of cranial changes related to the agricultural transition. However, the influence of subsistence strategy on global cranial variation is less clear, having been confirmed only for the mandible, and dietary effects beyond agriculture are often neglected. Here we identify global patterns of subsistence-related human cranial shape variation. We analysed a worldwide sample of 15 populations (n = 255) with known subsistence strategies using 3-D landmark datasets designed to capture the shape of different units of the cranium. Results show significant correlations between global cranial shape and diet, especially for temporalis muscle shape and general cranial shape. Importantly, the differences between populations with either a plant- or an animal-based diet are more pronounced than those between agriculturalists and hunter-gatherers, suggesting that the influence of diet as driver of cranial variation is not limited to Holocene transitions to agricultural subsistence. Dental arch shape did not correlate with subsistence pattern, possibly indicating the high plasticity of this region of the face in relation to age, disease and individual use of the dentition. Our results highlight the importance of subsistence strategy as one of the factors underlying the evolution of human geographic cranial variation.

The taxonomic attribution of the 1.55 million year old young adult fossil calvaria KNM-ER 42700 from Ileret, Kenya, is subject to ongoing controversy. It has been attributed to H. erectus based on comparative description and linear measurements. However, 3-D geometric morphometric analysis found that this specimen fell outside the range of variation of H. erectus in its cranial shape, which was intermediate between H. erectus and modern humans. One problem is that analyses so far were conducted on the original specimen, which shows slight post-mortem distortion. Here we use a surface scan of a high resolution cast of KNM-ER 42700 to virtually reconstruct the calvaria and conduct a new 3D geometric morphometric analysis of both its original and its reconstructed shape. Our comparative sample included several specimens of H. erectus (s.l., including the subadult KNM-WT 15000), H. habilis, H. heidelbergenis (s.l.) and H. neanderthalensis, as well as early and Upper Paleolithic H. sapiens. Our principal component analysis results showed that, like the original specimen, our virtual reconstruction of KNM-ER 42700 is also intermediate in shape between fossil Homo and modern humans. Taphonomic distortion, therefore, appears to not have been a major factor affecting previous 3-D geometric morphometric analyses. The intermediate shape of KNM-ER 42700 might instead be related to the young developmental age of the specimen. Further work on reconstructing the original specimen or based on computed tomorgraphic scans is needed to confirm these results.

This chapter presents the genetic and fossil evidence concerning the origins and earliest dispersals of Homo sapiens across the Old World. The authors begin by reviewing the evidence for an African origin of Homo sapiens, and then consider the following questions: how many major dispersals of modern humans were there out of Africa, when did they occur, and by what routes? Keywords: Pleistocene; human evolution; Genetics; Asia; Africa

Currently in physical anthropology there is a need for reliable methods of sex estimation for immature individuals and highly fragmented remains. This study develops a sex estimation technique from discriminant function analysis of the bony labyrinth as it matures before puberty and can survive taphonomic conditions that would destroy most other skeletal material. The bony labyrinth contains the organs of hearing and balance. For this reason biologists and paleoanthropologists have undertaken research in this area to understand evolutionary changes in locomotion. Prior studies have found clear differences between species, but within-species variation has not been satisfactorily investigated. 3D segmentations of the left and right labyrinths of 94 individuals from a Cretan collection were generated and measured. Mean measurements of height, width, size, and shape indices were analyzed for sexual dimorphism, bilateral asymmetry, and measurement error. Significant sexual dimorphism was detected for several measurements. For sex estimation, the single best variable was the radius of curvature of the posterior semicircular canal, which achieved 76% accuracy. Two multivariate functions increased accuracy to 84%. Although these equations are less accurate than equations for complete long bones and crania, they appear to be as accurate as or better than other techniques for sexing immature individuals and temporal bones.

The Kalamakia cave, a Middle Paleolithic site on the western coast of the Mani peninsula, Greece, was excavated in 1993–2006 by an interdisciplinary team from the Ephoreia of Paleoanthropology and Speleology (Greek Ministry of Culture) and the Museum national d'Histoire naturelle (Paris). The site is dated to between ca. 100,000 and >39,000 years BP (Before Present) and has yielded Mousterian lithics, a rich fauna, and human remains from several layers. The latter include 10 isolated teeth, a cranial fragment and three postcranial elements. The remains represent at least eight individuals, two of them subadults, and show both carnivore and anthropogenic modifications. They can be identified as Neanderthal on the basis of diagnostic morphology on most specimens. A diet similar to that of Neanderthals from mixed habitat is suggested by our analysis of dental wear (occlusal fingerprint analysis) and microwear (occlusal texture microwear analysis), in agreement with the faunal and palynological analyses of the site. These new fossils significantly expand the Neanderthal sample known from Greece. Together with the human specimens from Lakonis and Apidima, the Kalamakia human remains add to the growing evidence of a strong Neanderthal presence in the Mani region during the Late Pleistocene.

The Manyara Beds in the area of Makuyuni Village in the Lake Manyara Basin, Tanzania have been studied for nearly a century, but interpretations of their age have ranged from Middle Pleistocene to Late Pliocene. New geological, paleontological and archeological fieldwork was conducted at the site and has provided siginificant new evidence, including refined stratigraphy, radiometric age estimates, preliminary paleomagnetic analysis, significant new faunal collections, as well as stratigraphic context for some of the lithic artifacts in stratigraphic context. These efforts have succesfully constrained the geological age estimates for the Manyara Beds to between less than 0.63 to 1.3 Ma, and the age of the two hominin bearing localities to between 0.63 and 0.78 Ma. This new chronology may impact the taxonomic interpretation of the hominin remains recovered from the site. They also suggest that two mammalian taxa, Metridiochoerus compactus and Eurygnathohippus, may have some of their youngest known occurences in the Manyara Beds. Acheulean lithics were also found in stratigraphic context during this more narrow time interval. Furthermore, the presence of potential cut-marks on the surface of a bovid mandible may represent the first evidence for human modification of bones from the Manyara Beds.

Facial morphology comprises some of the most distinctive features of early modern humans. The rich fossil record of Morocco allows assessing changes in facial morphology from the late Middle Pleistocene through the Late Pleistocene. Specimens associated with the Aterian industry in Morocco were originally thought to be relatively recent (40–20 ka), but could be much older (35–90 ka). Predating this population are the late Middle Pleistocene specimens of Irhoud. Later in the same geographical area, larger samples are represented by the Iberomaurusian series. We conducted a 3D geometric morphometric analysis of the facial shape of the Aterian specimen Dar es-Soltan II-5, with the aim of deciphering the affinities of this specimen with earlier North African and Levantine fossils, later Upper Paleolithic Eurasian specimens, as well as later North African populations. We used a large comparative sample (n = 191) comprising seven geographic populations of recent humans, Iberomaurusians from Afalou and Taforalt (n = 22), and Middle and Late Pleistocene Eurasian and African fossils. The 3D coordinates of 19 facial landmarks were collected. Specimen landmark configurations were processed with Generalized Procrustes Analysis. Principal Components, Canonical Variates, and cluster analyses were performed and Procrustes distances and Mahalanobis squared distances were calculated. Both Irhoud 1 and Dar es-Soltan II-5 are similar to the early anatomically modern humans from Qafzeh, and the Iberomaurusian sample is closely connected to the Upper Paleolithic European sample.

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 hypotheses about brain evolution in Homo. Three-dimensional geometric morphometric analyses of endobasicranial shape reveal previously undocumented details of evolutionary changes in Homo sapiens. Larger olfactory bulbs, relatively wider orbitofrontal cortex, relatively increased and forward projecting temporal lobe poles appear unique to modern humans. Such brain reorganization, beside physical consequences for overall skull shape, might have contributed to the evolution of H. sapiens' learning and social capacities, in which higher olfactory functions and its cognitive, neurological behavioral implications could have been hitherto underestimated factors. © 2011 Macmillan Publishers Limited. All rights reserved.