Reconstructing the Deep Population History of Central and South America.

We report genome-wide ancient DNA from 49 individuals forming four parallel time transects in Belize, Brazil, the Central Andes, and the Southern Cone, each dating to at least ∼9,000 years ago. The common ancestral population radiated rapidly from just one of the two early branches that contributed to Native Americans today. We document two previously unappreciated streams of gene flow between North and South America. One affected the Central Andes by ∼4,200 years ago, while the other explains an affinity between the oldest North American genome associated with the Clovis culture and the oldest Central and South Americans from Chile, Brazil, and Belize. However, this was not the primary source for later South Americans, as the other ancient individuals derive from lineages without specific affinity to the Clovis-associated genome, suggesting a population replacement that began at least 9,000 years ago and was followed by substantial population continuity in multiple regions.

Apidima Cave fossils provide earliest evidence of Homo sapiens in Eurasia.

Two fossilized human crania (Apidima 1 and Apidima 2) from Apidima Cave, southern Greece, were discovered in the late 1970s but have remained enigmatic owing to their incomplete nature, taphonomic distortion and lack of archaeological context and chronology. Here we virtually reconstruct both crania, provide detailed comparative descriptions and analyses, and date them using U-series radiometric methods. Apidima 2 dates to more than 170 thousand years ago and has a Neanderthal-like morphological pattern. By contrast, Apidima 1 dates to more than 210 thousand years ago and presents a mixture of modern human and primitive features. These results suggest that two late Middle Pleistocene human groups were present at this site-an early Homo sapiens population, followed by a Neanderthal population. Our findings support multiple dispersals of early modern humans out of Africa, and highlight the complex demographic processes that characterized Pleistocene human evolution and modern human presence in southeast Europe.

Virtual reconstruction and geometric morphometric analysis of the Kocabas hominin fossil from Turkey: Implications for taxonomy and evolutionary significance.

The Kocabas specimen comes from a travertine quarry near the homonymous village in the Denizli basin (Turkey). The specimen comprises three main fragments: portions of the right and left parietal and left and right parts of the frontal bone. The fossil was assumed to belong to the Homo erectus s.l. hypodigm by some authors, whereas others see similarities with Middle Pleistocene fossils (Broken Hill 1/Kabwe, Bodo, or Ceprano). Here, we present the first attempt to make a complete reconstruction of the missing medial portion of the frontal bone and a comprehensive geometric morphometric analysis of this bone. We restored the calotte by aligning and mirroring the three preserved fragments. Afterward, we restored the missing portion by applying the thin-plate spline interpolation algorithm of target fossils onto the reconstructed Kocabas specimen. For the geometric morphometric analyses, we collected 80 landmarks on the frontal bone (11 osteometric points, 14 bilateral curve semilandmarks, and 41 surface semilandmarks). The comparative sample includes 21 fossils from different chronological periods and geographical areas and 30 adult modern humans from different populations. Shape analyses highlighted the presence in Kocabas of features usually related to Middle Pleistocene Homo, such as a developed supraorbital torus associated with a relatively short frontal squama and reduced post-toral sulcus. Cluster analysis and linear discriminant analysis classification procedure suggest Kocabas being part of the same taxonomic unit of Eurasian and African Middle Pleistocene Homo. In light of our results, we consider that attributing the Kocabas hominin to H. erectus s.l. may be unwarranted. Results of our analyses are compatible with different evolutionary scenarios, but a more precise chronological framework is needed for a thorough discussion of the evolutionary significance of this specimen. Future work should clarify its geological age, given uncertainties regarding its stratigraphic provenance.

Similar cranial trauma prevalence among Neanderthals and Upper Palaeolithic modern humans.

Neanderthals are commonly depicted as leading dangerous lives and permanently struggling for survival. This view largely relies on the high incidences of trauma that have been reported1,2 and have variously been attributed to violent social behaviour3,4, highly mobile hunter-gatherer lifestyles2 or attacks by carnivores5. The described Neanderthal pattern of predominantly cranial injuries is further thought to reflect violent encounters with large prey mammals, resulting from the use of close-range hunting weapons1. These interpretations directly shape our understanding of Neanderthal lifestyles, health and hunting abilities, yet mainly rest on descriptive, case-based evidence. Quantitative, population-level studies of traumatic injuries are rare. Here we reassess the hypothesis of higher cranial trauma prevalence among Neanderthals using a population-level approach-accounting for preservation bias and other contextual data-and an exhaustive fossil database. We show that Neanderthals and early Upper Palaeolithic anatomically modern humans exhibit similar overall incidences of cranial trauma, which are higher for males in both taxa, consistent with patterns shown by later populations of modern humans. Beyond these similarities, we observed species-specific, age-related variation in trauma prevalence, suggesting that there were differences in the timing of injuries during life or that there was a differential mortality risk of trauma survivors in the two groups. Finally, our results highlight the importance of preservation bias in studies of trauma prevalence.

Author Correction: New fossils from Jebel Irhoud, Morocco and the pan-African origin of Homo sapiens.

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: x̅ = 33.15, σ = 3.26 for height (rather than x̅ = 34.23, σ = 4.57); and x̅ = 16.25, σ = 1.28 for breadth (rather than x̅ = 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.

The evolution and changing ecology of the African hominid oral microbiome.

The oral microbiome plays key roles in human biology, health, and disease, but little is known about the global diversity, variation, or evolution of this microbial community. To better understand the evolution and changing ecology of the human oral microbiome, we analyzed 124 dental biofilm metagenomes from humans, including Neanderthals and Late Pleistocene to present-day modern humans, chimpanzees, and gorillas, as well as New World howler monkeys for comparison. We find that a core microbiome of primarily biofilm structural taxa has been maintained throughout African hominid evolution, and these microbial groups are also shared with howler monkeys, suggesting that they have been important oral members since before the catarrhine-platyrrhine split ca. 40 Mya. However, community structure and individual microbial phylogenies do not closely reflect host relationships, and the dental biofilms of Homo and chimpanzees are distinguished by major taxonomic and functional differences. Reconstructing oral metagenomes from up to 100 thousand years ago, we show that the microbial profiles of both Neanderthals and modern humans are highly similar, sharing functional adaptations in nutrient metabolism. These include an apparent Homo-specific acquisition of salivary amylase-binding capability by oral streptococci, suggesting microbial coadaptation with host diet. We additionally find evidence of shared genetic diversity in the oral bacteria of Neanderthal and Upper Paleolithic modern humans that is not observed in later modern human populations. Differences in the oral microbiomes of African hominids provide insights into human evolution, the ancestral state of the human microbiome, and a temporal framework for understanding microbial health and disease.

Icex: Advances in the automatic extraction and volume calculation of cranial cavities.

The use of non-destructive approaches for digital acquisition (e.g. computerised tomography-CT) allows detailed qualitative and quantitative study of internal structures of skeletal material. Here, we present a new R-based software tool, Icex, applicable to the study of the sizes and shapes of skeletal cavities and fossae in 3D digital images. Traditional methods of volume extraction involve the manual labelling (i.e. segmentation) of the areas of interest on each section of the image stack. This is time-consuming, error-prone and challenging to apply to complex cavities. Icex facilitates rapid quantification of such structures. We describe and detail its application to the isolation and calculation of volumes of various cranial cavities. The R tool is used here to automatically extract the orbital volumes, the paranasal sinuses, the nasal cavity and the upper oral volumes, based on the coordinates of 18 cranial anatomical points used to define their limits, from 3D cranial surface meshes obtained by segmenting CT scans. Icex includes an algorithm (Icv) for the calculation of volumes by defining a 3D convex hull of the extracted cavity. We demonstrate the use of Icex on an ontogenetic sample (0-19 years) of modern humans and on the fossil hominin crania Kabwe (Broken Hill) 1, Gibraltar (Forbes' Quarry) and Guattari 1. We also test the tool on three species of non-human primates. In the modern human subsample, Icex allowed us to perform a preliminary analysis on the absolute and relative expansion of cranial sinuses and pneumatisations during growth. The performance of Icex, applied to diverse crania, shows the potential for an extensive evaluation of the developmental and/or evolutionary significance of hollow cranial structures. Furthermore, being open source, Icex is a fully customisable tool, easily applicable to other taxa and skeletal regions.

New fossils from Jebel Irhoud, Morocco and the pan-African origin of Homo sapiens.

Fossil evidence points to an African origin of Homo sapiens from a group called either H. heidelbergensis or H. rhodesiensis. However, the exact place and time of emergence of H. sapiens remain obscure because the fossil record is scarce and the chronological age of many key specimens remains uncertain. In particular, it is unclear whether the present day 'modern' morphology rapidly emerged approximately 200 thousand years ago (ka) among earlier representatives of H. sapiens or evolved gradually over the last 400 thousand years. Here we report newly discovered human fossils from Jebel Irhoud, Morocco, and interpret the affinities of the hominins from this site with other archaic and recent human groups. We identified a mosaic of features including facial, mandibular and dental morphology that aligns the Jebel Irhoud material with early or recent anatomically modern humans and more primitive neurocranial and endocranial morphology. In combination with an age of 315 ± 34 thousand years (as determined by thermoluminescence dating), this evidence makes Jebel Irhoud the oldest and richest African Middle Stone Age hominin site that documents early stages of the H. sapiens clade in which key features of modern morphology were established. Furthermore, it shows that the evolutionary processes behind the emergence of H. sapiens involved the whole African continent.

Neanderthal behaviour, diet, and disease inferred from ancient DNA in dental calculus.

Recent genomic data have revealed multiple interactions between Neanderthals and modern humans, but there is currently little genetic evidence regarding Neanderthal behaviour, diet, or disease. Here we describe the shotgun-sequencing of ancient DNA from five specimens of Neanderthal calcified dental plaque (calculus) and the characterization of regional differences in Neanderthal ecology. At Spy cave, Belgium, Neanderthal diet was heavily meat based and included woolly rhinoceros and wild sheep (mouflon), characteristic of a steppe environment. In contrast, no meat was detected in the diet of Neanderthals from El Sidrón cave, Spain, and dietary components of mushrooms, pine nuts, and moss reflected forest gathering. Differences in diet were also linked to an overall shift in the oral bacterial community (microbiota) and suggested that meat consumption contributed to substantial variation within Neanderthal microbiota. Evidence for self-medication was detected in an El Sidrón Neanderthal with a dental abscess and a chronic gastrointestinal pathogen (Enterocytozoon bieneusi). Metagenomic data from this individual also contained a nearly complete genome of the archaeal commensal Methanobrevibacter oralis (10.2× depth of coverage)-the oldest draft microbial genome generated to date, at around 48,000 years old. DNA preserved within dental calculus represents a notable source of information about the behaviour and health of ancient hominin specimens, as well as a unique system that is useful for the study of long-term microbial evolution.

Evolution of Homo in the Middle and Late Pleistocene.

The Middle and Late Pleistocene is arguably the most interesting period in human evolution. This broad period witnessed the evolution of our own lineage, as well as that of our sister taxon, the Neanderthals, and related Denisovans. It is exceptionally rich in both fossil and archaeological remains, and uniquely benefits from insights gained through molecular approaches, such as paleogenetics and paleoproteomics, that are currently not widely applicable in earlier contexts. This wealth of information paints a highly complex picture, often described as 'the Muddle in the Middle,' defying the common adage that 'more evidence is needed' to resolve it. Here we review competing phylogenetic scenarios and the historical and theoretical developments that shaped our approaches to the fossil record, as well as some of the many remaining open questions associated with this period. We propose that advancing our understanding of this critical time requires more than the addition of data and will necessitate a major shift in our conceptual and theoretical framework.

First record of Macaca (Cercopithecidae, Primates) in the Middle Pleistocene of Greece.

In this article, we describe an almost complete macaque mandible from the Middle Pleistocene locality Marathousa 1 in the Megalopolis Basin of southern Greece. The mandible belonged to a male individual of advanced ontogenetic age and of estimated body mass ∼13 kg. Comparative metric analysis of its teeth permits its attribution to the Barbary macaque Macaca sylvanus, a species that was geographically widely distributed in Western Eurasia during the Plio-Pleistocene. The dental dimensions of the Marathousa 1 macaque fit better within the variation of the Early Pleistocene M. s. florentina and the Middle to Late Pleistocene M. s. pliocena rather than with the extant representative M. s. sylvanus. Moreover, principal component analysis reveals a better match with M. s. pliocena. However, because no clear-cut diagnostic criteria have been defined to differentiate these European fossil subspecies, we attribute the Marathousa 1 specimen to M. s. cf. pliocena, in agreement with the chronology of the locality. Previously known only from the Early Pleistocene of Greece by some isolated teeth, this is the first record of Macaca in the Middle Pleistocene of the country and one of very few in the eastern sector of the peri-Mediterranean region. We discuss the presence of macaques in the paleolake environment of Marathousa 1, as well as their predation risks from both carnivores and hominins present at the locality.

New horizons in reconstructing past human behavior: Introducing the "Tübingen University Validated Entheses-based Reconstruction of Activity" method.

An accurate reconstruction of habitual activities in past populations and extinct hominin species is a paramount goal of paleoanthropological research, as it can elucidate the evolution of human behavior and the relationship between culture and biology. Variation in muscle attachment (entheseal) morphology has been considered an indicator of habitual activity, and many attempts have been made to use it for this purpose. However, its interpretation remains equivocal due to methodological shortcomings and a paucity of supportive experimental data. Through a series of studies, we have introduced a novel and precise methodology that focuses on reconstructing muscle synergies based on three-dimensional and multivariate analyses among entheses. This approach was validated using uniquely documented anthropological samples, experimental animal studies, histological observations, and geometric morphometrics. Here, we detail, synthesize, and critically discuss the findings of these studies, which overall point to the great potential of entheses in elucidating aspects of past human behavior.

Prevalence of cranial trauma in Eurasian Upper Paleolithic humans.

OBJECTIVES: This study characterizes patterns of cranial trauma prevalence in a large sample of Upper Paleolithic (UP) fossil specimens (40,000-10,000 BP). MATERIALS AND METHODS: Our sample comprised 234 individual crania (specimens), representing 1,285 cranial bones (skeletal elements), from 101 Eurasian UP sites. We used generalized linear mixed models (GLMMs) to assess trauma prevalence in relation to age-at-death, sex, anatomical distribution, and between pre- and post-Last Glacial Maximum (LGM) samples, while accounting for skeletal preservation. RESULTS: Models predicted a mean cranial trauma prevalence of 0.07 (95% CI 0.003-0.19) at the level of skeletal elements, and of 0.26 (95% CI 0.08-0.48) at the level of specimens, each when 76-100% complete. Trauma prevalence increased with skeletal preservation. Across specimen and skeletal element datasets, trauma prevalence tended to be higher for males, and was consistently higher in the old age group. We found no time-specific trauma prevalence patterns for the two sexes or age cohorts when comparing samples from before and after the LGM. Samples showed higher trauma prevalence in the vault than in the face, with vault remains being affected predominantly in males. DISCUSSION: Cranial trauma prevalence in UP humans falls within the variation described for Mesolithic and Neolithic samples. According to our current dataset, UP males and females were exposed to slightly different injury risks and trauma distributions, potentially due to different activities or behaviors, yet both sexes exhibit more trauma among the old. Environmental stressors associated with climatic changes of the LGM are not reflected in cranial trauma prevalence. To analyze trauma in incomplete skeletal remains we propose GLMMs as an informative alternative to crude frequency calculations.

Biocultural evidence of precise manual activities in an Early Holocene individual of the high-altitude Peruvian Andes.

OBJECTIVES: Cuncaicha, a rockshelter site in the southern Peruvian Andes, has yielded archaeological evidence for human occupation at high elevation (4,480 masl) during the Terminal Pleistocene (12,500-11,200 cal BP), Early Holocene (9,500-9,000 cal BP), and later periods. One of the excavated human burials (Feature 15-06), corresponding to a middle-aged female dated to ~8,500 cal BP, exhibits skeletal osteoarthritic lesions previously proposed to reflect habitual loading and specialized crafting labor. Three small tools found in association with this burial are hypothesized to be associated with precise manual dexterity. MATERIALS AND METHODS: Here, we tested this functional hypothesis through the application of a novel multivariate methodology for the three-dimensional analysis of muscle attachment surfaces (entheses). This original approach has been recently validated on both lifelong-documented anthropological samples as well as experimental studies in nonhuman laboratory samples. Additionally, we analyzed the three-dimensional entheseal shape and resulting moment arms for muscle opponens pollicis. RESULTS: Results show that Cuncaicha individual 15-06 shows a distinctive entheseal pattern associated with habitual precision grasping via thumb-index finger coordination, which is shared exclusively with documented long-term precision workers from recent historical collections. The separate geometric morphometric analysis revealed that the individual's opponens pollicis enthesis presents a highly projecting morphology, which was found to strongly correlate with long joint moment arms (a fundamental component of force-producing capacity), closely resembling the form of Paleolithic hunter-gatherers from diverse geo-chronological contexts of Eurasia and North Africa. DISCUSSION: Overall, our findings provide the first biocultural evidence to confirm that the lifestyle of some of the earliest Andean inhabitants relied on habitual and forceful precision grasping tasks.

2000-year-old pathogen genomes reconstructed from metagenomic analysis of Egyptian mummified individuals.

BACKGROUND: Recent advances in sequencing have facilitated large-scale analyses of the metagenomic composition of different samples, including the environmental microbiome of air, water, and soil, as well as the microbiome of living humans and other animals. Analyses of the microbiome of ancient human samples may provide insights into human health and disease, as well as pathogen evolution, but the field is still in its very early stages and considered highly challenging. RESULTS: The metagenomic and pathogen content of Egyptian mummified individuals from different time periods was investigated via genetic analysis of the microbial composition of various tissues. The analysis of the dental calculus' microbiome identified Red Complex bacteria, which are correlated with periodontal diseases. From bone and soft tissue, genomes of two ancient pathogens, a 2200-year-old Mycobacterium leprae strain and a 2000-year-old human hepatitis B virus, were successfully reconstructed. CONCLUSIONS: The results show the reliability of metagenomic studies on Egyptian mummified individuals and the potential to use them as a source for the extraction of ancient pathogen DNA.

A virtual assessment of the suprainiac depressions on the Eyasi I (Tanzania) and Aduma ADU-VP-1/3 (Ethiopia) Pleistocene hominin crania.

Despite a steady increase in our understanding of the phenotypic variation of Pleistocene Homo, debate continues over phylogenetically informative features. One such trait is the suprainiac fossa, a depression on the occipital bone above inion that is commonly considered an autapomorphy of the Neanderthal lineage. Challenging this convention, depressions in the suprainiac region have also been described for two Pleistocene hominin crania from sub-Saharan Africa: Eyasi I (Tanzania) and ADU-VP-1/3 (Ethiopia). Here, we use a combined quantitative and qualitative approach, using µCT imaging, to investigate the occipital depressions on these specimens. The results show that neither the external nor the internal morphologies of these depressions bear any resemblance to the Neanderthal condition. A principal component analysis based on multiple thickness measurements along the occipital squama demonstrates that the relative thickness values for the internal structures in Eyasi I and ADU-VP-1/3 are within the range of Homo sapiens. Thus, our results support the autapomorphic status of the Neanderthal suprainiac fossa and highlight the need to use nuanced approaches and multiple lines of evidence.

Experimental evidence that physical activity affects the multivariate associations among muscle attachments (entheses).

The morphology of entheses (muscle/tendon attachment sites) on bones is routinely used in paleontological and bioarcheological studies to infer the physical activity patterns of ancient vertebrate species including hominins. However, such inferences have often been disputed owing to limitations of the quantitative methods commonly employed and a lack of experimental evidence demonstrating direct effects of physical activity on entheseal morphology. Recently, we introduced a new and improved method of quantifying and analyzing entheseal morphology that involves repeatable three-dimensional measurements combined with multivariate statistics focused on associations among multiple entheses. Here, to assess the validity of our method for investigating variation in entheseal morphology related to physical activity patterns, we analyzed femora of growing turkeys that were experimentally exercised for 10 weeks on either an inclined or declined treadmill or served as controls (N=15 individuals, 5 per group). Our multivariate approach identified certain patterns involving three different entheses (associated with the gluteus primus, medial gastrocnemius, vastus medialis and adductor magnus muscles) that clearly differentiated controls from runners. Importantly, these differences were not observable when comparing groups within each of the three entheseal structures separately. Body mass was not correlated with the resulting multivariate patterns. These results provide the first experimental evidence that variation in physical activity patterns has a direct influence on entheseal morphology. Moreover, our findings highlight the promise of our newly developed quantitative methods for analyzing the morphology of entheses to reconstruct the behavior of extinct vertebrate species based on their skeletal remains.

Basicranial ontogeny comparison in Pan troglodytes and Homo sapiens and its use for developmental stage definition of KNM-ER 42700.

OBJECTIVE: This study aims to develop a comparative basis for assessing the developmental stage of KNM-ER 42700 based on the ontogenetic pattern of the ectocranial surface of the basicranium in modern humans and chimpanzees. MATERIALS AND METHODS: A total of 33 landmarks were collected from an ontogenetic sample of modern humans (80), chimpanzees (51), and 12 individuals classified as Homo erectus s.l. Ontogenetic trajectories were analyzed, and common aspects were extracted for the purpose of discriminating age groups. A regression of size on the extracted shape variables was used to investigate common ontogenetic allometry. RESULTS: The basicranial development of chimpanzees and humans follows different trajectories; however, similarities are also present. The common shape component of development extracted can be used to define age groups in both chimpanzees and modern humans. The extracted shape component presents a similar ontogenetic and static-allometric pattern in these two species. The developmental stages of H. erectus s.l. specimens were attributed following these common traits. Our analysis correctly assigned developmental stages to those specimens of H. erectus for which developmental ages are known. DISCUSSION: The component used for assessing the developmental stage has an ontogenetic allometric component. However, this shape component can discriminate age group irrespective of size and is no longer related to size when static allometry is considered. Adult H. erectus s.l. specimens were attributed to the adult category. KNM-WT 15000 fell with the late juvenile age group, whereas D2700 plotted in the region of overlap between the juvenile and adult age groups and Mojokerto with the younger age groups, as predicted by their known developmental ages. KNM-ER 42700 fell within the adult variability despite its incompletely fused spheno-occipital synchondrosis.

A virtual assessment of the proposed suprainiac fossa on the early modern European calvaria from Cioclovina, Romania.

OBJECTIVES: The calvaria from Cioclovina (Romania) has been argued to possess some traits commonly ascribed to individuals belonging to the Neanderthal lineage, including a suprainiac fossa. However, its supranuchal morphology has only been evaluated with a qualitative analysis of the ectocranial surface. We evaluate whether the morphology of the supranuchal area of this specimen is homologous to the Neanderthal condition. MATERIALS AND METHODS: We described in detail the external morphology, and, using computed tomography, investigated the internal morphology of the Cioclovina supranuchal area. We took measurements of the internal structures and calculated their relative contributions to total cranial vault thickness, which were compared to published data and evaluated with a principal component analysis (PCA). RESULTS: The Cioclovina supranuchal region is characterized by superficial resorption present on the outer layer of the external table. Neither the diploic layer nor the external table decrease in relative thickness in the area above inion. In the PCA, Cioclovina falls within the convex hulls of recent modern Homo sapiens. DISCUSSION: Our results show that the morphology of the Cioclovina supranuchal region does not correspond to the external and internal morphology of the typical Neanderthal suprainiac fossa. It cannot be characterized as a depression but rather as an area presenting superficial bone turnover. Together with earlier results, there is little phenotypic evidence that Cioclovina has high levels of Neanderthal ancestry. Our study demonstrates the usefulness of this quantitative method in assessing proposed Neanderthal-like suprainiac depressions in Upper Paleolithic and other fossil specimens.

Population history of southern Italy during Greek colonization inferred from dental remains.

OBJECTIVES: We are testing competing scenarios regarding the population history of the ancient Greek colonization of southern Italy using dental phenotypic evidence. MATERIALS AND METHODS: We collected dental metric and nonmetric trait data for 481 human skeletons from six archaeological sites along the Gulf of Taranto, dating to pre-colonial (900-700 BC) and post-colonial periods (700-200 BC). We are evaluating scenarios through an individual-level biodistance analysis using a three-pronged approach: (a) by analyzing levels of mobility in pre- and post-colonial periods under a model of isolation-by-distance; (b) by quantifying differences in group means and variances in pre- and post-colonial periods utilizing permutational multivariate analysis of variance and Betadisper analyses; and (c) by identifying ancestries of post-colonial individuals using naïve Bayes classification. RESULTS: Southern Italy during pre-colonial times was characterized by low levels of mobility and marked differences in group means and variances. During post-colonial times, mobility increased and there were no differences in group means and variances. About 18% of the people in post-colonial times were of Greek ancestry and lived equally distributed across Greek colonies and indigenous villages. Nevertheless, the overall biological composition and variability of southern Italy remained relatively unchanged across pre- and post-colonial periods. DISCUSSION: Our results support a scenario in which only few Greek colonists migrated to southern Italy and lived in smaller numbers alongside indigenous people in Greek colonies as well as in indigenous villages. Our results contradict a scenario in which large numbers of Greek invaders founded biologically isolated and substantially homogeneous colonial enclaves within conquered territories.