The influence of climate and population structure on East Asian skeletal morphology.

Recent studies have shown that global variation in body proportions is more complex than previously thought as some traits formerly associated with climate adaptation are better explained by geographic proximity and neutral evolutionary forces. While the recent incorporation of quantitative genetic methodologies has improved understanding of patterns related to climate in Africa, Europe, and the Americas, Asia remains underrepresented in recent and historic studies of body form. As ecogeographic studies tend to focus on male morphology, potential sex differences in features influenced by climate remain largely unexplored. Skeletal measurements encompassing the dimensions of the skull, pelvis, limbs, hands, and feet were collected from male (n = 459) and female (n = 442) remains curated in 13 collections across seven countries in East Asia (n = 901). Osteological data were analyzed with sex and minimum temperature as covariates adjusted by autosomal single-nucleotide polymorphism population genetic distance using univariate Bayesian linear mixed models, and credible intervals were calculated for each trait. Analysis supports a relationship between specific traits and climate as well as providing the magnitude of response in both sexes. After accounting for genetic distance between populations, greater association between climate and morphology was found in postcranial traits, with the relationship between climate and the skull limited primarily to breadth measurements. Larger body size is associated with colder climates with most measurements increasing with decreased temperature. The same traits were not always associated with climate for males and females nor correlated with the same intensity for both sexes. The varied directional association with climate for different regions of the skeleton and between the sexes underscores the necessity of future ecogeographic research to holistically evaluate body form and to look for sex-specific patterns to better understand population responses to environmental stresses.

The Dmanisi Equus: Systematics, biogeography, and paleoecology.

The Equus datum has been established as a geochronologic 'instantaneous' migratory event of a North American Equus species into Eurasia at the beginning of the Pleistocene (2.58 Ma). A remarkable radiation of Equus followed across Eurasia and Africa. Dmanisi includes excellent remains of Equus, well calibrated between 1.85 and 1.76 Ma. Our morphologic and morphometric analyses of the augmented Dmanisi Equus sample support the co-occurrence of Equus stenonis and Equus altidens in the sequence. Dmanisi E. stenonis is found to be morphologically similar to the European E. stenonis populations and represents the best well-dated easternmost occurrence of this species in Eurasia. The Dmanisi E. altidens represents the oldest well-calibrated occurrence of this species in Western Eurasia. Our analyses demonstrate that E. altidens extended its range westward from west Asia to Greece, Germany, Italy, Spain, and possibly France. Our results do not support distinguishing multiple subspecies of E. altidens, including E. altidens altidens, E. altidens granatensis and E. stenonis mygdoniensis. The Dmanisi cranial and postcranial samples exhibit morphologies close both to extant hemiones and zebras. Equus altidens is believed to have been well adapted to newly emergent arid environments in western Eurasia during the late Early and early Middle Pleistocene. The first occurrence of E. altidens at Dmanisi marks an important turnover in the horse communities of the late Early Pleistocene, with a dispersion of this species from West Asia to West Europe ca. 1.8 Ma.

Population level comparisons in central Mexico using cranial nonmetric traits.

OBJECTIVES: We study the genetic diversity between Classic Teotihuacan and its neighboring towns trying to understand how far or close they are at the genetic level. MATERIALS AND METHODS: We use cranial nonmetric traits to study a sample of 280 adult skulls from archaeological sites running from the late Preclassic to the early Postclassic. Samples of Classic Teotihuacan were studied for La Ventilla and San Sebastián Xolalpan neighbors. For the Epiclassic period, samples from Xaltocan, Toluca valley, Mogotes and Xico were used. For the Preclassic and Postclassic samples from Xico were also used. We used a parametric bootstrap for the mean measure of divergence for the statistical analysis. RESULTS: Samples from Xico have small biodistance from Preclassic to Postclassic. Samples from Los Mogotes differ depending on the functional context of deposition, with individuals from household burials (funerary) differing from non-funerary, ceremonial interments and exhibiting affinities to Epiclassic samples from Toluca valley. Epiclassic populations from Xaltocan vary significantly from any samples analyzed. Samples from Classic period Teotihuacan vary considerably among them but form a separate genetic group from all the other populations under study. CONCLUSIONS: The great biodistance separation among Classic Teotihuacan and its neighbor villages of central Mexico let us conclude that, contrary from the classical idea that those villages were confirmed by the inhabitants of Teotihuacan's collapse: They indeed remain as separate populations by themselves.

Importance of the postcranial skeleton in eusuchian phylogeny: Reassessing the systematics of allodaposuchid crocodylians.

Our current knowledge on the crocodyliform evolution is strongly biased towards the skull morphology, and the postcranial skeleton is usually neglected in many taxonomic descriptions. However, it is logical to expect that it can contribute with its own phylogenetic signal. In this paper, the changes in the tree topology caused by the addition of the postcranial information are analysed for the family Allodaposuchidae, the most representative eusuchians in the latest Cretaceous of Europe. At present, different phylogenetic hypotheses have been proposed for this group without reaching a consensus. The results of this paper evidence a shift in the phylogenetic position when the postcranium is included in the dataset, pointing to a relevant phylogenetic signal in the postcranial elements. Finally, the phylogenetic relationships of allodaposuchids within Eusuchia are reassessed; and the internal relationships within Allodaposuchidae are also reconsidered after an exhaustive revision of the morphological data. New and improved diagnoses for each species are here provided.

The exceptional finding of Locus 2 at Dehesilla Cave and the Middle Neolithic ritual funerary practices of the Iberian Peninsula.

There is a significant number of funerary contexts for the Early Neolithic in the Iberian Peninsula, and the body of information is much larger for the Late Neolithic. In contrast, the archaeological information available for the period in between (ca. 4800-4400/4200 cal BC) is scarce. This period, generally called Middle Neolithic, is the least well-known of the peninsular Neolithic sequence, and at present there is no specific synthesis on this topic at the peninsular scale. In 2017, an exceptional funerary context was discovered at Dehesilla Cave (Sierra de Cádiz, Southern Iberian Peninsula), providing radiocarbon dates which place it at the beginning of this little-known Middle Neolithic period, specifically between ca. 4800-4550 cal BC. Locus 2 is a deposition constituted by two adult human skulls and the skeleton of a very young sheep/goat, associated with stone structures and a hearth, and a number of pots, stone and bone tools and charred plant remains. The objectives of this paper are, firstly, to present the new archaeological context documented at Dehesilla Cave, supported by a wide range of data provided by interdisciplinary methods. The dataset is diverse in nature: stratigraphic, osteological, isotopic, zoological, artifactual, botanical and radiocarbon results are presented together. Secondly, to place this finding within the general context of the contemporaneous sites known in the Iberian Peninsula through a systematic review of the available evidence. This enables not only the formulation of explanations of the singular new context, but also to infer the possible ritual funerary behaviours and practices in the 5th millennium cal BC in the Iberian Peninsula.

Coalescing traditions-Coalescing people: Community formation in Pannonia after the decline of the Roman Empire.

The decline of the Roman rule caused significant political instability and led to the emergence of various 'Barbarian' powers. While the names of the involved groups appeared in written sources, it is largely unknown how these changes affected the daily lives of the people during the 5th century AD. Did late Roman traditions persist, did new customs emerge, and did both amalgamate into new cultural expressions? A prime area to investigate these population and settlement historical changes is the Carpathian Basin (Hungary). Particularly, we studied archaeological and anthropological evidence, as well as radiogenic and stable isotope ratios of strontium, carbon, and nitrogen of human remains from 96 graves at the cemetery of Mözs-Icsei dulo. Integrated data analysis suggests that most members of the founder generation at the site exhibited burial practises of late Antique traditions, even though they were heterogeneous regarding their places of origin and dietary habits. Furthermore, the isotope data disclosed a nonlocal group of people with similar dietary habits. According to the archaeological evidence, they joined the community a few decades after the founder generation and followed mainly foreign traditions with artificial skull modification as their most prominent characteristic. Moreover, individuals with modified skulls and late Antique grave attributes attest to deliberate cultural amalgamation, whereas burials of largely different isotope ratios underline the recipient habitus of the community. The integration of archaeological and bioarchaeological information at the individual level discloses the complex coalescence of people and traditions during the 5th century.

Predicting final lesion characteristics during MR-guided focused ultrasound pallidotomy for treatment of Parkinson's disease.

OBJECTIVE: Magnetic resonance-guided focused ultrasound (MRgFUS) ablation of the globus pallidus interna (GPi) is being investigated for the treatment of advanced Parkinson's disease symptoms. However, GPi lesioning presents unique challenges due to the off-midline location of the target. Furthermore, it remains uncertain whether intraprocedural MR thermometry data can predict final lesion characteristics. METHODS: The authors first performed temperature simulations of GPi pallidotomy and compared the results with those of actual cases and the results of ventral intermediate nucleus (VIM) thalamotomy performed for essential tremor treatment. Next, thermometry data from 13 MRgFUS pallidotomy procedures performed at their institution were analyzed using 46°C, 48°C, 50°C, and 52°C temperature thresholds. The resulting thermal models were compared with resulting GPi lesions noted on postprocedure days 1 and 30. Finally, the treatment efficiency (energy per temperature rise) of pallidotomy was evaluated. RESULTS: The authors' modeled acoustic intensity maps correctly demonstrate the elongated, ellipsoid lesions noted during GPi pallidotomy. In treated patients, the 48°C temperature threshold maps most accurately predicted postprocedure day 1 lesion size, while no correlation was found for day 30 lesions. The average energy/temperature rise of pallidotomy was higher (612 J/°C) than what had been noted for VIM thalamotomy and varied with the patients' skull density ratios (SDRs). CONCLUSIONS: The authors' acoustic simulations accurately depicted the characteristics of thermal lesions encountered following MRgFUS pallidotomy. MR thermometry data can predict postprocedure day 1 GPi lesion characteristics using a 48°C threshold model. Finally, the lower treatment efficiency of pallidotomy may make GPi lesioning challenging in patients with a low SDR.

The Bronze and Iron Age populations of the Armenian Highland in the genetic history of Armenians.

OBJECTIVES: To investigate the biological diversity of the late Bronze and Iron Age populations in the Armenian Highland by nonmetric cranial traits, evaluate the genetic continuity in the development of the modern Armenian gene pool, and compare the results obtained with genetic data. MATERIALS AND METHODS: Twenty-eight nonmetric cranial traits were scored on 498 adult crania from different late Bronze and Iron Age cemeteries, as well as from modern Armenians and other European populations. We carried out a biodistance analysis between populations using the mean measure of divergence (MMD) statistics, tested the spatial-temporal model of population structure, and assessed the diversity within the late Bronze and early Iron Ages by using the values of variability index (Fst). RESULTS: The biodistance analysis revealed a close relationship among different ancient Armenian populations and between the average frequencies of the three sequential periods (late Bronze Age, early Iron Age I and II) and modern Armenians. A gradual increase of variability (Fst) within the three successive periods was observed. DISCUSSION: The analysis of nonmetric trait data reflects deep roots and continuity in the formation of the Armenian population. Since at least the Late Bronze Age, owing to permanent isolation, no significant changes have occurred in the Armenian gene pool. An increase in variability over the successive periods reflects the process of population differentiation from a single gene pool while maintaining average trait frequencies. The congruence of the results obtained with the genetic data confirms, once more, the possibility of using nonmetric cranial traits as a proxy for genetic markers.

The human remains from Axlor (Dima, Biscay, northern Iberian Peninsula).

OBJECTIVES: We provide the description and comparative analysis of all the human fossil remains found at Axlor during the excavations carried out by J. M. de Barandiarán from 1967 to 1974: a cranial vault fragment and seven teeth, five of which likely belonged to the same individual, although two are currently lost. Our goal is to describe in detail all these human remains and discuss both their taxonomic attribution and their stratigraphic context. MATERIALS AND METHODS: We describe external and internal anatomy, and use classic and geometric morphometrics. The teeth from Axlor are compared to Neandertals, Upper Paleolithic, and recent modern humans. RESULTS: Two teeth (a left dm2 , a left di1 ) and the parietal fragment show morphological features consistent with a Neandertal classification, and were found in an undisturbed Mousterian context. The remaining three teeth (plus the two lost ones), initially classified as Neandertals, show morphological features and a general size that are more compatible with their classification as modern humans. DISCUSSION: A left parietal fragment (Level VIII) from a single probably adult Neandertal individual was recovered during the old excavations performed by Barandiarán. Additionally, two different Neandertal children lost deciduous teeth during the formations of levels V (left di1 ) and IV (right dm2 ). In addition, a modern human individual is represented by five remains (two currently lost) from a complex stratigraphic setting. Some of the morphological features of these remains suggest that they may represent one of the scarce examples of Upper Paleolithic modern human remains in the northern Iberian Peninsula, which should be confirmed by direct dating.

Human cranium of Candonga Cave site and its implications for the initial peopling of South America.

The aim of this paper is to analyze the morphology of a skull from Candonga Cave (central Argentina), dating to the Pleistocene-Holocene transition, and to discuss its implications for the early peopling of South-America by Palaeoamerican and Amerindian populations. Although this cranium is fragmented, impeding the standard morphometric analysis, an alternative methodology ad hoc was implemented in order to obtain relevant information about the South American people origins. Results of measurements provide new information to reinforce the hypothesis that Paleoamerican crania possess similar morphological traits to those of current Fueguian populations, as previous studies have claimed, and to suggest that the cranium of Candonga has strong morphological affinities with Early Holocene specimens from Lagoa Santa (Brazil) and Late Holocene of Beagle Channel (Argentina), both belonging to Paleoamerican morphology.

Masticatory properties in pre-modern Holocene populations from Northern China.

Recent studies indicate that evolution of the craniofacial skeleton is influenced by dietary behavior, which in turn alters masticatory efficacy and temporomandibular joint (TMJ) stability. In this study the mechanical properties of the masticatory system and the integrity of the TMJ in human populations from Northern China, dated to between 3800 BCE and 100 CE, were assessed. The results demonstrate that the mechanical efficiency is comparable to other modern human populations, though variations are present across different populations. While the ratio of overall weighted muscle efficiency for incisor loading vs. molar loading in pastoral and some recent agricultural groups is similar to early Homo sapiens, the ratio in more ancient agricultural groups is similar to the ratio in populations with heavy anterior paramasticatory activities, such as Neandertals, Inuits, and Native Americans. The TMJ vulnerability negatively correlates with the maxillary dental arch size, and positively with the condylar size. These findings suggest that there are multi-directional strategies in adaptation to heavy anterior teeth loading, such as increasing anterior teeth loading efficiency, increasing facial height, increasing facial breath and facial orthognathy, or decreasing anterior facial length. Furthermore, populations or individuals with a smaller dental arch and high biting efficiency could more easily injure the TMJ during unilateral loadings, which may explain the higher prevalence of TMJ disorders in modern humans, especially in women. These findings further reflect the impact of diachronic changes of the masticatory apparatus and lifestyle and their impact on oral health during recent human history.

State of the art forensic techniques reveal evidence of interpersonal violence ca. 30,000 years ago.

The Cioclovina (Romania) calvaria, dated to ca. 33 cal ka BP and thought to be associated with the Aurignacian lithic industry, is one of the few relatively well preserved representatives of the earliest modern Europeans. Two large fractures on this specimen have been described as taphonomic modifications. Here we used gross and virtual forensic criteria and experimental simulations on synthetic bone models, to investigate their nature. Both forensic trauma pattern analysis and experimental models exclude a postmortem origin for the Cioclovina fractures. Rather, they indicate two incidents of blunt force trauma, the second clearly inflicted with a club-like object. The magnitude and extent of the lesions and the lack of signs of healing indicate a fatal injury. The Upper Paleolithic period is noted for intensified technological innovation, increased symbolic behavior, and cultural complexity. We show that the behavioural repertoire of the earliest modern Europeans also comprised violent inter-personal interactions and murder.

Taking a big bite: Working together to better understand the evolution of feeding in primates.

The study of adaptation requires the integration of an array of different types of data. A single individual can find such integration daunting, if not impossible. In an effort to clarify the role of diet in the evolution of the primate craniofacial and dental apparatus, we assembled a team of researchers that have various types and degrees of expertise. This interaction has provided a range of insights for all contributors, and this has helped to refine questions, clarify the possibilities and limitations that laboratory and field settings offer, and further explore the ways in which laboratory and field data can be suitably integrated. A complete and accurate picture of dietary adaptation cannot be gained in isolation. Collaboration provides the bridge to a more holistic view of primate biology and evolution.

Diachronic changes in craniofacial morphology among the middle-late Holocene populations from Hehuang region, Northwest China.

OBJECTIVES: This study analyzes craniofacial shape variation in the Hehuang region of Northwest China within a population genetic framework, and takes a diachronic approach to explore the relationship betwee cultural discontinuity and biological continuity/discontinuity in the Hehuang region during the middle to late Holocene. MATERIALS AND METHODS: The sample comprises 76 adult skulls from five archaeological sites, ranging from 4,500 to 1,530 BP. 3D geometric morphometrics, multivariate statistics, quantitative evolutionary genetic and biodistance analyses were performed to study the diachronic variation in craniofacial morphology. Analyses were performed on two cranial modules: the face and the cranial vault, across three major diachronic groups representing the late Neolithic (LNA), the Bronze Age (BA), and the Han-Jin dynasty (HD). RESULTS: Average regional FST for both cranial modules was low, indicating relatively greater variation within diachronic groups than among them. While the LNA and BA groups did not show any significant differences in facial and vault shape, significant craniofacial shape changes were detected between the BA and HD groups. DISCUSSION: The consistent craniofacial morphology during the LNA and the BA, and the shift in morphology between the BA and the HD indicates that cultural discontinuity does not always coincide with biological discontinuity. The Hehuang population evolved in situ with few changes, despite cultural and dietary changes, until the HD when migrations from the Central Plains are associated with extra-local gene flow to the area.

Unusual osteological findings in sacred relics from the LESJE monastery in central Serbia.

An adult male skeleton was submitted to the Department of Anatomy at the University of Belgrade for evaluation. It was believed to represent the remains of a second to third century Christian saint from the Lesje Monastery in central Serbia. Examination of the remains revealed an old crush fracture of a thoracic vertebra and an unusual, probably congenital, malformation of the atlanto-occipital joint with deformation of the left occipital condyle and resultant narrowing of the foramen magnum. Although the occipital malformations were most likely congenital, they may still have caused, or contributed to, death by compression of the underlying upper cervical spinal cord.

Morphological variability of Upper Paleolithic and Mesolithic skulls from Sicily.

Scenarios for the dispersal of Homo sapiens in Southern Europe and in the Mediterranean basin have been uncertain, given the scarceness of osteological samples and the simplicity of the proposed archaeologically-based settlement hypotheses. According to available data, the first anatomically modern humans entered Sicily during the Late Pleistocene, coming from the Italian peninsula. A presumably small Late Epigravettian population colonised coastal sites. Later, North-Western archaeological horizons gave hospitality to a significant Mesolithic expansion. In order to verify a hypothesis of continuity in the peopling of the island, we analyzed Sicilian skulls from the Late Epigravettian site of San Teodoro, Eastern Sicily (AMS 14C dated at 14,500 BP) and from the Mesolithic period (14C dated from 9,500 to 8,500 BP) coming from various sites (Uzzo, Molara, Grotta d'Oriente) located on the North Western coast of the island. The aims were to test the biological variability through time within the island as well as to evaluate the relationships of Sicilian Pleistocene hunter-gatherers with Old World populations. We also evaluated the Sicilian Mesolithic uniformity especially between the Uzzo and Grotta d'Oriente sites, given their vicinity and accessibility during the Early Holocene. We applied 3D geometric morphometric methods to assess shape variation as well as geographic and diachronic morphological patterns. All analyzed specimens, plus a comparative sample from the Old World dated from the Upper Paleolithic to recent periods, were transformed in digital images and standard craniofacial landmarks were extracted from the 3D models. Our results underline a high variability among the Mesolithic specimens, as well as a large craniometric distance from the presumed founder Paleolithic settler representatives (San Teodoro specimens) that have closer morphological affinities with other European Upper Paleolithic specimens.

Mitogenomic data indicate admixture components of Central-Inner Asian and Srubnaya origin in the conquering Hungarians.

It has been widely accepted that the Finno-Ugric Hungarian language, originated from proto Uralic people, was brought into the Carpathian Basin by the conquering Hungarians. From the middle of the 19th century this view prevailed against the deep-rooted Hungarian Hun tradition, maintained in folk memory as well as in Hungarian and foreign written medieval sources, which claimed that Hungarians were kinsfolk of the Huns. In order to shed light on the genetic origin of the Conquerors we sequenced 102 mitogenomes from early Conqueror cemeteries and compared them to sequences of all available databases. We applied novel population genetic algorithms, named Shared Haplogroup Distance and MITOMIX, to reveal past admixture of maternal lineages. Our results show that the Conquerors assembled from various nomadic groups of the Eurasian steppe. Population genetic results indicate that they had closest connection to the Onogur-Bulgar ancestors of Volga Tatars. Phylogenetic results reveal that more than one third of the Conqueror maternal lineages were derived from Central-Inner Asia and their most probable ultimate sources were the Asian Scythians and Asian Huns, giving support to the Hungarian Hun tradition. The rest of the lineages most likely originated from the Bronze Age Potapovka-Poltavka-Srubnaya cultures of the Pontic-Caspian steppe. Available data imply that the Conquerors did not have a major contribution to the gene pool of the Carpathian Basin.

Lagoa Santa's contribution to the origins and life of early Americans.

The region of Lagoa Santa, Central-Eastern Brazil, provides an exceptional archeological record about Late Pleistocene/Early Holocene occupation of the Americas. Since the first interventions made by the Danish naturalist Peter Lund in the 19th century, hundreds of human skeletons have been exhumed in the region. These skeletons are complemented by a rich botanic, faunal, technological, and geomorphological archeological record. We explore here the contributions of Lagoa Santa material to the origins and lifestyle of early Americans, providing an historic background. Cranial morphology of Lagoa Santa skeletons allowed the proposition of a model of two biological components for the occupation of the Americas, in which early Americans are morphologically similar to people of African and Australo-Melanesian origin. Furthermore, the archeological record in the region has revealed an intense use of plant resources, a restricted spatial distribution, and the symbolic elaboration of local hunter-gatherers, unveiling a distinct lifestyle compared to early North American populations.

Neurocranial anatomy of an enigmatic Early Devonian fish sheds light on early osteichthyan evolution.

The skull of 'Ligulalepis' from the Early Devonian of Australia (AM-F101607) has significantly expanded our knowledge of early osteichthyan anatomy, but its phylogenetic position has remained uncertain. We herein describe a second skull of 'Ligulalepis' and present micro-CT data on both specimens to reveal novel anatomical features, including cranial endocasts. Several features previously considered to link 'Ligulalepis' with actinopterygians are now considered generalized osteichthyan characters or of uncertain polarity. The presence of a lateral cranial canal is shown to be variable in its development between specimens. Other notable new features include the presence of a pineal foramen, the some detail of skull roof sutures, the shape of the nasal capsules, a placoderm-like hypophysial vein, and a chondrichthyan-like labyrinth system. New phylogenetic analyses place 'Ligulalepis' as a stem osteichthyan, specifically as the sister taxon to 'psarolepids' plus crown osteichthyans. The precise position of 'psarolepids' differs between parsimony and Bayesian analyses.